smithsonian contributions to zoology • number 650

Smithsonian Institution

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Smithsonian Institution

Scholarly Press

American Recent 

Eulipotyphla

Nesophontids, Solenodons, Moles, 

and Shrews in the New World

Neal Woodman
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smithsonian contributions to zoology • numb e r 6 5 0

American Recent 

Eulipotyphla

Nesophontids, Solenodons, Moles, 

and Shrews in the New World

Neal Woodman

WASHINGTON, D.C.

2018
ABSTRACT

Woodman, Neal. American Recent Eulipotyphla: Nesophontids, Solenodons, Moles, and Shrews in the New World. 

Smithsonian Contributions to Zoology, number 650, vi + 107 pages, 27 figures, 1 table, 4 appendixes, 2 indexes, 

2018. — The mammalian taxonomic order Eulipotyphla comprises the living families Erinaceidae (gymnures, hedgehogs, 

and moonrats), Solenodontidae (solenodons), Soricidae (shrews), and Talpidae (desmans and moles), as well as the recently 

extinct Nesophontidae (nesophontids). Morphological and molecular studies continue to alter our view of relationships 

within and among these families, and this research has added considerably to our understanding of the diversity, distributions, and relationships of many of the New World (i.e., North and South America and associated islands) species that 

comprise them. Currently, there are more than 450 recognized living species worldwide, making the Eulipotyphla the third 

most speciose order of mammals. In this work, I attempt to summarize the taxonomic results of recent studies, provide 

a guide to the most appropriate current applications of New World taxonomic names and their synonyms, and indicate 

current understanding of their distributions. The eulipotyphlans of this region currently include 111 recognized species of 

shrews, seven species of moles, and both living species of solenodons.

Cover images, from left to right: Solenodon cubanus (Figure 7, top), Neurotrichus gibbsii (Figure 12), Sorex brevicaudatus

(detail from Figure 17).

Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS

P.O. Box 37012, MRC 957, Washington, D.C. 20013-7012

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Compilation copyright © 2018 Smithsonian Institution

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Library of Congress Cataloging-in-Publication Data

Names: Woodman, Neal, author. | Smithsonian Institution Scholarly Press, publisher.

Title: American recent Eulipotyphla : Nesophontids, Solenodons, Moles, and Shrews in the New World / 

Neal Woodman.

Other titles: Smithsonian contributions to zoology ; no. 650. 0081-0282

Description: Washington, D.C. : Smithsonian Institution Scholarly Press, 2018. | Series: Smithsonian contributions 

to zoology, ISSN 0081-0282 ; number 650 | Includes bibliographical references and index. | Smithsonian Institution 

Compilation copyright 2018

Identifiers: LCCN 2018038627

Subjects: LCSH: Solenodontidae—North America. | Solenodontidae—South America. | Moles (Animals)—North 

America. | Moles (Animals)—South America. | Shrews—North America. | Shrews—South America.

Classification: LCC QL737.S7 W66 2018 | DDC 599.33097—dc23 | SUDOC SI 1.27:650

LC record available at https://lccn.loc.gov/2018038627

ISSN: 1943-6696 (online); 0081-0282 (print)

Publication date (online): 16 November 2018

Ó The paper used in this publication meets the minimum requirements of the American National Standard for Permanence 

of Paper for Printed Library Materials Z39.48–1992.
Contents

LIST OF FIGURES v

INTRODUCTION 1

Phylogenetic Relationships of the Eulipotyphla 2

What Happened to the “Insectivora”? 3

Organization of the List 4

EARLY NAMES FOR NORTH AMERICAN EULIPOTYPHLANS 5

RECENT AMERICAN EULIPOTYPHLA 10

Order Eulipotyphla 10

Family Nesophontidae Anthony, 1916 10

Genus Nesophontes Anthony, 1916 10

Family Solenodontidae Gill, 1872 12

Genus Solenodon Brandt, 1833 12

Family Talpidae G. Fischer, 1814 14

Tribe Condylurini Gill, 1875 14

Genus Condylura Illiger, 1811 14

Tribe Neurotrichini Hutterer, 2005 17

Genus Neurotrichus Günther, 1880 17

Tribe Scalopini Gill, 1875 19

Genus Parascalops True, 1894 19

Genus Scalopus É. Geoffroy Saint-Hilaire, 1803 20

Genus Scapanus Pomel, 1848 25

Family Soricidae G. Fischer, 1814 28

Subfamily Soricinae G. Fischer, 1814 28

Tribe Blarinini Kretzoi, 1965 28

Genus Blarina Gray, 1838 28

Genus Cryptotis Pomel, 1848 34

Tribe Notiosoricini Reumer, 1984 45

Genus Megasorex Hibbard, 1950 45

Genus Notiosorex Coues, 1877 45

Tribe Soricini G. Fischer, 1814 46

Genus Sorex Linnaeus, 1758 46

Acknowledgments 76
i v • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

APPENDIX A: PHYLOGENETIC LISTING OF SPECIES 77

APPENDIX B: LITERATURE ABBREVIATIONS 81

APPENDIX C: INSTITUTIONAL ABBREVIATIONS 87

APPENDIX D: DISTRIBUTION OF TYPE LOCALITIES 89

REFERENCES 91

TAXONOMIC INDEX 101

AUTHORITY INDEX 105
1. Numbers of recognized species of New World Soricidae through time 2

2. Phylogenetic relationships of the families of Eulipotyphla 3

3. Sorex avellanarum É. Geoffroy Saint-Hilaire, 1803 7

4. John Bachman’s (1837) shrews: Sorex richardsonii Bachman, 1837 

(=Sorex arcticus Kerr, 1792); Sorex forsteri Richardson, 1828 (=Sorex 

cinereus Kerr, 1792); Sorex cooperi Bachman, 1837 (=Sorex cinereus

Kerr, 1792); Sorex fimbripes Bachman, 1837 8

5. Pen-over-pencil sketch of the “black-eared shrew” or “corn mouse,” 

Sorex melanotis Rafinesque, 1818b 9

6. The “red mole,” Seba’s (1734) Talpa, rubra, Americana 10

7. The Cuban solenodon, Solenodon cubanus Peters, 1861 13

8. The Hispaniolan solenodon, Solenodon paradoxus Brandt, 1833 14

9. “La Taupe du Canada,” Talpa canadensis De la Faille, 1769 

(=Condylura cristata (Linnaeus 1758)) 15

10. Condylura macroura Harlan, 1825 (=Condylura cristata

(Linnaeus, 1758)) 16

11. Spencer Baird’s shrews and mole: Blarina exilipes Baird, 1857 

(=Sorex parvus Say, 1822 = Cryptotis parvus); Blarina berlandieri 

Baird, 1857 (=Cryptotis berlandieri); Sorex hoyi Baird, 1857; Sorex 

platyrhinus De Kay, 1842 (=Sorex cinereus Kerr, 1792); Urotrichus 

gibbsii Baird, 1857 (=Neurotrichus gibbsii) 18

12. Neurotrichus gibbsii (Baird, 1857) 19

13. Thomas Pennant’s (1771) “radiated mole” (=Sorex cristatus Linnaeus, 

1758 = Condylura cristata) and “long-tailed mole” (=Scalops breweri

Bachman, 1841= Parascalops breweri) 21

14. Ventral view of the “purple mole,” Seba’s (1734) Talpa, Virginianus,

niger, supinus (=Sorex aquaticus Linnaeus, 1758 = Scalopus aquaticus) 23

15. Sorex aquaticus Linnaeus, 1758 (=Scalopus aquaticus) 23

16. Sorex brevicaudus Say, 1822 (=Blarina brevicauda) and Sorex parvus

Say, 1822 (=Cryptotis parvus) 29

17. Sorex brevicaudatus Duvernoy, 1842 (=Sorex brevicaudus Say, 

1842 = Blarina brevicauda) 30

18. Benjamin Barton’s (1806) “black shrew” from the vicinity of 

Philadelphia, the basis for Sorex niger Ord, 1815 (=Sorex talpoides

Gapper, 1830 = Blarina brevicauda talpoides) 31

Figures
v i • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

19. Sorex talpoides Gapper, 1830 (=Blarina brevicauda talpoides) 31

20. John Bachman’s (1837) shrews: Sorex carolinensis Bachman, 1837 

(=Blarina carolinensis); Sorex longirostris Bachman, 1837; Sorex 

cinereus Bachman, 1837 (=Blarina floridana Merriam, 

1895 = Cryptotis parvus floridanus); Sorex dekayi Bachman, 

1837 (=Sorex talpoides Gapper, 1830 = Blarina brevicauda talpoides) 33

21. Sorex harlani Duvernoy, 1842 (=Cryptotis parvus (Say, 1822)) 42

22. Sorex veraepacis Alston, 1877 and Blarina micrura Alston, 1877 

(=Blarina tropicalis Merriam, 1895 = Cryptotis tropicalis) 45

23. Spencer Baird’s shrews: Sorex thompsoni Baird, 1857 (=Sorex hoyi 

thompsoni ); Sorex haydeni Baird, 1857; Sorex suckleyi Baird, 1857 

(=Sorex vagrans Baird, 1857); Sorex pachyurus Baird, 1857 

(=Sorex arcticus laricorum Jackson, 1925) 50

24. Sorex forsteri Richardson, 1828 (=Sorex cinereus Kerr, 1792) 52

25. Spencer Baird’s shrews: Sorex cooperi Bachman, 1837 (=Sorex cinereus

Kerr, 1792); Sorex vagrans Baird, 1857; Sorex trowbridgii Baird, 1857; 

Neosorex navigator Baird, 1857 (=Sorex navigator) 53

26. Otisorex platyrhinus De Kay, 1842 (=Sorex cinereus Kerr, 1792) 54

27. Amphisorex lesueurii Duvernoy, 1842 (=Sorex cinereus Kerr, 1792) 56
INTRODUCTION

This work summarizes the taxonomic status of Recent Eulipotyphla inhabiting 

North and South America, including the Caribbean region, up to approximately 1 January 2018. The name Eulipotyphla Waddell et al., 1999 refers to the mammalian orderlevel clade that includes the living families Erinaceidae G. Fischer, 1814 (gymnures, 

hedgehogs, and moonrats), Solenodontidae Gill, 1872 (solenodons), Soricidae G. Fischer, 

1814 (shrews), and Talpidae G. Fischer, 1814 (desmans and moles). By extension, the 

order also includes the recently extinct Nesophontidae, which forms a natural grouping 

with the Solenodontidae that Brace et al. (2016) termed the Solenodonata. My original 

intent was to provide an updated accounting of the taxonomy of North American and 

Caribbean Eulipotyphla using Miller and Kellogg’s (1955) section on the order Insectivora as a model and guide. However, only a single eulipotyphlan genus, the soricid 

Cryptotis Pomel, 1848, occurs on continental regions south of Honduras, making it logical and useful to expand geographical coverage to provide a comprehensive overview of 

all New World (i.e., North and South America and associated islands) Eulipotyphla. The 

extinct family Nesophontidae is included because this taxon was covered by Miller and 

Kellogg (1955) and because many nesophontids are now known to have survived into the 

post-Columbian era (Miller, 1930; MacPhee et al., 1999; Brace et al., 2016).

With more than 500 living species worldwide, Eulipotyphla includes approximately 

8% of all living mammals, making it the third most speciose order of Recent Mammalia 

after Rodentia (rodents) and Chiroptera (bats). Within the order, Soricidae is the largest 

family, with nearly 85% of living eulipotyphlan species, followed by Talpidae (10%), 

Erinaceidae (<5%), and Solenodontidae (<0.5%; Wilson and Reeder, 2005; Burgin et al., 

2018). The Pan-American contribution to Eulipotyphla currently includes 111 species of 

shrews, eight species of moles, both living species of solenodons, and no erinaceids. This 

enumeration of species reflects considerable change in our understanding of the diversity 

of New World Eulipotyphla since Miller and Kellogg (1955; plus Tate, 1932 for South 

American shrews), who recognized 77 soricids, seven talpids, and three solenodontids (in 

two genera). That our varying view of the diversity of this group has not been linear can 

be seen in Figure 1. This plot tallies recognized species of soricids through time since the 

publication of the 10th edition of Systema Naturae (Linnaeus, 1758), which serves as a 

American Recent Eulipotyphla: 

Nesophontids, Solenodons, Moles, 

and Shrews in the New World

Neal Woodman

U.S. Geological Survey, Biological Survey Unit, 

Patuxent Wildlife Research Center, MRC 108, 

National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, 

D.C. 20013-7012, USA.

Correspondence: woodmann@si.edu

Manuscript received 16 November 2017; accepted 9 April 2018
2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

convenient, if artificial, starting point for modern biological taxonomy. Linnaeus (1758) recognized only two New World species—identified by him as belonging to the genus Sorex but both 

later recognized as talpids—of a worldwide total of five known 

eulipotyphlan species. From there, the number of New World 

species generally trends upward as new species were discovered 

and described. The strong downward trend from the 1950s until 

about 1980, followed by a strong upsurge represents, in part, 

changing philosophies regarding concepts of species and the 

practical application of those concepts to living organisms. 

Phylogenetic Relationships of the Eulipotyphla

Molecular analyses indicate that Eulipotyphla represents 

the earliest branch within the Laurasiatheria clade of mammals 

(Figure 2), making it the sister to all other laurasiatherian orders (Chiroptera, Cetartiodactyla, Perissodactyla, Carnivora, 

Pholidota; Tarver et al., 2016). Molecular clock model estimates 

generally place the divergence of the Eulipotyphla from the rest 

of the Laurasiatheria in the Late Cretaceous (Douady and Douzery, 2003: 94–75 million years ago [mya]; Springer et al., 2003: 

95–76 mya; Bininda-Emonds et al., 2007: 94.4–89.2 mya; Tarver 

et al., 2016: 73.75–81.96 mya). In contrast, phenotypic analysis 

indicates a later origin that contributed to an early Paleocene 

radiation of mammals (O’Leary et al., 2013: 58.9–57.8 mya).

Various calibrated molecular models of the timings of lineage divergences within Eulipotyphla were summarized recently 

by Springer et al. (2018: table 1). The majority of these models 

indicate that the most basal branch of Recent Eulipotyphla, the 

Solenodonata clade (Solenodontidae + Nesophontidae), most 

likely diverged from other Eulipotyphla in the Late Cretaceous 

about 86–66 mya and certainly before the end of the Paleocene 

(Sato et al., 2016: 60.8–57.3 mya; Brandt et al., 2017: 98.6–62.1 

mya). Among the remaining eulipotyphlans, the Talpidae was the 

next Recent family to branch off, at about 85–61 mya or possibly later (dos Reis et al., 2012: 62.0–60.9 mya; Sato et al., 2016: 

60.8–57.3 mya). This left the Soricidae and Erinaceidae, which 

diverged an estimated 71–61 mya (Sato et al., 2016: 60.4–57.3 

FIGURE 1. Graph of the numbers of recognized species of New World Soricidae through 

time. Sources: Linnaeus, 1758; Pennant, 1771; Erxleben, 1777; Zimmermann, 1777; Gmelin, 

1788; Kerr, 1792; Shaw, 1800; Turton, 1802; Illiger, 1815; Ord, 1815; Harlan, 1825; Godman, 1826; Richardson, 1837; Bachman, 1837; Baird, 1857; True, 1884; Merriam, 1895a, 

1895b; Miller and Rehn, 1901 (+Thomas, 1898); Miller, 1912a (+Thomas, 1898); Miller, 1924 

(+Thomas, 1921); Cabrera, 1925; Miller and Kellogg, 1955 (+Tate, 1932); Hall and Kelson, 

1959 (+Cabrera, 1958); Hall, 1981 (+Cabrera, 1958); Honacki et al., 1982; Hutterer, 1993, 

2005; this work.
NUMBER 650 • 3

mya). The separation of Nesophontidae from Solenodontidae 

may have been the last major branching event among the Recent 

families, occurring 68.6–43.6 mya (Brace et al., 2016). The topology and timing are obviously simplified because they ignore 

relevant diverse fossil clades, such as Proscalopidae Reed, 1961 

and Heterosoricidae Viret and Zapfe, 1951.

Relationships within Nesophontidae and within Solenodontidae remain unresolved, whereas those within Soricidae 

and Talpidae continue to be modified as additional taxa and 

characters are added to phylogenetic analyses. Integration of genetic and morphological analyses continues to be a goal (He et 

al., 2015; Schwermann and Thompson, 2015). Among talpids, 

tribal groupings used herein follow Hutterer (2005), but I ignore the subfamily group because inferred tree topologies vary 

with the analytical methods used, resulting in continued uncertainty about relationships among tribes and the composition of 

subfamilies (Shinohara et al., 2003; Crumpton and Thompson, 

2013; Piras et al., 2015; Schwermann and Thompson, 2015; He 

et al., 2016). Divergence time estimates from He et al. (2016) 

indicate that the Talpidae lineage split from other eulipotyphlans 

in the Late Cretaceous, which is in agreement with Brace et al.’s 

(2016: 75.6–65.2 mya) and Springer et al.’s (2018: 74.0–63.9 

mya) time frames. He et al. (2016) dated the most recent common ancestor of the modern members of the family to near the 

Eocene boundary (57–42 mya), when the ancestor of Uropsilus diverged from the rest of the family. Other tribal divergences 

began in the late Eocene (39.5–35.3 mya), when the Scalopini 

split from the rest of the moles, and continued through the Oligocene (28.1–23.0 mya), when the Neurotrichini and Urotrichini 

diverged. For a different view of the timing of divergences, see 

Piras et al. (2015).

Of the three living subfamilies of the Soricidae (Crocidurinae, Myosoricinae, Soricinae), only members of the Soricinae 

occur in the Western Hemisphere. My definition of tribes within 

the subfamily follows Hutterer (2005). Divergence time estimates indicate separation of the tribal lineages took place before 

the end of the middle Miocene, commencing 20.5–12.5 mya and 

finishing by 17.4–10.2 mya (Dubey et al., 2007). 

What Happened to the “Insectivora”?

I have been queried by a number of colleagues regarding the 

use of the name Eulipotyphla (Waddell et al., 1999) instead of the 

other names that have been applied to the ordinal grouping that 

includes the Soricidae. These include Lipotyphla Haeckel, 1866, 

Soricomorpha Gregory, 1910, and the traditional Insectivora 

Bowdich, 1821. The International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 

[ICZN], 1999) does not specifically address standards for the 

use of names higher than the family level, and in the absence of 

such formal guidance, I use the name whose original composition most closely matches the monophyletic extant group that 

is currently deemed to comprise the lineages within the order. 

In this case, it is the ordinal grouping that includes Erinaceidae, 

Solenodontidae, Soricidae, and Talpidae and that excludes other 

families that would render the order polyphyletic.

Insectivora Bowdich, 1821 was originally proposed as a 

family in the order Sarcophaga (Table 1). This name may have 

been derived from the Insectivores of Blainville (1816a, 1816b) 

and Cuvier (1816), a subdivision of their order Carnassiers. The 

full content of Blainville’s Insectivores is uncertain, but it did not 

include talpids, which he placed in the separate suborder Anomaux, rather than with the Insectivores in suborder Normaux. 

Bowdich’s (1821) Insectivora more closely matched Cuvier’s 

(1816) Insectivores in content. On the basis of the list of genera 

Bowdich (1821) assigned to Insectivora, the original grouping 

included Erinaceidae, Soricidae, and Talpidae but also Chrysochloridae (golden moles) and Tenrecidae (tenrecs), which now 

comprise the separate group Afrosoricida (Gatesy et al., 2017), 

thereby making the Insectivora polyphyletic. The first solenodon,

Solenodon paradoxus Brandt, 1833, was not yet known to natural 

historians, so the absence of the Solenodontidae at that time is irrelevant. Through time, additional families—the Macroscelididae 

FIGURE 2. Phylogenetic relationships of the families of Eulipotyphla (based on Douady and Douzery, 2003; Springer et al., 2003; 

Brace et al., 2016).
4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(elephant shrews) and Tupaiidae (treeshrews)—were added to 

the Insectivora (Gregory, 1910). The order gained the reputation among many taxonomists as a “wastebasket” group composed of families of uncertain relationships whose members were 

often considered “primitive” in general nature relative to other 

Eutherians.

Haeckel (1866) erected Lipotyphla and Menotyphla as the 

two suborders of the order Insectivora (Table 1). Lipotyphla originally included genera that today are distributed among six different families: Chrysochloridae, Erinaceidae, Solenodontidae, 

Soricidae, Talpidae, and Tenrecidae. The suborder Menotyphla 

contained the Macroscelididae and Tupaiidae. The inclusion of 

the Afrosoricidae (Chrysochloridae and Tenrecidae) in the Lipotyphla makes that original group polyphyletic.

Soricomorpha Gregory, 1910 originated as one of three 

“sections” of the suborder Lipotyphla within the order Insectivora (Table 1). Soricomorpha included only the two families 

Soricidae and Talpidae. Section Erinaceomorpha was erected 

for the single family Erinaceidae, whereas the Solenodontidae, 

Chrysochloridae, and Tenrecidae together formed the section 

Zalambdodonta (Gregory, 1910). This classification separated 

Erinaceidae and Soricidae, which molecular studies indicate are 

sister taxa, thus making Soricomorpha paraphyletic (Douady et 

al., 2002; Douady and Douzery, 2003; Gatesy et al., 2017). 

Waddell et al. (1999) coined Eulipotyphla as an ordinallevel grouping specifically for the clade containing “hedgehogs, 

shrews, solenodon, and moles” (Waddell et al., 1999:1). Although the use of undefined English common names lacks the 

specificity preferable in the naming of a new taxonomic grouping in the late twentieth century, the content can be interpreted 

to include the Erinaceidae, Solenodontidae, Soricidae, and Talpidae (Table 1). This is the only one of the four ordinal names 

whose original content is neither polyphyletic nor paraphyletic 

under our current understanding of mammalian relationships 

(Asher, 2007; Bininda-Emonds et al., 2007; O’Leary et al., 

2013; Brace et al., 2016; Sato et al., 2016; Gatesy et al., 2017). 

Given this criterion, Eulipotyphla is the only acceptable name 

for the order.

Organization of the List

In the following list, the sequence of the families within the 

order Eulipotyphla is phylogenetic (Figure 2). To ease the location of a particular taxon, however, the sequences of tribes and 

genera within a family, species within a genus, and subspecies 

within a species are alphabetical. A phylogenetic listing of eulipotyphlan taxa is presented in Appendix A.

Synonymies for genera and species-level groups (i.e., species 

and subspecies) follow the format and recommendations outlined 

in Gardner and Hayssen (2004). For species with recognized subspecies, the synonymy is placed with the nominate subspecies. 

Synonymies cite the original description and the first use of each 

name or name combination (synonyms) that are associated with a 

TABLE 1. Overview of changes in taxonomy of eulipotyphlans (Erinaceidae, Solenodontidae, Soricidae, Talpidae) through time. Names 

in bold type are groups names that have been used for the taxonomic 

order that includes the Soricidae. See Gregory (1910) and McKenna 

and Bell (1997) for a more complete review of Mammalia.

Taxonomy Current family

Bowdich, 1821:24, 31–32

Order Sarcophaga

 Family Insectivora

 Erinaceus Erinaceidae

 Sorex Soricidae

 Mygale, Scalops, Talpa Talpidae

 Chrysochloris Chrysochloridae

 Centenes [sic] Tenrecidae

Haeckel, 1866: clx

Order Insectivora

Suborder Menotyphla

 Family Macroscelidia Macroscelididae

 Family Cladobatida Tupaiidae

Suborder Lipotyphla 

 Family Erinaceidea Erinaceidae

 Family Soricida Soricidae

 Family Talpida Talpidae and Chrysochloridae

 Family Centetida Tenrecidae and Solenodontidae 

Gregory, 1910

Order Insectivora

Suborder Lipotyphla

 Section Zalambdodonta

 Family Centetidae Tenrecidae

 Family Potamogalidae Tenrecidae

 Family Solenodontidae Solenodontidae

 Family Chrysochloridae Chrysochloridae

 Section Erinaceomorpha

 Family Erinaceidae Erinaceidae

 Section Soricomorpha

 Family Soricidae Soricidae

 Family Talpidae Talpidae

Waddell et al., 1999

Order Eulipotyphlaa

 “Hedgehogs” Erinaceidae

 “Shrews” Soricidae

 “Solenodon” Solenodontidae

 “Moles” Talpidae

a Waddell et al. (1999) used nonspecific common names to designate the contents 

of their order Eulipotyphla, rather than using defined family names. The quotation marks indicate that these are the actual terms they used and they communicate the rather loose definition those names provide.
NUMBER 650 • 5

currently valid taxon and note the current status of the name. The 

name of an original describer or author of a unique taxon name 

(including an incorrect subsequent spelling) is directly associated 

with the taxon name as originally proposed with no intervening 

punctuation. The names of subsequent authors (e.g., authors of 

unique name combinations) are separated from the taxon name 

by a colon. A list of abbreviations used for literature citations in 

the synonymies can be found in Appendix B. References forming 

part of a synonymy are not included in the References unless they 

are also cited in the text. Where type specimens (i.e., holotypes, 

lectotypes, neotypes) are known, I provide the institutional abbreviation (explained in Appendix C) and catalog number for 

the type specimens. Type localities in quotation marks are given 

as originally published, with additional or clarifying information provided afterward. Additions may be included in brackets within quotes, but I generally attempted to avoid this except 

when filling out abbreviations in the original. The distribution 

of type localities is provided in Appendix D. Additional useful 

information on type specimens of eulipotyphlans can be found in 

Sanborn (1947), Jones and Genoways (1969), Youngman (1973), 

Koopman (1976), Baranova et al. (1981), Hafner (1993), Lawrence (1993), Helgen and McFadden (2001), Turni et al. (2007), 

Fisher and Ludwig (2015), and similar published catalogs. 

Date of publication can be critical to determining the priority of a particular taxonomic name, but printed dates on journals 

and books often do not accurately reflect their public availability 

(e.g., Duncan, 1937; Woodman, 2010, 2013, 2017). I attempted 

to provide the most accurate date that I could determine for publications in which new taxa are described. Meisel (1924–1929) 

is particularly useful for determining correct dates of publication 

for some older North American publications. 

EARLY NAMES FOR NORTH AMERICAN 

EULIPOTYPHLANS

The first comprehensive systematic accounting of the terrestrial vertebrates of the New World by an American natural historian was published anonymously by the Philadelphia naturalist 

George Ord in the second volume of the 1815 American edition 

of “Guthrie’s Geography” (Guthrie, 1815; Woodman, 2013). At 

that time, Ord (1815) recognized two species of moles and six 

species of shrews (Soricidae) inhabiting the Western Hemisphere 

(Figure 1). Prior to that time, lists of New World vertebrates 

were generally limited to regional faunas (e.g., Hernandez, 1651; 

Pennant, 1784) or were enveloped within more comprehensive 

systematic treatises (e.g., Seba, 1734; Schreber, 1778). One exception is Forster’s (1771) A Catalogue of the Animals of North 

America, for which the mammals were derived from Pennant’s 

(1771) Synopsis of Quadrupeds, but Forster’s list is an incomplete representation even of the latter work.

Ord’s (1815) list appeared in an era when the community 

of naturalists in North America was just beginning to reach the 

critical mass necessary to organize scientific societies, found 

journals, and transition individual cabinets of curiosities into 

scientifically useful collections. Early American naturalists were 

anxious to earn the respect of their European counterparts, but 

they had the added onus of establishing that the local fauna and 

flora were worthy of serious study (Semonin, 2000). Buffon’s 

(1761) theory of New World degeneracy had argued that the 

inferior climate and soils of the New World resulted in a depleted 

American biota of plants, animals, and humans that were smaller, 

less vigorous, less fertile versions of Old World species. Although 

Buffon (1780) later repudiated his theory, its effects lingered, as 

did notions of a shared, if impoverished, European fauna in the 

Americas. Partly for this reason, early lists of American mammals often include European species (Ord, 1815; Harlan, 1825). 

At the same time, new species unique to the New World were 

being noticed. Efforts to document diversity here and elsewhere 

were hampered by the dearth of adequate libraries, the paucity 

of publication outlets, the slow speed of communication, and 

the lack of widely accepted standards for naming and classifying taxa. Many species were described on the basis of published 

accounts and/or images provided by explorers and travelers, the 

describer never having seen the animal either alive or as a specimen. It was not uncommon for the same traveler’s account to be 

used by as the basis for multiple new species by different authors.

Ultimately, not all new species names maintained currency, 

and a number no longer appear in standard synonymies, making 

it difficult to trace their origins or understand their historical or 

biological context. To assist with understanding early names potentially referring to eulipotyphlans, I compiled the following alphabetical listing of taxonomic names that may be encountered 

in the early literature. The following accounts also provide an 

object lesson regarding the value of requiring a physical holotype 

for describing a new species or subspecies and the continuing 

usefulness of preserving type specimens.

Blarina pyrrhonota Jentinck, 1910.

1888. Blarina pyrrhonota Jentinck, Catalogue systématique des mammifères, 

p. 131 (nomen nudum).

1910. Blarina pyrrhonota Jentinck, Notes from the Leyden Museum 32:167.

1925. Cryptotis pyrhonota: Cabrera, Gen. Mamm. Insect. Galeop., p. 135; 

29 November. Incorrect subsequent spelling of Blarina pyrrhonota

Jentinck, 1910.

1958. Cryptotis surinamensis: Cabrera, Rev. Mus. Argentino 4(1):47 (not 

Sorex surinamensis Gmelin, 1788).

1963. Sorex araneus: Husson, Studies on the Fauna of Suriname and Other 

Guyanas 5(13):336; January.

Type Locality: “Suriname.”

Comments: The name Blarina pyrrhonota first appeared in 

the published catalog of the Muséum d’Histoire Naturelle 

des Pays-Bas (Jentinck, 1888:131), in which a specimen is 

listed as the type of the species. The species was not described, however, relegating the name to the status of nomen 
6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

nudum. The name was subsequently made available when 

Jentinck (1910) formally described B. pyrrhonota. Cabrera 

(1958:47) treated the name as a junior synonym of Sorex

surinamensis Gmelin, 1788, which he referred to the genus 

Cryptotis. After study of the holotype and its history, Husson (1963) concluded that the holotype of B. pyrrhonota is 

a common European shrew, Sorex araneus Linnaeus, 1758, 

that was mislabeled as to its origin.

Condylura fissipes

1815. Condylura fissipes: Illiger, Abhand. Physik. Klasse, p. 70.

1821. [Talpasorex] fissipes: Schinz, Das Thierreich, p. 191.

1821. [Talpa] fissipes: Schinz, Das Thierreich, p. 191.

1821. Codylura fissipes: Schinz, Das Thierreich, p. 191.

Comments: The origin of the name Condylura fissipes remains 

obscure; neither True (1896) nor Jackson (1915) could determine its author. The earliest mention that I discovered 

was Illiger (1815:70), who included the name C. fissipes

(but lacking description or synonymy) in his list of mammal 

species indigenous to North America. The name is lacking 

from an earlier taxonomic classification (Illiger, 1811:125–

126), indicating it came to his attention between 1811 and 

1815. Schinz (1821:191) and Minding (1829:64) referred 

the taxon to the genus Talpasorex, and the name has generally been treated as a junior synonym of Condylura cristata (Linnaeus, 1758). If the name proves to be other than 

a nomen nudum, however, it is more likely to refer to one 

of the other species of northeastern North American moles.

Musaraneus brasiliensis

(see Sorex americanus)

Sorex americanus Müller, 1776

1648. Mus araneus Marcgrave, in Piso, Historia Naturalis Brasiliae, p. 229 

(pre-Linnaean work).

1756. Musaraneus brasiliensis Brisson, Regnum animale, p. 180 (preLinnaean work).

1762. Musaraneus brasiliensis Brisson, Regnum animale, p. 127 (excluded 

work).

1776. Sorex americanus Müller, Voll. Natursyst. Suppl., p. 36. 

1777. [Sorex] brasiliensis: Erxleben, Syst. Reg. Anim., p. 127.

1777. Sorex Brasiliensis: Zimmermann, Spec. Zool. Geog., p. 502.

Type Locality: “Brasilien.” Thomas (1911:124) noted that 

Marcgrave stayed at Moritzstadt (Mauritsstad, Stadt Mauritius, or Cidade Maurícia; now Recife) in the state of Pernambuco most of the time he was in Brazil (1640–1644), 

and this should be taken as the central locality for most of 

his specimens.

Comments: Sorex americanus was based on the “Musaraigne 

du Bresil” of Buffon (1767:160) and the “Brasiliaansche 

Spitsmuis” of Boddaert (1772:52), which in turn were based 

on the earlier description of what Marcgrave (1648:229) 

called Mus araneus. This name is of interest because Mus 

araneus (“spider mouse”) is a classical Latin term for 

“shrew,” and it is clear from later authors’ referral of the 

animal to the genus Sorex that they considered it a soricid. Musaraneus brasiliensis, however, was an animal from 

Brazil, where soricids are not known to occur. Marcgrave’s 

(1648) original description is now recognized as referring 

to the South American didelphid marsupial, Monodelphis 

americana (Müller, 1776). For the complete synonymy for 

that species, see Pine and Handley (2008).

Sorex avellanarum É. Geoffroy Saint-Hilaire, 1803

FIGURE 3

1734. Glis, seu Mus avellanarum Seba, Thesaurus, vol. 1, p. 50, tab. 31: fig. 

7 (pre-Linnaean work).

1803. Sorex avellanarum É. Geoffroy Saint-Hilaire, Cat. Mamm. Mus. Natl.

Hist. Nat., p. 74.

Holotype: “No. CLXVI. Cet individu très-précieux est conservé dans la liqueur; il a été donné par M. Van-Marum, 

qui l’avoit acquis à la vente du cabinet de Meyer: c’est sans 

doute l’original de la figure qu’en a donnée Seba” (No. 

CLXVI. This very precious individual is preserved in fluid;

it was donated by M. Van-Marum, who had acquired it at

the sale of Meyer’s cabinet; it is doubtless the original of the

figure given by Seba).

Type Locality: “L’Amérique, selon Seba.”

Comments: É. Geoffroy Saint-Hilaire (1803) named “la musaraigne blanche” (the white shrew) based on Seba’s (1734:50) 

description and illustration (Figure 3) of Glis, seu Mus Avellanarum (Glis or Mus Avellanarum) and a specimen in the 

Muséum National d’Histoire Naturelle of Paris upon which 

Geoffroy Saint-Hilaire thought Seba had based his account. 

The illustration shows a somewhat stocky animal with 

relatively large (for a soricine) ears and eyes and a robust, 

tapering tail that, in overall appearance, looks very much 

like a very pale, leucistic, or albinistic crocidurine shrew. If 

so, it could not be from the New World. The Latin name 

Mus avellanarum was earlier applied to certain dormice 

(Gliridae: Gesner, 1560:115; Ray, 1693:219), particularly 

the hazel dormouse, Muscardinus avellanarius (Linnaeus, 

1758:62).

Sorex cerulescens Rafinesque, 1818

1818. Sorex cerulescens Rafinesque, Am. Month. Mag. 3:446.

Type Locality: USA: “The western states” [Kentucky].

Comments: The “bluish shrew mouse” is one of 10 species of 

mammals (and 20 other animals) that were invented by John 

James Audubon and described to Constantine S. Rafinesque 

as a prank during the latter’s visit to Audubon’s home in 

Henderson, Kentucky, in 1818. Unfortunately, Rafinesque 

naively took Audubon at his word and subsequently described the faux animals as new species (Woodman, 2016).
NUMBER 650 • 7

Sorex constrictus Hermann, 1780.

1780. Sorex constrictus Hermann, in Zimmermann, Geog. Gesch. Mensch.

Vierf. Thiere 2:383.

Comments: Sorex constrictus is a junior synonym of the western European and north African species Sorex russula Hermann, 1780 (see Hutterer, 2005). On the basis of a specimen 

(PM 895) in Peale’s Philadelphia Museum that had been 

obtained near Philadelphia, Harlan (1825) indicated that 

S. constrictus occurred in North America as well as in Europe. Unfortunately, no Philadelphia Museum specimens of 

mammals are known to have survived (Woodman, 2009), so 

it is now impossible to know the true identity of the animal.

Sorex dichrurus Rafinesque, 1833

1833. Sorex dichrurus Rafinesque, Atlantic J. 1(6):175, summer.

1840. Sorex dicrurus Rafinesque, The Good Book and Amenities of Nature, 

p. 68. Incorrect subsequent spelling of Sorex dichrurus Rafinesque, 

1833.

Holotype: A specimen, now probably lost, then in the Niagara 

Falls Museum (Woodman, 2012b).

Type Locality: USA: “the falls of Niagara; . . . Goat Island in 

the middle of the falls,” Niagara Co., New York.

Remarks: This “shrew” is a misidentified meadow jumping 

mouse, Dipus hudsonius Zimmermann, 1780:358 (=Zapus 

hudsonius) (Woodman, 2012b).

Sorex exilis Gmelin, 1788

1788. [Sorex] exilis Gmelin, Systema Naturae, 13th ed., p. 115.

Type Locality: Russia: “Sibiria, ad fluvium Jenisei” (Siberia, at 

the Yenisei River).

Comments: A junior synonym of the Eurasian pygmy shrew, 

Sorex minutus Linnaeus, 1766, Ord (1815) listed S. exilis as 

part of the North America vertebrate fauna.

Sorex fimbripes Bachman, 1837

FIGURE 4

1837. Sorex fimbripes Bachman, J. Acad. Nat. Sci. Phila. 7(2):391. 

1863. Neosorex(?) fimbripes: Verrill, Proc. Boston Soc. Nat. Hist. 9:167; 

February.

1868. Crossopus fimbripes: Fitzinger, Sitzungsber. Math.-natürwissensch.

Classe Kaiserl. Akad. Wissensch. 57(1):631.

1868. Crocidura fimbripes: Fitzinger, Sitzungsber. Math.-natürwissensch.

Classe Kaiserl. Akad. Wissensch. 57(1):631 (in synonymy).

1868. Croscopus? fimbripes: Fitzinger, Sitzungsber. Math.-natürwissensch.

Classe Kaiserl. Akad. Wissensch. 57(1):632 (in synonymy).

1877. Sorex fimbriata Holder et al., Hist. Am. Fauna, vol. 3, p. xxxii. Incorrect subsequent spelling of Sorex fimbripes Bachman, 1837. 

1877. Sorex fimbriata Richardson et al., Mus. Nat. Hist., vol. 3, p. xxxii. 

Incorrect subsequent spelling of Sorex fimbripes Bachman, 1837. 

1930. Sorex palustris fimbriata: Green, Contr. Mamm. N. Mtn. Reg., p. 11 

(31 March).

Holotype: None known to exist. Baird (1861:55–56) stated 

that he inspected “the type of fimbripes” in the Academy 

of Natural Sciences of Philadelphia (ANSP) in 1861. Unfortunately, any such specimen of S. fimbripes no longer 

exists there (Ned Gilmore, Academy of Natural Sciences 

of Drexel University, Philadelphia, email communication, 

FIGURE 3. Sorex avellanarum É. Geoffroy-Saint-Hilaire, 1803, from Seba (1734: pl. 31, fig. 7). Image courtesy of Biodiversity Heritage Library 

(https://www.biodiversitylibrary.org/), contributed by Smithsonian Institution Libraries, Washington.
8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

14 February 2018; see also Koopman, 1976). Lyon and 

Osgood (1909:243) reported the supposed rediscovery of 

the “type” of S. fimbripes (USNM 84556) in the collection of the National Museum of Natural History, Washington, D.C. (see also Poole and Schantz, “1942:185; Fisher 

and Ludwig, 2015:43). However, the only information associated with this specimen was an old label of unknown 

original bearing the handwritten words “Sorex fimbripes. 

Type.” Although it is possible that Bachman’s original 

holotype was returned to ANSP, from which it was subsequently loaned to Baird at the USNM, there is no known 

correspondence or other record that supports this scenario. 

Hollister (1911:381) subsequently identified USNM 84556 

as Sorex fumeus Miller, 1895, an identification with which I 

concur. He discounted that this specimen was the holotype, 

as it did not, in his estimation, match Bachman’s (1837) 

description. Hollister (1911) further noted that in the time 

of Baird, the word type could simply refer to an example of 

a species, and Baird (1857) himself did not designate holotypes. Moreover, my comparison of external measurements 

of USNM 84556 with those reported by Bachman (1837) 

for the holotype and Baird (1861) for the ANSP “type” 

suggests that none are from the same specimen. It seems 

more likely that the “types” of S. fimbripes in both ANSP 

and USNM were simply the examples each institution had 

of that species. The holotype was probably destroyed during the American Civil War.

Type Locality: USA: “high table-land on a branch of Drury’s 

Run, a tributary of the west branch of the Susquehannah 

river,” Clinton County, Pennsylvania.

Remarks: John Bachman described S. fimbripes from a specimen sent to him from the ANSP. Bachman’s (1837:393) 

characterization of the “fringe-footed shrew” as having 

feet whose “edges on the lower surface are fringed considerably” and fore feet that bear “some resemblance to the 

fins of a turtle” strongly suggests the animal was a water 

shrew (Sorex palustris group). This impression is further 

substantiated by Bachman’s statement that the species bore 

some resemblance to “the Genus Mygale of Cuvier,” an 

aquatic desman (Mygale Cuvier, 1800 = Desmana Güldenstaedt, 1777). However, Bachman’s (1837:392, pl. 24, 

fig.8) description also stated that the body of S. fimbripes

was “a little less in size than that of Forster’s Shrew,” Sorex 

forsteri Richardson, 1828 (=S. cinereus Kerr, 1792), much 

too small for any known species of North American semiaquatic shrew, and his illustration of the animal suggests a 

small-sized terrestrial species (Figure 4). Since then, Sorex 

fimbripes has been treated variably as a species of semiaquatic shrew (Pomel, 1848; Baird, 1857; Fitzinger, 1858); 

a species of ambulatory shrew (Baird, 1861); an eastern 

subspecies of the water shrew, Sorex palustris Richardson, 

1828 (Green, 1930); a synonym of S. personatus I. Geoffroy Saint-Hilaire, 1827 (=S. cinereus Kerr, 1792) (Miller, 

1895); a synonym of S. c. cinereus Kerr, 1792 (Jackson, 

1928; Hall, 1981; Hutterer, 2005); and entirely unidentifiable (Hollister, 1911; Handley and Varn, 1994). Lacking a 

clear holotype, the last option is the most appropriate for 

this species.

FIGURE 4. John Bachman’s (1837) shrews: from top to bottom, 

Sorex richardsonii Bachman, 1837 (=Sorex arcticus Kerr, 1792); 

Sorex forsteri Richardson, 1828 (=Sorex cinereus Kerr, 1792); Sorex 

cooperi Bachman, 1837 (=Sorex cinereus Kerr, 1792); Sorex fimbripes Bachman, 1837. Image from Bachman (1837: pl. 24).
NUMBER 650 • 9

Sorex melanotis Rafinesque, 1818

FIGURE 5

1818. Sorex melanotis Rafinesque, Am. Month. Mag. 3:446.

Type Locality: USA: “The western states.” [Kentucky].

Comments: The “black-eared shrew” or “corn mouse” is 

a species that was invented by John James Audubon and 

described to Constantine S. Rafinesque as a prank during 

the latter’s 1818 visit to Audubon’s home in Henderson, 

Kentucky. Unfortunately, the naïve Rafinesque took Audubon at his word and described the animal as new species 

(Woodman, 2016).

Sorex mexicanus Kerr, 1792

1792. Sorex mexicanus Kerr, Animal Kingdom, p. 207.

Type Locality: “Mexico and New Spain.”

Comments: Ord (1815) included the “Mexican shrew,” Sorex 

mexicanus, in his comprehensive listing of the North American terrestrial vertebrate fauna on the basis of Turton 

(1802:72). Turton probably based his description on the 

work of Kerr (1792), who referenced the “Mexican shrew” 

of Pennant (1771:309), an account by Buffon (1767:159), 

and the original account of the “tucan” by Hernandez 

(1651: book 1, tract 1, p. 7, chapter XXIV). Hernandez 

considered the animal to be a kind of mole, and Buffon 

thought it might be the same as Seba’s (1734) “red mole.” 

More likely, as noted by Rhoads (1894a, 1894b: Appendix, 

p. 15), the species described by Hernandez was a pocket 

gopher (Geomyidae). Alfred L. Gardner (U. S. Geological 

Survey, Biological Survey Unit, personal communication, 

16 October 2017) pointed out that “tucan” is likely a variant or misunderstanding of tuza, a local common name for 

the pocket gopher in much of Mexico and Central America. 

Lichtenstein (1830:113) thought “tucan” derived from “der 

entstellte spanische des Maulwurfs Tuçe oder Toço” (the 

distorted Spanish for the mole, Tuçe or Toço). There may be 

an additional etymological connection to tuco-tuco, a local 

common name for a southern South American burrowing 

rodent (Ctenomyidae: Ctenomys Blainville, 1826).

Sorex surinamensis Gmelin, 1788

1788. [Sorex] surinamensis Gmelin, Systema Naturae, 13th ed., p. 114.

1868. Crocidura surinamensis: Fitzinger, Sitzungsber. Math.-natürwissensch.

Classe Kaiserl. Akad. Wissensch. 57(1):633 (in synonymy).

1888. Didelphys brevicaudata: Thomas, Cat. Marsup. Monotrem., p. 357.

1919. Peramys brevicaudatus: Cabrera, Gen. Mamm. Monot. Marsup., p. 42.

1958. Cryptotis surinamensis: Cabrera, Rev. Mus. Argentino 4(1):47.

Type Locality: “Suriname.”

Comments: Relying on Gmelin’s (1788) description, Thomas 

(1888:357) and Cabrera (1919:42) believed that Sorex surinamensis represented the didelphid mouse opossum, Monodelphis brevicaudata (Erxleben, 1777). In contrast, Tate 

(1932:223) thought Gmelin’s description could “apply only 

to a shrew,” but “the description is too general for specific 

recognition to be possible.” He entertained the possibility 

that S. surinamensis and Blarina pyrrhonota Jentinck, 1910 

were based on the same specimen because both were thought 

to be shrews found in Suriname, although he also cast doubt 

on their provenances. Cabrera (1958) also subsequently 

treated S. surinamensis as a shrew. Husson (1963:35) was 

FIGURE 5. Pen-over-pencil sketch of the “black-eared shrew” or “corn mouse,” Sorex melanotis Rafinesque, 1818b from Rafinesque’s field 

notebook 17. Image SIA 2012-6096, courtesy of Smithsonian Institution Archives, Washington, D.C.
1 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the first to study the holotype of B. pyrrhonota since Jentinck (1910) described the species, and he wrote that there 

is no evidence supporting the supposition that the holotype 

of B. pyrrhonota is also the holotype of S. surinamensis. 

The holotype of S. surinamensis is lost, but no species of 

soricids are known from Suriname, and the description is 

almost certainly that of a mouse opossum.

Talpa rubra Erxleben, 1777

FIGURE 6

1734. Talpa, rubra, Americana Seba, Thesaurus, vol. 1, tab. XXXII, no. 2 

(pre-Linnaean work).

1777. [Talpa] rubra Erxleben, Syst. Reg. Anim., p. 119.

1800. Talpa rufa Shaw, Gen. Zool. 1(2):522 (replacement name for Talpa

rubra Erxleben, 1777). 

1820. Chrysochloris rufa: Desmarest, Mammalogie 1:156.

Type Locality: “America.”

Comments: Ord (1815) included Talpa rubra in his list of 

North American terrestrial vertebrates. Talpa rubra Erxleben, 1777 is based on the “red mole” of Pennant (1771:315) 

and “Roode Mol” of Boddaert (1772:51). It originated in 

the Talpa, rubra, Americana of Seba (1734), which was described as having three toes on the fore feet and four toes on 

the hind feet (Figure 6). Buffon (1767) conflated this with 

Hernandez’s (1651) “tucan” (see Sorex mexicanus Kerr, 

1792); hence, it may also be based partly on the description 

of a geomyid (Rhoads, 1894a, 1894b). Talpa rubra was justifiably considered unidentifiable by Rhoads (1894b).

RECENT AMERICAN EULIPOTYPHLA

Order Eulipotyphla Waddell, Okada, 

and Hasegawa, 1999

1999. Eulipotyphla Waddell, Okada, and Hasegawa, Syst. Biol. 48(1):1–5; 

March.

Family Nesophontidae Anthony, 1916

1916. Nesophontidae Anthony, Bull. Am. Mus. Nat. Hist. 35:728;

16 November.

Genus Nesophontes Anthony, 1916

1916. Nesophontes Anthony, Bull. Am. Mus. Nat. Hist. 35:725; 

16 November.

FIGURE 6. The “red mole,” Seba’s (1734: pl. 32, fig. 2) Talpa, rubra, Americana. Image courtesy of Biodiversity Heritage Library (https://www

.biodiversitylibrary.org/), contributed by Smithsonian Institution Libraries, Washington, D.C.
NUMBER 650 • 1 1

Type Species: Nesophontes edithae Anthony, 1916 (by original 

designation).

Remarks: The genus Nesophontes includes six described species and possibly two undescribed species (one from Grand 

Cayman, the other from Cayman Brac; Morgan, 1994; 

MacPhee et al., 1999). Known primarily from skeletal remains, these species are all extinct, although stratigraphic 

associations with introduced exotic species and evidence 

from radiocarbon dating indicate that at least four species (Nesophontes hypomicrus, N. micrus, N. paramicrus, 

N. zamicrus) survived into post-Columbian time (Miller, 

1929, 1930; MacPhee et al., 1999; Brace et al., 2016). The 

genus was reviewed by Hall (1981) and the Cuban species 

were revised by Condis Fernández et al. (2005).

Nesophontes edithae Anthony, 1916

1916. Nesophontes edithae Anthony, Bull. Am. Mus. Nat. Hist. 35:725; 

16 November.

Holotype: AMNH VP 14174.

Type Locality: Puerto Rico: “Cueva Catedral, near Morovis, 

Porto Rico.” 

Distribution: Puerto Rico, Vieques, St. Johns, St. Thomas (MacPhee et al., 1999).

Remarks: This is the type species of the genus Nesophontes

Anthony, 1916. Nesophontes edithae is extinct and known 

only from skeletal remains.

Nesophontes hypomicrus Miller, 1929

1929. Nesophontes hypomicrus Miller, Smithson. Misc. Coll. 81(9):4; 

30 March.

Holotype: USNM PAL 367223 (previously USNM 253077).

Type Locality: Haiti: “the deep cave near the Atalaye plantation, Haiti.” Miller and Kellogg (1955:53) amended the 

type locality to “cave near Atalaye plantation, about 4 miles 

east of St. Michel.” 

Distribution: Hispaniola and Gonâve Island (Miller, 1930; 

MacPhee et al., 1999).

Remarks: Nesophontes hypomicrus is extinct and known only 

from skeletal remains, although there is evidence the species survived into post-Columbian time. Remains of Nesophontes hypomicrus, N. paramicrus, and N. zamicrus were 

found mixed with those of the introduced rodents Rattus 

rattus and Mus musculus in seemingly undisturbed cave sediments on Haiti (Miller, 1929). Bones of these three species 

of Nesophontes, together with those of R. rattus, were also 

discovered in a dried mass of partly disintegrated owl pellets in a shelter on the north flank of Monte Culo de Maco, 

about 10 km southwest of Constanza, Dominican Republic (Miller, 1930). A skull of N. hypomicrus from Cueva 

de Bosque Humedo, Dominican Republic, was accelerator 

mass spectrometry (AMS) radiocarbon dated to 734 ± 24 

calendar years before present (cal ybp; circa ad 1247–1294; 

Brace et al., 2016).

Nesophontes major Arredondo, 1970

1970. Nesophontes major Arredondo, Mem. Soc. Cienc. Nat. La Salle

30(86):126; August.

Holotype: GEC P682.

Type Locality: Cuba: “Cueva de La Santa, Bacuranao, provincia de La Habana.”

Distribution: Cuba (MacPhee et al., 1999).

Remarks: This species is extinct and is known only from skeletal remains.

Nesophontes micrus G. M. Allen, 1917

1917. Nesophontes micrus G. M. Allen, Bull. Mus. Comp. Zool. 61:5;

January.

1919. Nesophontes longirostris Anthony, Bull. Am. Mus. Nat. Hist. 41:633; 

30 December. Holotype: AMNH VP 17626, from “a cave near the 

beach at Daiquiri, Cuba.”

1970. Nesophontes submicrus Arredondo, Mem. Soc. Cienc. Nat. La Salle

30(86):137; August. Holotype: GEC 740, from “Cueva de La Santa, 

Bacuranao, provincia de La Habana,” Cuba. 

1977. Nesophontes superstes K. Fischer, Zeit. Geol. Wiss. 5(2):221; February. Holotype: MB Ma. 1967/258/1, from “Cueva de la Ventana, 

über der Pio Domingo-Höhle gelegen, Sumerido, Provinz Pinar del 

Rio,” Cuba.

Holotype: MCZ 9600.

Type Locality: Cuba: “cavern in the Sierra of Hato-Nuevo, 

Province of Matanzas.”

Distribution: Cuba, Hispaniola, and the Isle of Pines (MacPhee 

et al., 1999).

Remarks: This species is extinct and is known only from skeletal remains. Remains from Cuba dated to as late as 590 ± 

50 cal ybp (circa ad 1400), indicating that it survived into 

the late Holocene (MacPhee et al., 1999). Nesophontes longirostris, N. submicrus, and N. superstes were synonymized 

with N. micrus by Condis Fernández et al. (2005).

Nesophontes paramicrus Miller, 1929

1929. Nesophontes paramicrus Miller, Smithson. Misc. Coll. 81(9):3; 

30 March.

Holotype: USNM PAL 367224 [previously USNM 253063].

Type Locality: Haiti: “front of large cave near St. Michel, 

Haiti.” Miller and Kellogg (1955:53) amended the type locality to “cave near Atalaye plantation, about 4 miles east 

of St. Michel.” 

Distribution: Hispaniola (MacPhee et al., 1999).

Remarks: Nesophontes paramicrus is extinct and is known 

only from skeletal remains, although there is evidence it 

survived into post-Columbian time. Remains of this species, 
1 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

N. hypomicrus, and N. zamicrus were found mixed with 

those of the introduced rodents Rattus rattus and Mus musculus in seemingly undisturbed cave sediments on Haiti 

(Miller, 1929). Bones of the three species of Nesophontes, together with those of R. rattus, were also discovered in a dried 

mass of partly disintegrated owl pellets in a shelter on the 

north flank of Monte Culo de Maco, about 10 km southwest 

of Constanza, Dominican Republic (Miller, 1930). Remains 

of N. paramicrus were AMS radiocarbon dated to as late as 

680 ± 50 cal ybp (circa ad 1295; MacPhee et al., 1999).

Nesophontes zamicrus Miller, 1929

1929. Nesophontes zamicrus Miller, Smithson. Misc. Coll. 81(9):7; 30 March.

Holotype: USNM PAL 367225 [previously USNM 253090].

Type Locality: Haiti: “large cave near St. Michel, Haiti.” Miller 

and Kellogg (1955:53) amended the type locality to “cave 

near Atalaye plantation, about 4 miles east of St. Michel.” 

Distribution: Hispaniola (MacPhee et al., 1999).

Remarks: This species is extinct and known only from skeletal remains, although it appears to have survived into 

post-Columbian time. Remains of Nesophontes hypomicrus, N. paramicrus, and N. zamicrus were found mixed 

with those of the introduced rodents Rattus rattus and Mus 

musculus in seemingly undisturbed cave sediments on Haiti 

(Miller, 1929). Bones of the three species of Nesophontes, 

together with those of R. rattus, were also discovered in a 

dried mass of partly disintegrated owl pellets in a shelter on 

the north flank of Monte Culo de Maco, about 10 km southwest of Constanza, Dominican Republic (Miller, 1930).

Family Solenodontidae Gill, 1872

1872. Solenodontinae Gill, Smithson. Misc. Coll. 11(1):19; November. 

Proposed as a subfamily.

Genus Solenodon Brandt, 1833

1833. Solenodon Brandt, Mém. Acad. Imp. Sci. St. Petersbourg, ser. 6, Sci.

Math. Phys. et Nat. 2:459, tabs. 1–2. 

1925. Atopogale Cabrera, Gen. Mamm. Insect. Galeop., p. 177; 29 November. Type species: Solenodon cubanus Peters, 1861, by original 

designation.

1962. Antillogale Patterson, Breviora 165:2; 22 August. Type species: Antillogale marcanoi Patterson, 1962, by original designation.

Type Species: Solenodon paradoxus Brandt, 1833 (by monotypy).

Remarks: The genus includes four described species, two 

of which are extinct. It was reviewed by Hall and Kelson 

(1959), Hall (1981), and Ottenwalder (2001).

Solenodon arredondoi Morgan and Ottenwalder, 1993

1993. Solenodon arredondoi Morgan and Ottenwalder, Ann. Carnegie Mus. 

62(2):154; 28 May.

Holotype: MNHNC 421/123.

Type Locality: Cuba: “Cueva Paredones, 3 km SW Ceiba del 

Agua San Antonio de los Baños, La Habana Province, Cuba.”

Distribution: Western Cuba: known only from La Habana and 

Pinar del Río provinces (Ottenwalder, 2001).

Remarks: Solenodon arredondoi is extinct. It is known only 

from skeletal remains that are considered to be late Pleistocene in age (Ottenwalder, 2001).

Solenodon cubanus Peters, 1861

FIGURE 7

1861. Solenodon cubanus Peters, Monatsb. Preuss. Akad. Wiss. Berlin

1862:169.

1925. Atopogale cubana: Cabrera, Gen. Mamm. Insect. Galeop., p. 177; 29 

November.

1944. Solenodon poeyanus Barbour, Proc. New Engl. Zool. Club 23:6; 7 

March. Holotype: MCZ 6957, from “vicinity of Nipe Bay,” northeastern Cuba.

Lectotype: ZMB 2761, designated by Turni et al. (2007).

Type Locality: Cuba: “Aguilera, am Abhange der Sierra 

Maestra, zwischen Cap Maisi und Cap Cruz.”

Distribution: Southeastern Cuba.

Remarks: Peters’s (1861) original description of Solenodon cubanus was rather cursory. He subsequently provided a more 

complete account with figures (Peters, 1863). This is the 

type species of Atopogale Cabrera, 1925.

Solenodon marcanoi (Patterson, 1962)

1962. Antillogale marcanoi Patterson, Breviora 165:3; 22 August.

Holotype: MCZ 7261.

Type Locality: Dominican Republic: “unnamed cave 2 kilometers SE of Rancho La Guardia, Municipio de Hondo Valle, 

Provincia de San Rafael.” 

Distribution: Southern Hispaniola: known from La Hotte and 

La Selle massifs, Haiti, and Sierra de Neiba, Dominican Republic (Ottenwalder, 2001).

Remarks: This is the type species for Antillogale Patterson, 

1962. The species is extinct, although well-preserved remains were found together with those of the introduced rodent Rattus, indicating that it survived into post-Columbian 

times (Ottenwalder, 2001).

Solenodon paradoxus Brandt, 1833

FIGURE 8

Remarks: This is the type species for Solenodon Brandt, 1833. 

Ottenwalder (2001) showed that S. paradoxus comprised 

two distinct allopatric subspecies, S. p. paradoxus north 

of the Neiba Valley and S. p. woodi south of that valley. 

This division is provisionally supported by mitogenomic 

data (Brandt et al., 2017). A combined morphological 

and molecular study by Turvey et al. (2016) purported to 

show a similar deep division within S. p. woodi that further 
NUMBER 650 • 1 3

differentiated a population restricted to La Hotte massif 

in southwestern Haiti. They called this population S. p.

haitiensis but failed to publish the distinguishing morphological or molecular characters, to designate a holotype, or 

to otherwise describe the new taxon, rendering the name 

unavailable. Notwithstanding the lack of adequate methodology, it is possible that S. p. woodi may be divisible into 

two genetically distinctive populations.

Solenodon paradoxus paradoxus Brandt, 1933

FIGURE 8

1833. Solenodon paradoxus Brandt, Mém. Acad. Imp. Sci. St. Petersbourg,

ser. 6, Sci. Math. Phys. et Nat. 2:459.

2001. Solenodon paradoxus paradoxus: Ottenwalder, in Woods and Sergile, 

Biogeography of the West Indies, p. 301.

Holotype: RAS 982 (Baranova et al., 1981).

FIGURE 7. External view (top) and skeleton (bottom) of the Cuban solenodon, Solenodon cubanus Peters, 1861. Images from 

Peters (1861: pls. 1, 3).
1 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Type Locality: “Insula Hispaniola.”

Distribution: Hispaniola: populations north of the Neiba Valley (Ottenwalder, 2001).

Solenodon paradoxus woodi Ottenwalder, 2001

2001. Solenodon paradoxus woodi Ottenwalder, in Woods and Sergile, Biogeography of the West Indies, p. 299.

2016. Solenodon paradoxus haitiensis Turvey et al., Diver. Distr. 22:597 

(unavailable name).

Holotype: UF 30135.

Type Locality: Dominican Republic: “Bucan de Tuí, S. Oviedo, 

Península de Barahona, Provincia Pedernales.”

Distribution: Hispaniola: south of the Neiba Valley (Ottenwalder, 2001; Turvey et al., 2016).

Family Talpidae G. Fischer, 1814

1814. Talpidae G. Fischer, Zoognosia 3:x.

Remarks: North American talpids were revised by True (1896) 

and Jackson (1915). On the basis of the molecular phylogenetic study by Shinohara et al. (2003), Hutterer (2005) 

recognized three subfamilies within the Talpidae: Scalopinae (Condylura, Parascalops, Scalopus, Scapanulus, 

Scapanus), Talpinae (Desmana, Dymecodon, Euroscaptor, 

Galemys, Mogera, Neurotrichus, Parascaptor, Scaptochirus, 

Scaptonyx, Talpa, Urotrichus), and Uropsilinae (Uropsilus). 

Subsequent studies have shown different arrangements of 

tribes and genera within the Talpidae that generally render 

this concept of the Scalopinae and Talpinae polyphyletic 

(Schwermann and Thompson, 2015; He et al., 2016; Hooker, 

2016). Because the relationships among the tribes and genera 

remain uncertain, I do not recognize subfamilies here.

Tribe Condylurini Gill, 1875

1875. Condylurae Gill, Synop.Insectiv. Mamm., p. 110; 14 May. Proposed 

as a suprageneric taxon of unidentified rank.

Genus Condylura Illiger, 1811

1811. Condylura Illiger, Prodr. Syst. Mamm. Avium, p. 125. 

1821. Talpasorex Schinz, Theirreich 1:191 (not Talpasorex Lesson, 1827). 

Type species: Sorex cristatus Linnaeus, 1758.

1821. Codylura Schinz, Theirreich 1:191. Incorrect subsequent spelling of 

Condylura Illiger, 1811.

1825. Astromycter Harris, East. Star 2(24 or 25):??; May. Type species: 

Condylura prasinata Harris, 1825 (=Sorex cristatus Linnaeus, 1758). A 

summary notice by Benjamin Silliman in the June 1825 issue of American Journal of Science (Harris, 1825c) is typically cited as the original 

description of Astromycter, but Harris (1825a) published a more complete description in May.

1830. Rhinaster Wagler, Natürl. Syst. p. 14. Type species: Sorex cristatus

Linnaeus, 1758.

1838. Condylurus: Blainville, Ann. Franç. Anat. Phys. 2:219. Unjustified 

emendation of Condylura Illiger, 1811. Proposed as a subgenus of 

Talpa Linnaeus, 1758.

1839. Condytura Bell, Cyclop. Anat. Phys. 2:994. Incorrect subsequent 

spelling of Condylura Illiger, 1811.

FIGURE 8. The Hispaniolan solenodon, Solenodon paradoxus Brandt, 1833. Image from Brandt (1833: tab. 1).
NUMBER 650 • 1 5

1843. Astromyctes Gray, Spec. Mamm. Brit. Mus., p. 76. Incorrect subsequent spelling of Astromycter Harris, 1825.

1863. Astromydes Blyth, Cat. Mamm. Asia. Soc. Mus., p. 87. Incorrect subsequent spelling of Astromycter Harris, 1825.

1913. Condylus Van Hyning, Science 38(972):243; 15 August. Incorrect subsequent spelling of Condylura Illiger, 1811.

Type Species: Sorex cristatus Linnaeus, 1758. Illiger (1811) 

included two species in his new genus Condylura: Sorex 

cristatus Linnaeus, 1758 and Talpa longicaudata Erxleben, 

1777 (=Scalops breweri Bachman, 1841).

Remarks: Condylura was revised by True (1896) and Jackson 

(1915). It was reviewed by Hall and Kelson (1959), Petersen 

and Yates (1980), and Hall (1981). Sansalone et al. (2016b) 

reviewed fossil Condylura, including a middle Miocene 

specimen from Kazakhstan that supports a Eurasian, rather 

than North American, origin for the genus. The genus has 

also been recorded from the Plio-Pleistocene of Poland 

(Skoczeń, 1993; Sansalone et al., 2016a).

Condylura cristata (Linnaeus, 1758)

FIGURES 9, 10

Remarks: This is the type species for the genus Condylura Illiger, 1811.

Condylura cristata cristata (Linnaeus, 1758)

1758. [Sorex] cristatus Linnaeus, Systema Naturae, 10th ed., 1:53.

1769. Talpa Canadensis De la Faille, Essai Hist. Nat. Taupe, p. 40, pl. 1. 

Holotype: a dried specimen (probably no longer extant) in De la Faille’s 

collection, obtained from M. Dabbadie, governor of Louisiana from 

“Canada.”

1777. Talpa Cristata: Zimmermann, Spec. Zool. Geog., p. 496.

1800. Talpa radiata Shaw, Gen. Zool. 1(2):523; based on Pennant’s (1771) 

radiated mole and Sorex cristatus Linnaeus, 1758.

1800. Sorex radiatus Shaw, Gen. Zool. 1(2):531; based on the description and 

figure of Talpa canadensis De La Faille, 1778 from Canada (Figure 9).

1803. Scalopus cristatus: É. Geoffroy Saint-Hilaire, Cat. Mamm. Mus. Natl.

Hist. Nat., p. 77.

1814. [Scalops] cristatus: G. Fischer, Zoognosia 3:156.

1819. Condylura cristata: Desmarest, J. Phys. Chim. Hist. Nat. Arts 89:230, 

pl. 2; September (first use of current name combination for the species).

1821. Tal[pasorex]. cristata: Schinz, Thierreich 1:191.

1821. Talpa flava: Schinz, Thierreich 1:191. Part; not Talpa flava Zimmermann, 1777 = Sorex aquaticus Linnaeus, 1758 (=Scalopus aquaticus).

1821. [Talpa] purpurascens: Schinz, Thierreich 1:191. Part; not Talpa purpurascens Shaw, 1800 = Sorex aquaticus Linnaeus, 1758 = Scalopus aquaticus. 

1825. Condylura macroura Harlan, Fauna Am., p. 39. Holotype: PM 866 

(lost or destroyed: Woodman, 2009), from “the United States, abound 

in New Jersey.” 

FIGURE 9. “La Taupe du Canada,” Talpa canadensis De la Faille, 1769 (=Condylura cristata (Linnaeus, 1758)). Image from De la Faille 

(1778), courtesy of the Joseph F. Cullman 3rd Library of Natural History, Smithsonian Institution Libraries.
1 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1825. Astromycter prasinatus: Harris, East. Star 2(24 or 25):??; May. Type 

locality: “Machias, in the state of Maine.” A summary notice by Benjamin Silliman in the American Journal of Science for June 1825 (Harris, 1825c) is typically cited as the original description of Harris’ green 

mole, but Harris (1825a, 1825b) published more complete descriptions 

under the names Astromycter prasinatus and Condylura prasinata, respectively, in May 1825.

1825. [Condylura] prasinata Harris, Boston J. Philos. Arts 2(6):582; May. 

1832. astromycter prarinatus Rafinesque, Atlantic J. 1:61; summer. Incorrect 

subsequent spelling of Astromycter prasinatus Harris, 1825.

1839. Condytura cristata: Bell, Cyclop. Anat. Phys. 2:994.

1841. Rh[inaster]. cristatus: Wagner, Die Säugthiere Suppl. 2:114.

1841. Rh[inaster]. macrurus Wagner, Die Säugthiere Suppl. 2:115. Incorrect 

subsequent spelling of C. macroura Harlan, 1825. 

1841. Rh[inaster]. macroura: Wagner, Die Säugthiere Suppl. 2:117. 

1841. Condylura macrura Wagner, Die Säugthiere Suppl. 2: pl. 156A. Incorrect subsequent spelling of C. macroura Harlan, 1825. 

1863. Astromydes cristatus: Blyth, Cat. Mamm. Asia. Soc. Mus., p. 87.

1863. Condylura Christata Gilpin, Proc. Trans. Nova Scot. Inst. Nat. Sci. 

1(2):4. Incorrect subsequent spelling of Sorex cristatus Linnaeus, 1758 

(=Condylura cristata).

1913. Condylus cristata: Van Hyning, Science 38(972):243; 15 August. 

1940. Condylura cristata nigra R. W. Smith, Am. Midl. Nat. 24(1):218; 

31 July. Holotype: MVZ 86603, from “Wolfville, Kings County, Nova 

Scotia,” Canada.

1940. Condylura cristata cristata R. W. Smith, Am. Midl. Nat. 24(1):218; 

31 July. First use of current name combination.

Holotype: None known to exist (Wallin, 2001). In describing 

the species, Linnaeus (1758:53) referenced Pehr Kalm, a 

Swedish-Finnish explorer, who visited what is now the eastern United States and probably provided him with a description of the species. 

Type Locality: USA: “Pensylvania”; restricted to “Eastern Pennsylvania” by Jackson (1915:87). If this is the animal described 

by Kalm (1753–1756, 2:310–312; 1770–1771, 1:190–191) 

in his journal for 12 October 1748, the type locality can be 

further restricted to the Schuylkill River valley near Philadelphia. However, Linnaeus (1758) also referenced Kalm for 

Sorex aquaticus Linnaeus, 1758.

Distribution: Canada and USA: from the Atlantic coast of Labrador to east central Manitoba and south to southeastern North 

FIGURE 10. Condylura macroura Harlan, 1825 (=Condylura cristata (Linnaeus, 1758)). Image from Wagner (1855: pl. 156A), courtesy of 

Biodiversity Heritage Library (https://www.biodiversitylibrary.org/), contributed by Ernst Mayer Library, Harvard University, Cambridge.
NUMBER 650 • 1 7

Dakota, central Wisconsin, southern Indiana, northeastern 

Ohio, central West Virginia, and central Virginia (Hall, 1981).

Remarks: Although Hall (1981) recognized C. cristata nigra as 

a distinct form, Petersen and Yates (1980) considered it a 

synonym of C. c. cristata. 

Condylura cristata parva Paradiso, 1959

1959. Condylura cristata parva Paradiso, Proc. Biol. Soc. Wash. 72:103; 

24 July.

Holotype: USNM 293291.

Type Locality: USA: “5 mi. NW. of Stuart, Patrick Co., Virginia.” 

Distribution: USA: central West Virginia and central Virginia 

south to eastern Tennessee and western South Carolina and 

south along the coast to the southeastern corner of Georgia 

(Hall, 1981).

Remarks: Jackson (1915) and Petersen and Yates (1980) noted 

that C. cristata is generally larger in the northern part of 

its geographical distribution and smaller to the south. The 

subspecies C. cristata parva represents the smaller southern 

individuals in what may be a gradual north–south size cline. 

If this proves to be correct, the separation of C. cristata into 

subspecies is probably unwarranted. 

Hartman (1999) stated that C. cristata nigra is a synonym of C. cristata parva, in which case the former name 

would have priority (see Hutterer, 2005:301). However, the 

type localities of the two taxa are >1,600 km apart, and nowhere would their proposed distributions come into contact 

(see Paradiso, 1959:105; Hall, 1981:76, map 44).

Tribe Neurotrichini Hutterer, 2005

2005. Neurotrichini Hutterer, in MSW3, p. 303.

Genus Neurotrichus Günther, 1880

1857. Urotrichus: Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):76, pl. 28 (not 

Urotrichus Temminck, 1841).

1880. Neürotrichus Günther, Proc. Zool. Soc. Lond. 1880(3):441; October.

1880. Nëurotrichus: Günther, Proc. Zool. Soc. Lond. 1880(3): pl. 42; October. Incorrect subsequent spelling of Neürotrichus Gunther, 1880.

1881. Neourotrichus: Rye, Zool. Record 17: index, p. 8; October. Incorrect 

subsequent spelling of Neürotrichus Gunther, 1880.

1881. Neurotrichus: Forbes, Zool. Record 17: Mammalia, p. 14. Justified 

emendation of Neürotrichus Gunther, 1880.

Type Species: Urotrichus gibbsii Baird, 1857, by monotypy.

Remarks: Neurotrichus was revised by True (1896) and Jackson (1915) and reviewed by Hall and Kelson (1959), Hall 

(1981), Carraway and Verts (1991), and Yates (1999). Two 

Plio-Pleistocene species of talpids from Poland that had 

been referred to Neurotrichus were recently recognized as 

belonging to a distinct fossil genus, Rzebikia Sansalone, 

Kotsakis, and Piras, 2016 (Sansalone et al., 2016a).

Neurotrichus gibbsii (Baird, 1857)

FIGURES 11, 12

Remarks: Neurotrichus gibbsii is the type species for the genus

Neurotrichus Günther, 1880.

Neurotrichus gibbsii gibbsii (Baird, 1857)

1857. Urotrichus gibbsii Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):76, pl. 28.

1866. Urotrichus Gibsii Lord, Naturalist in Vancouver Island and British 

Columbia, p. 338. Incorrect subsequent spelling of Urotrichus gibbsii

Baird, 1857.

1880. [Neürotrichus] gibbsii: Günther, Proc. Zool. Soc. Lond. 1880(3):441; 

October.

1880. Nëurotrichus gibbsii: Günther, Proc. Zool. Soc. Lond. 1880(3): pl. 42; 

October.

1891. Neurotrichus gibbsii: Bryant, Zoe 1(12):359; February.

1899. Neurotrichus gibbsi major Merriam, N. Am. Fauna 16:88; 28 October. Holotype: USNM 65321, from “Carberry Ranch, Shasta County, 

Calif. (alt. 4,100 ft., between Mts. Shasta and Lassen).”

1912. Neürotrichus gibbsii gibbsii: Miller, Bull. U.S. Natl. Mus. 79:11; December. First use of current name combination.

1978. Neurotrichus gibbsii destructionensis Yates, Ph.D. dissertation, Texas 

Tech Univ., p. 267. Holotype: USNM 273081 (determined by me from 

reported measurements), from “Destruction Island, Jefferson County, 

Washington.”

Holotype: USNM 662/1843.

Type Locality: USA: “White River, Cascade Mountains, 

W[estern]. T[erritories]”; restricted to “Naches Pass, 4,500 

foot elevation, Pierce County, Washington,” by Dalquest 

and Burgner (1941:12). 

Distribution: Canada and USA: southwestern British Columbia, across west central Washington to coastal western 

Oregon and coastal northwestern California; elevational 

distribution: from near sea level to 2,075 m (Grinnell, 1933; 

Hall, 1981).

Neurotrichus gibbsii hyacinthinus Bangs, 1897

1897. Neürotrichus gibbsi hyacinthinus Bangs, Am. Nat. 31:240; March.

1901. Neurotrichus gibbsii hyacinthinus: Miller and Rehn, Proc. Boston Soc.

Nat. Hist. 30:254; 27 December (first correctly spelled use of current 

name combination).

1905. Neürotrichus gibbsi hyacinthinus: Elliot, Field Col. Mus. Publ. 105,

Zool. Ser. 6:467.

1978. Neurotrichus gibbsii intermedius Yates, Ph.D. dissertation, Texas Tech 

Univ., p. 273. Holotype: USNM 160900 (not 130217 as reported by 

Yates, 1978), from “5 mi. S Aptos, Santa Cruz County, California.”

1978. N[eurotrichus]. g[ibbsii]. intermedus Yates, Ph.D. dissertation, Texas 

Tech Univ., p. 274. Incorrect subsequent spelling of Neurotrichus gibbsii intermedius Yates, 1978.

Holotype: MCZ BANGS-1240.

Type Locality: USA: “Nicasio, Marin Co., Cal[ifornia].” 

Distribution: USA: Narrow humid coastal belt from about central Humboldt County south to northern Monterey County, 
1 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 11. Spencer Baird’s shrews and mole: from top to bottom, USNM 2157, Blarina exilipes Baird, 1857 

(=Sorex parvus Say, 1822 = Cryptotis parvus); USNM 2159, Blarina berlandieri Baird, 1857 (=Cryptotis berlandieri); USNM 1688, Sorex hoyi Baird, 1857; USNM 1699, Sorex platyrhinus De Kay, 1842 (=Sorex cinereus Kerr, 

1792); Urotrichus gibbsii Baird, 1857 (=Neurotrichus gibbsii). Image from Baird (1857: pl. 18).
NUMBER 650 • 1 9

California; elevational distribution: below 245 m (Miller 

and Kellogg, 1955; Carraway and Verts, 1991).

Neurotrichus gibbsii minor Dalquest and Burgner, 1941

1941. Neurotrichus gibbsii minor Dalquest and Burgner, Murrelet 22(1):12; 

30 April.

Holotype: MVZ 94857.

Type Locality: USA: “the University of Washington Campus, 

Seattle, Washington”; King County. 

Distribution: Canada and USA: Lowlands of western Washington and southwesternmost corner of British Columbia 

(Dalquest and Burgner, 1941; Hall, 1981).

Tribe Scalopini Gill, 1875

1875. Scalopes Gill, Synop. Insectiv. Mamm., p. 110; 14 May. Proposed as 

a suprageneric taxon of unidentified rank.

Genus Parascalops True, 1894

1894. Parascalops True, Diagnoses of New North American Mammals, p. 2; 26 April (preprint of Proc. U.S. Natl. Mus. 17:242; 

15 November). 

1902. Perascalops Beddard, Cambr. Nat. Hist., p. 518. Incorrect subsequent 

spelling of Parascalops True, 1894.

Type Species: Scalops breweri Bachman, 1842 (by original 

designation).

Remarks: Parascalops was revised by True (1896) and Jackson (1915) and reviewed by Hall and Kelson (1959), Hallett (1978), and Hall (1981). A Plio-Pleistocene species of 

Parascalops has been reported from Poland (Skoczeń, 1993; 

Sansalone et al., 2016a).

Parascalops breweri (Bachman, 1841)

FIGURE 13

1777. Talpa longicaudata Erxleben, Syst. Reg. Anim., p. 118. Based on Pennant’s (1771:314; pl. 28, fig. 2) “long-tailed mole” from “New York.” 

1777. Talpa Caudata Zimmermann, Spec. Zool. Geog., p. 497. Based on 

Pennant’s (1771:314; pl. 28, fig. 2) “long-tailed mole” from “New 

York.” Part; not T. caudata Linnaeus, 1740. 

1784. Talpa longicauda Boddaert, Elenchus animalium 1:126. Unjustified 

emendation of T. longicaudata Erxleben, 1777.

1819. Condylura longicaudata: Desmarest, J. Phys. Chim. Hist. Nat. Arts

89:232; September.

1825. Talpasor[ex]. longicaudata: Schinz, Thierreich 4:312.

1825. Talpa Europea: Harlan, Fauna Am., p. 43. Part; not T. europaea Linnaeus, 1758.

1825. Talpa americana Harlan, Fauna Am., p. 43. Nomen nudum; a manuscript name for the “black mole” mentioned in an unpublished diary by 

William Bartram and listed in Harlan’s (1825:43) synonymy for Talpa 

europaea: 

That venerable naturalist, the late Mr. William Bartram, was 

accustomed to keep a diary, or “A calendar of natural history,” 

in which he carefully noted his daily observations, almost up to 

the hour of his death; these were never intended for publication, 

but since this work went to press, his relative, Maj. Robert Carr 

(the present proprietor of the Gardens,) has politely favoured 

me with a perusal of them.

I have not seen these notes personally, so I do not know whether the 

name Talpa americana originated with Bartram, but Harlan (1825) was 

the first to publish it.

FIGURE 12. Neurotrichus gibbsii (Baird, 1857) from Günther (1880: pl. 42, fig. A). Image courtesy of Biodiversity Heritage Library (https://

www.biodiversitylibrary.org/), contributed by The Natural History Museum Library, London.
2 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1841. S[calops]. Breweri Bachman, Proc. Boston Soc. Nat. Hist. 1:41. 

Bachman (1842) typically is given incorrectly as the original 

description.

1841. Rh[inaster]. longicaudatus: Wagner, Die Säugthiere Suppl. 2:115. 

1842. Scalops breweri: Bachman, Boston J. Nat. Hist. 4(1):32; January. 

1848. [Scapanus] breweri: Pomel, Arch. Sci. Phys. Nat. 9:247. 

1854. T[alpa]. reposta Le Conte, Proc. Acad. Nat. Sci. Phila. 6:327. 

Holotype: “There is in the Academy a specimen labelled ‘Scalops 

Breweri’ . . . Locality unknown.” The ANSP type is now lost or 

destroyed (Koopman, 1976). 

1854. T[alpa]. Breweri: Le Conte, Proc. Acad. Nat. Sci. Phila. 6:327.

1879. Scapanus americanus: Coues, Am. Nat. 13(3):190; March.

1884. Scapanus breweri: True, U.S. Natl. Mus. Circ. 29:22; 29 November.

1892. Scaphanus breweri: Herrick, Bull. Geol. Nat. Hist. Surv. Minn. 7:55.

1894. Parascalops breweri: True, Proc. U.S. Natl. Mus. 17:242; 26 April. 

First use of current name combination.

Holotype: “None known to exist” (Jackson, 1915:80). 

Type Locality: USA: eastern North America. The type was 

supposed by Bachman to have been taken on the island of 

Martha’s Vineyard, Massachusetts, where the species is not 

known to occur. 

Distribution: Canada and USA: southern Ontario, southern 

Quebec, and central Maine, south along the East Coast to 

eastern Massachusetts, central Connecticut, southeastern 

Pennsylvania, and along the Appalachians to northern South 

Carolina, and north through central Ohio (Hall, 1981).

Remarks: This is the type species of the monotypic genus Parascalops True, 1894. The original description of the species is 

typically cited as an article by Bachman (1842) in the Boston Journal of Natural History, but that work is preceded 

by an earlier description in the Proceedings of the Boston 

Society of Natural History (Bachman, 1841). 

Both Talpa longicaudata Erxleben, 1777 and Talpa 

caudata Zimmermann, 1777 were based on Pennant’s 

(1771) “long-tailed mole” (Figure 13), a species that has 

been equated alternatively with the modern species Scalopus aquaticus (Linnaeus, 1758) by Allen (1902) and 

Condylura cristata (Linnaeus, 1758) by Jackson (1915). 

According to Jackson (1915:89), Pennant (1771) described 

the long-tailed mole “as a mole with a radiated nose and a 

tail two inches long,” and Jackson therefore placed T. longicaudata and T. caudata in synonymy with Condylura 

cristata (Linnaeus, 1758). In fact, Pennant’s (1771:313) description of “nose long; the edges beset with radiated tendrils” refers to his “radiated mole.” His description of the 

long-tailed mole never mentions a radiated nose but does 

state “tail covered with short hair; the length two inches” 

(Pennant, 1771:314). An accompanying illustration figured 

both species (Pennant, 1771: pl. 28, fig. 2), clearly showing 

the radiated nose of the radiated mole and the prominent 

hairy tail of the long-tailed mole, thereby indicating that 

the latter animal was the hairy-tailed mole, Parascalops 

breweri (Figure 13). 

Genus Scalopus É. Geoffroy Saint-Hilaire, 1803

1776. Talva Müller, Voll. Natursyst. Suppl., p. 35. Incorrect subsequent spelling of Talpa Linnaeus, 1758.

1789. Talpa: Müller, Voll. Natursyst. Suppl., p. 35. 

1800. Scalops Cuvier, Leçons d’anatomie comparée, Vol. 1: tab. 1: Classification des mammifères. Nomen nudum.

1803. Scalopus É. Geoffroy Saint-Hilaire, Cat. Mamm. Mus. Natl. Hist.

Nat., p. 77. 

1811. Scalops Illiger, Prodr. Syst. Mamm. Avium, p. 126. Type species: Sorex 

aquaticus Linnaeus, 1758, by original designation.

1827. Talpasorex Lesson, Manuel de mammalogie, p. 124. Type species: 

Scalops pennsylvanica Harlan, 1825 (=Sorex aquaticus Linnaeus, 

1758) (not Talpasorex Schinz, 1821).

1911. Scalpos Brooks, Rep. W. Va. Board Agric., p. 28. Incorrect subsequent 

spelling of Scalops Illiger, 1811.

1941. Hesperoscalops Hibbard, Am. Midl. Nat. 26(2):337. Type species: Hesperoscalops rexroadi Hibbard, 1941 from the Pliocene of 

Kansas.

Type Species: Scalopus virginianus É. Geoffroy Saint-Hilaire, 

1803 (=Sorex aquaticus Linnaeus, 1758), designated by 

Palmer (1904:621).

Remarks: The genus was revised by True (1896), Jackson 

(1915), and Yates and Schmidly (1977) and reviewed by 

Hall and Kelson (1959), Yates and Schmidly (1978), and 

Hall (1981). The genus contains a single species with 16 

subspecies.

Scalopus aquaticus (Linnaeus, 1758)

Remarks: This is the type species for the genus Scalopus

É. Geoffroy Saint-Hilaire, 1803.

Scalopus aquaticus aereus (Bangs, 1896) 

1896. Scalops texanus aereus Bangs, Proc. Biol. Soc. Wash. 10:138; 

28 December.

1899. Scalops machrinus intermedius Elliot, Field Col. Mus. Publ. 37, Zool. 

Ser. 1(14):280; 9 May. Holotype: FMNH 6829 (not FMNH 6832, as 

suggested by Jackson, 1915:49), from “Alva, Oklahoma Territory”; 

Woods County, Oklahoma, USA. There was some question regarding 

which of two FMNH specimens represents the holotype of Scalops machrinus intermedius. Elliot (1899a:280) did not designate the holotype 

by number in his original description but stated that it was a male collected 23 February 1899 by Thaddeus Surber at Alva. It appears that 

Elliot incorrectly reported the date of collection (A. Ferguson, FMNH, 

email communication, 24 April 2017). 

1901. [Scalops] aereus: Elliot, Field Col. Mus. Publ. 45, Zool. Ser. 2(1):390.

1901. Scalops aquaticus aereus: Miller and Rehn, Proc. Boston Soc. Nat.

Hist. 30:250; 27 December.

1901. [Scalops aquaticus] intermedius: Elliot, Field Col. Mus. Publ. 45, Zool.

Ser. 2(1):390.

1905. Scalopus aquaticus intermedius: Bailey, N. Am. Fauna 25:207; 24 

October.

1905. Scalopus aereus: Elliot, Field Col. Mus. Publ. 105, Zool. Ser. 6:471.
NUMBER 650 • 2 1

1914. Scalopus aquaticus pulcher Jackson, Proc. Biol. Soc. Wash. 27:20; 2 

February. Holotype: USNM 170698, from “Delight, Arkansas”; Pike 

County. 

1912. Scalopus aquaticus aereus: Miller, Bull. U.S. Natl. Mus. 79:8; 31 December. First use of current name combination.

Holotype: MCZ BANGS-5475.

Type Locality: USA: “Stilwell, I[ndian]. T[erritory].”; Adair 

County, Oklahoma, USA. 

Distribution: USA: southeastern corner of Colorado to south 

central Kansas to northeastern Arkansas and south to 

southern Louisiana and portions of eastern and north central Texas, including the panhandle (Hall, 1981).

Scalopus aquaticus alleni R. H. Baker, 1951

1951. Scalopus aquaticus alleni R. H. Baker, Univ. Kansas Publ. Mus. Nat.

Hist. 5(2):22; 28 February.

Holotype: AMNH 7189/5788.

Type Locality: USA: “Rockport, Aransas County, Texas.” 

Distribution: USA: portions of southeastern and south central 

Texas (Hall, 1981).

FIGURE 13. Thomas Pennant’s (1771) “radiated mole” (=Sorex cristatus Linnaeus, 1758 = Condylura cristata) (top) and “longtailed mole” (=Scalops breweri Bachman, 1841 = Parascalops breweri) (bottom). 
2 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Scalopus aquaticus anastasae (Bangs, 1898) 

1898. Scalops anastasae Bangs, Proc. Boston Soc. Nat. Hist. 28(7):212; 

March.

1901. [Scalops] anastasiae Elliot, Field Col. Mus. Publ. 45, Zool. Ser. 

2(1):391; 6 March. Incorrect subsequent spelling of Scalops anastasae

Bangs, 1898.

1912. Scalopus anastasae: Cory, Field Mus. Nat. Hist. publ. 153, Zool. Ser. 

11:438; June.

1915. Scalopus aquaticus anastasae: Jackson, N. Am. Fauna 38:39; 30 September. First use of current name combination.

Holotype: MCZ 7192.

Type Locality: USA: “Point Romo, Anastasia Island, Florida”; 

St. Johns County.

Distribution: USA: known only from Anastasia Island.

Scalopus aquaticus aquaticus (Linnaeus, 1758) 

FIGURES 14, 15

1734. Talpa, Virginianus, niger Seba, Thesaurus, p. 51, tab. XXXII: fig. 3 

(pre-Linnaean work).

1758. [Sorex] aquaticus Linnaeus, Systema Naturae, 10th ed., 1:53.

1776. Talva virginea Müller, Voll. Natursyst. Suppl., p. 35. Based on Seba’s 

(1734) Talpa, Virginianus, niger from “Virginien.”

1777. Talpa europea flavescens Erxleben, Syst. Reg. Anim., p. 118; based 

on Pennant’s (1771) “yellow mole” from “New York” (see Pennant, 

1771:314); see also Allen (1902:16).

1777. Talpa Flava Zimmermann, Spec. Zool. Geog., p. 496; based on Pennant’s (1771:312) “yellow mole” from “New York” (see Pennant, 

1771: 314); see also Allen (1902:16) and Jackson (1915:89).

1777. Talpa fusca Zimmermann, Spec. Zool. Geog., p. 497; based on Pennant’s (1771:314) “brown mole” from “New York” (see Pennant, 

1771:314); see also Allen (1902:16).

1788. [Talpa europaea] flava: Gmelin, Systema Naturae, 13th ed., p. 110.

1789. Talpa virginea: Müller, Voll. Natursyst. Suppl., p. 35. 

1800. Talpa purpurascens Shaw, Gen. Zool. 1(2):521; based on Talpa, Virginianus, niger Seba, 1734 (not Talpa purpurascens Schinz, 1821 = 

Sorex cristatus Linnaeus, 1758).

1803. Scalopus virginianus É. Geoffroy Saint-Hilaire, Cat. Mamm. Mus.

Natl. Hist. Nat., p. 78. Based on Seba’s (1734) Talpa, Virginianus, niger

(Figure 14) and Sorex aquaticus Linnaeus, 1758.

1814. Talpa cupreata Rafinesque, Précis découv., p. 14. Type locality: 

“Amer[ica]. septentrionale”; restricted to “the Atlantic States” by 

Rafinesque (1832:61).

1820. Scalops canadensis Desmarest, Mammalogie 1:155; based on Sorex 

aquaticus Linnaeus, 1758.

1825. Scalops pennsylvanica Harlan, Fauna Am., p. 33. Holotype in Harlan’s 

personal collection (Harlan, 1825:33), now lost or destroyed; presumably from southeastern Pennsylvania (see Jackson, 1915). 

1825. Talpa europea: Harlan, Fauna Am., p. 43 (not Talpa europea Linnaeus, 1758).

1827. Talpasorex pensylvanica: Lesson, Manuel de mammalogie, p. 124. Incorrect subsequent spelling of S. pennsylvanica Harlan, 1825.

1829. Sc[alops]. aquaticus: J. B. Fischer, Synopsis mammalium, p. 249.

1838. Talpa virginiana: Blainville, Ann. Franç. Anat. Phys. 2:219.

1838. Talpa sorex pensylvancius: Blainville, Ann. Franç. Anat. Phys. 2:219. 

Incorrect subsequent spelling of Talpasorex pensylvanica Lesson, 1827. 

1854. T[alpa]. aquatica: Le Conte, Proc. Acad. Nat. Sci. Phila. 6:327.

1884. Scalops aquaticus aquaticus: True, U.S. Natl. Mus. Circ. 29:22; 29 

November.

1905. Scalopus aquaticus: Oberholser, Notes on the Mammals and Summer Birds of Western North Carolina (Biltmore, N.C.: Biltmore Forest 

School), p. 8; 30 September.

1911. Scalpos aquaticus: Brooks, Rep. W. Va. Board Agric., p. 28.

1912. Scalopus aquaticus aquaticus: Miller, Bull. U.S. Natl. Mus. 79:7. First 

use of current name combination.

1978. Scalopus aquaticus davisi Yates, Ph.D. dissertation, Texas Tech Univ., 

p. 229. Holotype: CM 18444, from “Yarmouth, Barnstable Co., 

Massachusetts.”

Holotype: None known to exist (Wallin, 2001). Linnaeus 

(1758:53) references Pehr Kalm, a Swedish-Finnish explorer, 

who probably provided him with a description of the species.

Type Locality: “America septentrionali” (North America); restricted to “Philadelphia, Pennsylvania,” USA, by Jackson 

(1915:33). This species (or Sorex cristatus Linnaeus, 1758) 

may be the animal described by Kalm (1753–1756, 2:310–

312; 1770–1771, 1:190–191) in his published journal for 

12 October 1748.

Distribution: USA: eastern states from eastern and southern 

Massachusetts, southeastern New York, southeastern Pennsylvania, and northeastern West Virginia, south through 

Virginia, and in Appalachian Mountains south through 

western North Carolina and eastern Tennessee (Hall, 1981); 

elevational distribution from near sea level to 1,220 m 

(USNM specimens; Oberholser, 1905).

Scalopus aquaticus australis (Chapman, 1893) 

1893. Scalops aquaticus australis Chapman, Bull. Am. Mus. Nat. Hist. 

5(21):339; 22 December.

1905. Scalopus aquaticus australis: Elliot, Field Col. Mus. Publ. 105, Zool.

Ser. 6:470. First use of current name combination.

Holotype: AMNH 3916/2990.

Type Locality: USA: “Gainesville, Florida”; Alachua County. 

Distribution: USA: southeastern Georgia and eastern portion of peninsular Florida south to Hypoluxo, Palm Beach 

County (Hall, 1981).

Scalopus aquaticus bassi A. H. Howell, 1939 

1939. Scalopus aquaticus bassi A. H. Howell, J. Mamm. 20(3):363; 14 

August.

Holotype: FMNH 43968.

Type Locality: USA: “Engelwood, Sarasota County, Florida.” 

Distribution: USA: known only from the type locality.

Scalopus aquaticus caryi Jackson, 1914

1914. Scalopus aquaticus caryi Jackson, Proc. Biol. Soc. Wash. 27:20; 2 February.
NUMBER 650 • 2 3

FIGURE 15. Sorex aquaticus Linnaeus, 1758 (=Scalopus aquaticus). Image from Wagner (1855: pl. 158), courtesy of Biodiversity 

Heritage Library (https://www.biodiversitylibrary.org/), contributed by the Ernst Mayer Library, Museum of Comparative Zoology, 

Harvard University, Cambridge, Massachusetts.

FIGURE 14. Ventral view of the “purple mole,” Seba’s (1734: pl. 32, fig. 3) Talpa, Virginianus, niger, supinus (=Sorex aquaticus

Linnaeus, 1758 = Scalopus aquaticus). Image courtesy of Biodiversity Heritage Library (https://www.biodiversitylibrary.org/), contributed by Smithsonian Institution Libraries, Washington, D.C.
2 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1978. Scalopus aquaticus nebrascensis Yates, Ph.D. dissertation, Texas Tech 

Univ., p. 239. Holotype: KU 50129, from “5 mi. N, 2 mi. W Parks, 

Dundy Co., Nebraska.”

Holotype: USNM 116799.

Type Locality: USA: “Neligh, Nebraska”; Antelope County. 

Distribution: USA: Arid and semiarid Great Plains region from 

southeastern Wyoming, central and western Nebraska, northeastern Colorado, and northwestern Kansas (Hall, 1981).

Scalopus aquaticus cryptus W. B. Davis, 1942

1942. Scalopus aquaticus nanus W. B. Davis, Am. Midl. Nat. 27(2):383; 

March. Holotype: TCWC 1785, from “13 miles east of Centerville, 

Leon County, Texas.”

1942. Scalopus aquaticus cryptus W. B. Davis, Am. Midl. Nat. 27(2):384; 

March.

Holotype: TCWC 1454.

Type Locality: USA: “College Station, Brazos County, Texas.”

Distribution: USA: portions of eastern Texas (Hall, 1981).

Remarks: Yates and Schmidly (1977) considered the names 

Scalopus aquaticus nanus and S. a. cryptus to represent a 

single entity, which, as first revisers, they called S. a. cryptus. 

Citing that work, Hall (1981) also treated the two names as 

representing a single population, but under the name S. a.

nanus, possibly because S. a. nanus appears before S. a.

cryptus in Davis (1942). However, because both names were 

published at the same time, neither has strict priority, and 

the name chosen by the first reviser (ICZN, 1999:24.2), in 

this case by Yates and Schmidly (1977), becomes the available name.

Scalopus aquaticus howelli Jackson, 1914

1914. Scalopus aquaticus howelli Jackson, Proc. Biol. Soc. Wash. 27:19; 2 

February.

Holotype: USNM 177931.

Type Locality: USA: “Autaugaville, Alabama”; Autauga 

County.

Distribution: USA: North Carolina (except in Appalachian 

Mountains), South Carolina, northern and western Georgia, the panhandle of Florida, Alabama, Mississippi, and 

Louisiana east of the Mississippi River (Hall, 1981).

Scalopus aquaticus inflatus Jackson, 1914

1914. Scalopus inflatus Jackson, Proc. Biol. Soc. Wash. 27:21; 2 February.

1977. Scalopus aquaticus inflatus: Yates and Schmidly, Occ. Pap. Mus. Texas 

Tech Univ. 45:28; 3 June. First use of current name combination.

Holotype: USNM 52709.

Type Locality: Mexico: “state of Tamaulipas, Mexico (45 miles 

from Brownsville, Texas).”

Distribution: USA and Mexico: southeastern tip of Texas to 

the type locality in Tamaulipas (Hall, 1981).

Scalopus aquaticus machrinoides Jackson, 1914

1914. Scalopus aquaticus machrinoides Jackson, Proc. Biol. Soc. Wash. 

27:19; 2 February.

Holotype: USNM 169717.

Type Locality: USA: “Manhattan, Kansas”; Riley County.

Distribution: USA: west of the Mississippi River, from central 

Minnesota, the southeastern corner of South Dakota, and 

eastern Nebraska, south through eastern Kansas to extreme

northern Arkansas (Hall, 1981).

Scalopus aquaticus machrinus (Rafinesque, 1832) 

1832. Talpa machrina Rafinesque, Atlantic J. 1:61.

1832. Talpa sericea Rafinesque, Atlantic J. 1:62. Type locality: “woods near 

Nicholasville and Harrodsburg,” Jessamine and Mercer Counties, Kentucky, respectively. True (1896:20) noted that Rafinesque’s description 

was of a young individual, and Jackson (1915:43) stated that this “species is clearly the young of . . . Talpa machrina.”

1842. Scalops argentatus Audubon and Bachman, J. Acad. Nat. Sci. Phila.

8(2):292. Type locality: “Michigan”; restricted to “southern Michigan” 

by Jackson (1915:42).

1854. Talpa Pennantii Le Conte, Proc. Acad. Nat. Sci. Phila. 6:327. Holotype: “A specimen in the Academy” [ANSP], locality unknown.

1877. Scalops aquaticus argentatus: Coues, Bull. U.S. Geol. Geog. Surv.

Terr. 3(3):633.

1896. Scalops aquaticus machrinus: True, Proc. U.S. Natl. Mus. 19:20; 21 

December.

1905. Scalopus aquaticus machrinus: Elliot, Field Col. Mus. Publ. 105, Zool.

Ser. 6:470. First use of current name combination.

Holotype: “None known to exist” (Jackson, 1915:43).

Type Locality: USA: “woods, gardens and fields, near Lexington,” Fayette County, Kentucky. 

Distribution: Canada and USA: east of the Mississippi River 

and west of the Appalachian Mountains from western Wisconsin, northern Illinois, southern Michigan, southwestern 

Ontario (Essex County), and northern Ohio, south to central and western Tennessee (Hall, 1981).

Scalopus aquaticus montanus R. H. Baker, 1951

1951. Scalopus montanus R. H. Baker, Univ. Kansas Publ. Mus. Nat. Hist. 

5(2):19; 28 February.

1977. Scalopus aquaticus montanus: Yates and Schmidly, Occ. Pap. Mus.

Texas Tech Univ. 45:29; 3 June. First use of current name combination.

Holotype: KU 35668.

Type Locality: Mexico: “Club Sierra del Carmen, 2 mi. N and 

6 mi. W Piedra Blanca, Coahuila.” 

Distribution: Mexico: isolated population known only from 

the type locality in the Sierra del Carmen of northern 

Coahuila.

Scalopus aquaticus parvus (Rhoads, 1894) 

1894. Scalops parvus Rhoads, Proc. Acad. Nat. Sci. Phila. 46:157. 
NUMBER 650 • 2 5

1901. [Scalops aquaticus] parvus: Elliot, Field Col. Mus. Publ. 45, Zool. Ser. 

2(1):390; 6 March.

1915. Scalopus aquaticus parvus: Jackson, N. Am. Fauna 38:41; 30 September. First use of current name combination. 

Holotype: ANSP 8468 (Koopman, 1976).

Type Locality: USA: “Tarpon Springs, Fla.”; Pinellas County. 

Distribution: USA: central western coast of peninsular Florida 

(Hall, 1981).

Scalopus aquaticus porteri Schwartz, 1952

1952. Scalopus aquaticus porteri Schwartz, J. Mamm. 33(3):381; 19 August.

Holotype: UMMZ 97718.

Type Locality: USA: “Uleta, Dade County, Florida.” 

Distribution: USA: eastern Dade County, Florida (Hall, 1981).

Scalopus aquaticus texanus (J. A. Allen, 1891)

1891. Scalops argentatus texanus J. A. Allen, Bull. Am. Mus. Nat. Hist. 

3(2):221; 29 April.

1893. Scalops texanus: J. A. Allen, Bull. Am. Mus. Nat. Hist. 5:200; 

18 August.

1896. Scalops aquaticus texanus: True, Proc. U.S. Natl. Mus. 19:21;

21 December.

1901. [Scalops aquaticus] texensis Elliot, Field Col. Mus. Publ. 45, Zool.

Ser. 2(1):390; 6 March. Incorrect subsequent spelling of S. aquaticus 

texanus J. A. Allen, 1891.

1905. Scalopus aquaticus texanus: Bailey, N. Am. Fauna 25:206; 24 October. 

First use of current name combination.

1905. Scalopus aquaticus texensis: Elliot, Field Col. Mus. Publ. 105, Zool.

Ser. 6:471. Incorrect subsequent spelling of S. aquaticus texanus J. A. 

Allen, 1891.

Holotype: AMNH 3488/2740.

Type Locality: USA: “Presidio Co., Texas.” Baker (1951:21) 

summarized the confusion regarding the location of the type 

locality in Presidio County. The holotype was collected by 

Wm. Lloyd in September 1887. In that same year, “Presidio 

County was reduced to its present size by the creation of 

Brewster and Jeff Davis counties. Lloyd might have taken 

the mole in any one of these three Trans-Pecos counties.” 

Distribution: USA: isolated population in the Big Bend region 

of Texas.

Genus Scapanus Pomel, 1848

1848. Scapanus Pomel, Arch. Sci. Phys. Nat. 9:247; November. 

1892. Scaphanus Herrick, Bull. Geol. Nat. Hist. Surv. Minn. 7:55. Incorrect 

subsequent spelling of Scapanus Pomel, 1848.

1902. Scapasius Beddard, Cambr. Nat. Hist., p. 518. Incorrect subsequent 

spelling of Scapanus Pomel, 1848.

Type Species: Scalops townsendii Bachman, 1839, designated 

by Elliot (1901:391).

Remarks: The genus was revised by True (1896) and Jackson 

(1915) and reviewed by Hall and Kelson (1959) and Hall 

(1981). Four modern species of Scapanus are recognized. 

Scapanus anthonyi and S. latimanus were revised by Palmer 

(1937) and Yates and Salazar-Bravo (2005) and reviewed 

by Verts and Carraway (2001). Scapanus orarius was reviewed by Hartman and Yates (1985), and S. townsendii

was reviewed by Carraway et al. (1993). Fossil species were 

reviewed by Hutchinson (1987).

Scapanus anthonyi J. A. Allen, 1893

1893. Scapanus anthonyi J. A. Allen, Bull. Am. Mus. Nat. Hist. 5:200; 18 

August.

1906. Scapanus californicus anthonyi: Stephens, California Mammals, p. 257.

1937. Scapanus latimanus anthonyi: F. G. Palmer, J. Mamm. 18(3):312; 

14 August. 

Holotype: AMNH 6313/4947.

Type Locality: Mexico: “San Pedro Martir Mountains (alt.

7000 ft.)”; Baja California. 

Distribution: Mexico: known only from Vallecitos south to La 

Grulla in the Sierra San Pedro Martir, Baja California (Yates 

and Salazar-Bravo, 2005); elevational distribution: 2130–

2285 m (Miller and Kellogg, 1955; Hall, 1981).

Remarks: Scapanus anthonyi previously was treated as a subspecies of S. latimanus (Palmer, 1937; Verts and Carraway, 

2001), but Yates and Salazar-Bravo (2005) determined it to 

be a separate species. It is noted for its small size and the 

consistent presence of only three premolars, rather than the 

usual four premolars typical of members of the genus (Yates 

and Salazar-Bravo, 2005).

Scapanus latimanus (Bachman, 1841)

Scapanus latimanus insularis F. G. Palmer, 1937

1937. Scapanus latimanus insularis F. G. Palmer, J. Mamm. 18(3):297; 14 

August.

Holotype: MVZ 68993.

Type Locality: USA: “Angel Island, San Francisco Bay, Marin 

County, California.”

Distribution: USA: known only from Angel Island; elevational 

distribution from near sea level to about 230 m (Miller and 

Kellogg, 1955; Hall, 1981).

Scapanus latimanus latimanus (Bachman, 1841) 

1841. S[calops]. latimanus Bachman, Proc. Boston Soc. Nat. Hist. 1:41.

1842. Scalops latimanus: Bachman, Boston J. Nat. Hist. 4(1):34; January 

(often incorrectly cited as the original description).

1856. Scalops Californicus Ayres, Proc. Calif. Acad. Sci. 1(1):54. Neotype: 

USNM 3111, from “San Francisco, California,” selected by Lyon and 

Osgood (1909:233–234), who stated, “this skeleton is one of Dr. Ayres’s 

original specimens, and probably the only one of them now in existence. It seems well to treat it as a type, although it was not so indicated 
2 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

by the original describer.” Jackson (1915:66–67) objected to this designation, stating, “there seems to be no good reason for designating this 

specimen as the type of californicus; . . . if Ayres set aside any specimen 

as the type, it was probably in the Museum of the California Academy of Sciences since preceding his original description the donation of 

five moles to the Academy was acknowledged,” and it is clearly stated 

that Ayres’s description was “in connection” with those five specimens 

(Ayres, 1856). In fact, Jackson (1915) was objecting to the treatment of 

USNM 3111 as though it were part of the original type series, which 

it was not. As suggested by Lyon and Osgood (1909), however, none 

of the five specimens donated to the California Academy of Sciences 

now exists. Nearly the entire collection of that institution was destroyed 

by fire following the 1906 San Francisco earthquake (Maureen Flannery, California Academy of Sciences, email communication, 31 July 

2017). Three USNM specimens (USNM 1288, 2673, 3111) of Scalops 

californicus were either collected by Ayres or obtained from him, the 

earliest of which was collected in September 1855, well after he read 

his description of Scalops californicus before the California Academy of 

Sciences on 21 May 1855 (Ayres, 1856; Jackson, 1915). Despite clearly 

not belonging to the original type series, the three USNM specimens 

were identified by Ayres as belonging to his species, and it is appropriate 

for one of them to have been designated the neotype.

1857. Scalops (Scapanus) townsendi: Baird, Rep. Expl. Surv. Railr. Pacif. 

8(1):65. Part; not Scalops townsendii Bachman, 1839.

1863. Scapanus townsendi: Peters, Monatsb. K. Preuss. Akad. Wiss. Berlin

1864:656. Part; not Scalops townsendii Bachman, 1839.

1887. Scapanus Townsendii: Townsend, Proc. U.S. Natl. Mus. 10:182 (not 

Scalops townsendii Bachman, 1839).

1894. Scapanus dilatus True, Diagnoses of New North American Mammals, 

p. 2; 26 April (preprint of Proc. U.S. Natl. Mus. 17:242; 15 November). 

Holotype: USNM 186628, from “Fort Klamath, Oreg[on].”; Klamath 

County, Oregon, USA.

1896. Scapanus californicus: True, Proc. U.S. Natl. Mus. 19:52; 21 December. Part. 

1897. Scapanus alpinus Merriam, Proc. Biol. Soc. Wash. 11:102; 26 April. 

Holotype: USNM 79967, from “Crater Lake, Mt. Mazama, Oregon. 

Altitude about 2130 meters [7000 ft.]”; Klamath County.

1897. Scapanus truei Merriam, Proc. Biol. Soc. Wash. 11:102; 26 April. 

Holotype: USNM 79290, from “Lake City, Modoc Co., Calif.”

1905. Scapanus truii: Elliot, Field Col. Mus. Publ. 105, Zool. Ser. 6:469. 

Incorrect subsequent spelling of Scapanus truei Merriam, 1897.

1906. Scapanus californicus truei: Stephens, California Mammals, p. 259.

1907. Scapanus latimanus: Osgood, Proc. Biol. Soc. Wash. 20:52; 18 April.

1912. Scapanus latimanus latimanus: Grinnell and Swarth, Univ. Calif. Publ.

Zool. 10(3):131; 13 April. First use of current name combination.

1913. Scapanus latimanus truei: Grinnell, Proc. Calif. Acad. Sci. 3:269;

28 August.

1915. Scapanus latimanus dilatus: Jackson, N. Am. Fauna 38:73; 

30 September. 

1915. Scapanus latimanus alpinus: Jackson, N. Am. Fauna 38:75; 

30 September.

1937. Scapanus latimanus caurinus F. G. Palmer, J. Mamm. 18(3):290; 

14 August. Holotype: MVZ 25388, from “Laytonville, Mendocino 

County, California,” USA.

Lectotype: ZMB 712, designated by Peters (1863:656) from two 

specimens examined by Bachman in the Zoological Museum 

of Humboldt University Berlin (Turni et al., 2007).

Type Locality: USA: Santa Clara, Santa Clara County, California. Peters (1863:656) publicized the fact that the holotype 

was collected by Ferdinand Deppe at Santa Clara in October 

1834, and Osgood (1907:52) confirmed that the locality is in 

California (see also Turni et al., 2007).

Distribution: USA: from south central Oregon through northern 

California, then south in the west along the coastal region of 

California to Santa Barbara; it occurs east of the Sacramento

Valley to the western edge of Nevada and south to Holbrook, 

Douglas County, Nevada (Hall, 1981; Yates and Salazar-Bravo, 

2005); elevational distribution: from near sea level to 2,590 m 

(Grinnell, 1933; Miller and Kellogg, 1955; Hall, 1981). 

Remarks: Scapanus latimanus caurinus and S. l. dilatus previously were considered distinct subspecies (Verts and Carraway, 2001) but were synonymized with S. l. latimanus by 

Yates and Salazar (2005).

Scapanus latimanus minusculus Bangs, 1899

1899. Scapanus californicus minusculus Bangs, Proc. New Engl. Zool. Club 

1:70; 31 July.

1912. Scapanus californicus (=latimanus) minusculus: Grinnell and Swarth, 

Univ. Calif. Publ. Zool. 10(3):133; 13 April.

1912. Scapanus latimanus minusculus: Miller, Bull. U.S. Natl. Mus. 79:10; 

31 December. First use of current name combination.

1913. Scapanus latimanus latimanus: Grinnell, Proc. Calif. Acad. Sci., ser. 4, 

3:269. Part; not Scalops latimanus Bachman, 1841 (=Scapanus latimanus latimanus).

1914. Scapanus latimanus sericatus Jackson, Proc. Biol. Soc. Wash. 27:55; 

20 March. Holotype: USNM 109548, from “Yosemite, Mariposa 

County, California,” USA.

1916. Scapanus latimanus campi Grinnell and Storer, Univ. Calif. Publ.

Zool. 17(1):1; 23 August. Holotype: MVZ 21520, from “Snelling, 250 

feet altitude, Merced County, California,” USA.

1918. Scapanus latimanus monoensis Grinnell, Univ. Calif. Publ. Zool. 

17(14):423; 25 April. Holotype: MVZ 25834, from “Taylor Ranch, 

two miles south of Benton Station, Mono County, California,” USA.

Holotype: MCZ BANGS 9189.

Type Locality: USA: “Fyffe, Eldorado Co., California.” 

Distribution: USA: east central California, from El Dorado 

County to Fresno County (Yates and Salazar-Bravo, 2005); 

elevational distribution: 60–2,900 m (Grinnell, 1933; Miller 

and Kellogg, 1955; Hall, 1981).

Remarks: Scapanus latimanus campi, S. l. monoensis, and S. l.

sericatus previously were considered distinct subspecies 

(Verts and Carraway, 2001) but were synonymized with S. l.

minusculus by Yates and Salazar-Bravo (2005).

Scapanus latimanus occultus Grinnell and Swarth, 1912

1912. Scapanus latimanus occultus Grinnell and Swarth, Univ. Calif. Publ.

Zool. 10(3):131; 13 April.
NUMBER 650 • 2 7

1914. Scapanus latimanus grinnelli Jackson, Proc. Biol. Soc. Wash. 27:56; 20 

March. Holotype: MVZ 17785, from “Independence (altitude 3,900 

feet), Inyo County, California,” USA. This was the site of the former 

Fort Independence, 3 km north of the town of Independence (Miller 

and Kellogg, 1955).

Holotype: MVZ 2369.

Type Locality: USA: “Santa Ana Cañon at 400 feet altitude, 

Orange County, California.” 

Distribution: Mexico and USA: from the Yosemite Valley, Mariposa County, California, south to Santa Barbara and then 

southeast to Laguna Hanson, Baja California (Yates and 

Salazar-Bravo, 2005); elevational distribution from near sea 

level to 2,990 m (Grinnell, 1933; Miller and Kellogg, 1955; 

Hall, 1981).

Remarks: Scapanus latimanus grinnelli previously was considered a distinct subspecies (Verts and Carraway, 2001) but 

was synonymized with S. l. occultus by Yates and SalazarBravo (2005).

Scapanus latimanus parvus F. G. Palmer, 1937

1937. Scapanus latimanus parvus F. G. Palmer, J. Mamm. 18(3):300; 

14 August.

Holotype: MVZ 30343.

Type Locality: USA: “Alameda, Alameda County, California.” 

Distribution: USA: known only from Alameda Island, California (Miller and Kellogg, 1955; Hall, 1981).

Scapanus orarius True, 1896

Scapanus orarius orarius True, 1896

1896. Scapanus orarius True, Proc. U.S. Natl. Mus. 19:52; 21 December.

1915. Scapanus orarius orarius: Jackson, N. Am. Fauna 38:61; 30 September. First use of current name combination.

Holotype: USNM 1381/37434. True (1896:52) incorrectly reported the holotype as USNM 381.

Type Locality: USA: “Shoalwater Bay, Washington” [=Willapa 

Bay], Pacific County.

Distribution: USA: humid Pacific coastal region from northern 

Washington south to northern California and eastward in 

Oregon to the central northern part of the state (Hartman 

and Yates, 1985).

Scapanus orarius schefferi Jackson, 1915

1915. Scapanus orarius schefferi Jackson, N. Am. Fauna 38:63; 30 September.

1944. Scapanus orarius yakimensis Dalquest and Scheffer, Murrelet 25(2):27; 

19 September. Holotype: MVZ 96354, from “3⁄4 mi. N Union Gap, 

Yakima County, Washington.”

Holotype: USNM 204997.

Type Locality: USA: “Walla Walla, Walla Walla County, 

Washington.” 

Distribution: Canada and USA: southwestern corner of British Columbia southeastward through central and western 

Washington and northwestern Oregon to west central Idaho 

(Hartman and Yates, 1985).

Scapanus townsendii (Bachman, 1839)

Remarks: This is the type species of the genus Scapanus Pomel, 

1848.

Scapanus townsendii olympicus M. L. Johnson and Yates, 

1980

1980. Scalops townsendii olympicus M. L. Johnson and Yates, Occ. Pap.

Mus. Texas Tech Univ. 63:1; 11 January.

Holotype: PSMNH 22735.

Type Locality: USA: “Hurricane Ridge, Olympic National 

Park, Clallam Co., Washington, elevation approximately 

1615 meters.”

Distribution: USA: known only from the Olympic Mountains, 

Clallam County, Washington (Johnson and Yates, 1980). 

Scapanus townsendii townsendii (Bachman, 1839) 

1829. Scalops canadensis Richardson, Fauna Bor.-Am. 1:9 (name preoccupied by Scalops canadensis Desmarest, 1820 and Scalops canadensis Harlan, 1825, both = Sorex aquaticus Linnaeus, 1758 = Scalopus 

aquaticus). Type locality: “the banks of the Columbia and the adjoining 

coasts of the Pacific.”

1839. Scalops Townsendii Bachman, J. Acad. Nat. Sci. Phila. 8(1):58.

1842. Scalops Townsendi Bachman, Boston J. Nat. Hist. 4(1):31; January. 

Incorrect subsequent spelling of Scalops townsendii Bachman, 1839 

(=Scapanus townsendii).

1848. Scapanus Towsendii Pomel, Arch. Sci. Phys. Nat. 9:247 Incorrect 

subsequent spelling of Scalops townsendii Bachman, 1839 (=Scapanus 

townsendii).

1854. Scalops metallescens Cassin, Proc. Acad. Nat. Sci. Phila. 6:242 (nomen 

nudum). This appears to be a preliminary working name for the specimen that became the holotype of Scalops aeneus Cassin, 1854. 

1854. Scalops aeneus Cassin, Proc. Acad. Nat. Sci. Phila. 6:299. Holotype: 

USNM 3725, from “Oregon.” This is one of two specimens originally 

reported as Scalops townsendii by Peale (1849:30), “one taken on the 

Columbia, the other on the Willamette River.” See also True (1896:63–

64) and Jackson (1915:59–60).

1854. [Talpa] aenea: Le Conte, Proc. Acad. Nat. Sci. Phila. 6:327.

1854. [Talpa] Townsendii: Le Conte, Proc. Acad. Nat. Sci. Phila. 6:327.

1854. [Talpa] taeniata Le Conte, Proc. Acad. Nat. Sci. Phila. 6:327. Holotype: ANSP 449, from “Banks of the Columbia river”; restricted by 

True (1898:63) to “Fort Vancouver,” Vancouver, Clark County, Washington. The holotype is also one of the two cotypes of Scalops townsendii Bachman, 1839 (Jackson, 1915:58–60).

1857. Scalops (Scapanus) townsendii: Baird, Rep. Expl. Surv. Railr. Pacif. 

8(1):65.

1980. Scalops townsendii townsendii: M. L. Johnson and Yates, Occ. Pap.

Mus. Texas Tech Univ. 63:1; 11 January. First use of current name 

combination.
2 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Lectotype: ANSP 450, designated by True (1896:63) from the 

two specimens upon which Bachman based the species, 

one collected by Thomas Nuttall (ANSP 450), the other by 

John Kirk Townsend (ANSP 449). According to Koopman 

(1976:3), the type is now lost.

Type Locality: USA: “Banks of the Columbia river”; restricted 

by True (1896:63) to “Fort Vancouver,” Vancouver, Clark 

County, Washington (see also Jackson, 1915:58–60). Bachman (1839) did not initially provide a locality for Nuttall’s specimen, but he later wrote, “the specimen of Mr. 

Townsend is labelled, ‘Banks of the Columbia River; May 

9th, 1835;’ that of Mr. Nuttall was, I believe, obtained in the 

same locality” (Bachman, 1842:32).

Distribution: Canada and USA: southwesternmost corner of 

British Columbia south through western Washington, western Oregon, and northwestern corner of California, west of 

the Cascade Mountains (Hall, 1981; Carraway et al., 1993).

Family Soricidae G. Fischer, 1814

1814. Soricinorum G. Fischer, Zoognosia 3:x.

Subfamily Soricinae G. Fischer, 1814

1814. Soricinorum G. Fischer, Zoognosia 3:x. Proposed as a family-level 

taxon 

Tribe Blarinini Kretzoi, 1965

1965. Blarinini Kretzoi, Vertebrata Hungarica, 7(1–2):124. 

Genus Blarina Gray, 1838

1838. Blarina Gray, Proc. Zool. Soc. London 1837(5):124; 14 June. Proposed as a subgenus of Corsira Gray, 1838 (=Sorex Linnaeus, 1758).

1842. Blarina: Lesson, Nouv. tableau mamm., p. 89. First use as a genus. 

1842. Brachysorex Duvernoy, Mag. Zool. Anat. Comp. Pal., ser. 2, 4(8):37. 

Proposed as a subgenus of Sorex Linnaeus, 1758. Type species: 

none selected; either Sorex (Brachysorex) brevicaudatus Duvernoy, 

1842 (=Sorex brevicaudus Say, 1822 = Blarina brevicauda) or Sorex 

(Brachysorex) harlani Duvernoy, 1842 (=Sorex parvus Say, 1822 = 

Cryptotis parvus). 

1843. Blaria: Gray, List Mamm. Brit. Mus., p. xxi. Incorrect subsequent 

spelling of Blarina Gray, 1838).

1848. Talposorex Pomel, Arch. Sci. Phys. Nat., 9:248. Type species: Talposorex platyurus Pomel, 1848 (=Sorex carolinensis Bachman, 1837 

= Blarina carolinensis), by original designation (not Talpasorex Schinz, 

1821 = Condylura Illiger, 1811; not Talpasorex Lesson, 1827 = Scalopus È. Geoffroy Saint-Hilaire, 1803).

1848. Galemys Pomel, Arch. Sci. Phys. Nat. 9:249. Part; also includes the 

subgenus Brachysorex, which contains the species Galemys micrurus

Pomel, 1848 (=Sorex talpoides Gapper, 1830 = Blarina brevicauda talpoides) and Galemys harlani Duvernoy, 1842 (=Sorex parvus Say, 1822 

= Cryptotis parvus).

1855. Anotus Wagner, Die Säugthiere Suppl. 5:550. Type species: Sorex carolinensis Bachman, 1837 (=Blarina carolinensis), by monotypy.

Type Species: Sorex talpoides Gapper, 1830 (=Blarina brevicauda talpoides), by original designation. 

Remarks: The genus was revised by Merriam (1895a) and 

Jones et al. (1984) and reviewed by Hall and Kelson (1959) 

and Hall (1981); phylogeography was studied by Brant and 

Orti (2002). Blarina brevicauda was revised by Webster et 

al. (2011); B. carolinensis was reviewed by Genoways and 

Choate (1998), and B. hylophaga was reviewed by George 

et al. (1981).

Blarina brevicauda (Say, 1822)

FIGURES 16–19 

Remarks: This is the type species for the genus Blarina Gray, 

1838. Webster et al. (2011) comprehensively revised the 

subspecies of Blarina brevicauda. The karyotype of the species is 2N = 48–50, FN = 48 (George et al., 1982; Thompson 

et al., 2011).

Blarina brevicauda aloga Bangs, 1902

1902. Blarina brevicauda aloga Bangs, Proc. New Engl. Zool. Club 3:76; 

31 March.

1902. Blarina brevicauda compacta Bangs, Proc. New Engl. Zool. Club

3:77; 31 March. Holotype: MCZ BANGS 9705, from “Nantucket, 

Massachusetts;” Nantucket County.

Holotype: MCZ BANGS 9727.

Type Locality: USA: “West Tisbury, Martha’s Vineyard, Massachusetts”; Dukes County.

Range: USA: Martha’s Vineyard and Nantucket islands, Massachusetts (Webster et al., 2011).

Remarks: Bangs (1902) distinguished the island forms Blarina 

brevicauda aloga (from Martha’s Vineyard) and B. brevicauda compacta (from Nantucket Island) from mainland 

B. brevicauda talpoides on the basis of their smaller size, 

and he distinguished the two subspecies from each other on 

the basis of pelage color. Hall (1981) recognized the two 

taxa as distinct, but Webster et al. (2011:30) synonymized 

them, noting that their “analyses indicate that individuals 

from Nantucket Island are slightly larger than those from 

Martha’s Vineyard.”

Blarina brevicauda brevicauda (Say, 1822)

FIGURES 16, 17

1822. Sorex brevicaudus Say, in James, Acct. Exped. Pittsb. Rocky Mtns., 

1:164; 31 December.

1842. [Sorex] Brachysorex brevicaudatus: Duvernoy, Mag. Zool. Anat.

Comp. Pal. 1842:38. Unjustified emendation.

1857. Blarina brevicauda: Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):42.

1891. Blarina costaricensis J. A. Allen, Bull. Am. Mus. Nat. Hist. 3:205; 

17 April. Holotype: AMNH 2800/3642, from “La Carpintera,” Cartago Province, Costa Rica. Merriam (1895a:10, 12–13) identified 
NUMBER 650 • 2 9

the holotype as Blarina brevicauda and thought it probably came 

from the Upper Mississippi Valley—possibly Iowa—where the collector, George K. Cherrie, had lived before he went to Costa Rica. 

Allen (1897:34) noted that the collector “still affirms that such an 

error was impossible, and that the specimen was actually taken at La 

Carpentera, Costa Rica.” Webster et al. (2011) confirmed that the 

specimen is B. brevicauda rather than B. hylophaga, which is also 

now known from Iowa.

1943. Blarina fossilis Hibbard, Univ. Kansas Sci. Bull. 29(2):238; 

15 October. Holotype: KUVP 6675, from mid-Pleistocene (Irvingtonian) sediments in “the Rezabek gravel pit in sec. 20, T.13S. 

R.11W., Lincoln county, Locality No. 5, Kansas.” Jones et al. 

(1984:84) stated that this taxon “most closely resembles Holocene 

B. b. brevicauda,” and Webster et al. (2011) placed it in synonymy 

with that taxon. 

1947. Blarina brevicauda manitobensis R. M. Anderson, Natl. Mus.

Canada Bull. 102(1946):23; 24 January. Holotype: CMN 8549, 

from “Max Lake, Turtle Mountains, Manitoba; latitude a little 

north of 49th parallel, longitude about 100 degrees west; altitude 

about 2,100 feet.”

Neotype: SMNH 37241, designated by Webster et al. (2011). 

The holotype (Figure 16) in the Philadelphia Museum (Godman, 1826:79–80 and accompanying plate) was lost or destroyed (Woodman, 2009). 

Type Locality: USA: “3 mi S, 4 mi E Fort Calhoun, Washington County, Nebraska, 41°24′43′′ N, 95°57′00′′ W.” The 

type locality of the neotype essentially matches the original type locality (“Engineer cantonment . . . taken near our 

cabins”), which was rediscovered by Carlson et al. (2004) 

and described by Genoways and Ratcliff (2008:6–7): “Engineer Cantonment is located in the extreme southeastern 

corner of modern Washington County, NE, at a place three 

miles south and four miles east of the town of Fort Calhoun 

(41°24′43′′ N latitude, 95°57′00.6′′ W longitude, SE 1⁄4 NW 

1⁄4 SE 1⁄4, Section 28, Township 17 N, Range 13 E). The site 

is 305 m in elevation.”

Distribution: Canada and USA: southeastern Saskatchewan 

and southern Manitoba south through eastern North Dakota, eastern South Dakota, eastern Nebraska, western and 

southern Minnesota, and the Mississippi Valley regions of 

FIGURE 16. Watercolor and ink of the holotypes of (top left) Sorex brevicaudus Say, 1822 (=Blarina brevicauda) and (bottom right) Sorex 

parvus Say, 1822 (=Cryptotis parvus) by Titian R. Peale. Image courtesy of the American Philosophical Society, Philadelphia (Titian Ramsay 

Peale Sketches).
3 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Wisconsin and Illinois to northern Missouri (Thompson et 

al., 2011; Webster et al., 2011).

Blarina brevicauda cumberlandensis Webster, Moncrief, 

Choate, and Genoways, 2011

2011. Blarina brevicauda cumberlandensis Webster, Moncrief, Choate, and 

Genoways, Virginia Mus. Nat. Hist. Mem. 10:31; 21 March.

Holotype: OMNH 37146.

Type Locality: USA: “Murfreesboro, Rutherford County, 

Tennessee.”

Distribution: USA: Cumberland Plateau region of central Tennessee and western and south central Kentucky (Webster et 

al., 2011).

Blarina brevicauda delmarvensis Webster, Moncrief, Choate, 

and Genoways, 2011

2011. Blarina brevicauda delmarvensis Webster, Moncrief, Choate, and Genoways, Virginia Mus. Nat. Hist. Mem. 10:32; 21 March.

Holotype: KU 45447.

Type Locality: USA: “Cambridge, Dorchester County, Maryland.”

Distribution: USA: southern half of Delaware and the eastern 

shore of Maryland and Virginia (Webster et al., 2011).

Blarina brevicauda jerryrchoatei Webster, Moncrief, and 

Genoways, 2011

2011. Blarina brevicauda jerryrchoatei Webster, Moncrief, and Genoways, in 

Webster et al., Virginia Mus. Nat. Hist. Mem. 10:34; 21 March. 

Holotype: KU 114306.

Type Locality: USA: “‘turnpike right-of-way 1 1⁄2 mi N, 1 1⁄

5 mi 

E Courthouse, Lawrence,’ Douglas County, Kansas.” 

Distribution: Kansas: valleys of the Kansas River and its tributaries in Douglas, Jefferson, and Leavenworth Counties; elevational distribution: 240–300 m.

Remarks: Where the distributions of Blarina brevicauda and 

B. hylophaga come into contact, the two species generally appear to have parapatric distributions with a narrow contact 

zone and minimal hybridization (Thompson et al., 2011). 

Blarina brevicauda jerryrchoatei represents an isolated population that is within the distribution of B. hylophaga. Webster 

et al. (2011:34) indicated that the taxon is “intermediate in 

most characters, not as massive as in specimens from elsewhere west of the Mississippi River.” Esselstyn and Timm 

(2006) reported that mitochondrial cytochrome b sequences 

from the population do not differ substantially from those 

of B. b. brevicauda. More complete study is needed to better 

elucidate the relationships of this Kansas River population.

Blarina brevicauda knoxjonesi Webster, 1996

1996. Blarina brevicauda knoxjonesi Webster, in Genoways and Baker, Contributions in Mammalogy, p. 52; 5 September.

Holotype: USNM 567349.

Type Locality: USA: “Carolina Beach, New Hanover Co., 

North Carolina (34°01′32′′N, 77°54′36′W).”

Distribution: USA: south of the Pamlico River in eastern North 

Carolina (Webster et al., 2011).

FIGURE 17. Sorex brevicaudatus Duvernoy, 1842 (=Sorex brevicaudus Say, 1842 = Blarina brevicauda). Image from Duvernoy (1842: pl. 52), 

courtesy of Biodiversity Heritage Library (https://www.biodiversitylibrary.org/), contributed by the Ernst Mayer Library, Museum of Comparative Zoology, Harvard University, Cambridge.
NUMBER 650 • 3 1

FIGURE 18. Copperplate engraving of Benjamin Barton’s (1806) “black shrew” from the vicinity of Philadelphia, the basis for Sorex 

niger Ord, 1815 (=Sorex talpoides Gapper, 1830 = Blarina brevicauda talpoides; Woodman, 2013). Image courtesy of the American 

Philosophical Society, Philadelphia, Pennsylvania (Benjamin Smith Barton Papers).

FIGURE 19. Sorex talpoides Gapper, 1830 (=Blarina brevicauda talpoides). From Gapper (1830: pl. 8), courtesy of Biodiversity Heritage Library (https://www.biodiversitylibrary.org/), contributed by The Natural History Museum Library, London.
3 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Blarina brevicauda talpoides (Gapper, 1830) 

FIGURES 18–20

1815. Sorex niger Ord, Zoology of North America, p. 291, in W. Guthrie 

[et al.], A New Geographical, Historical, and Commercial Grammar. 

Nomen nudum; based on Barton’s (1806) “black shrew” from “the 

vicinity of Philadelphia” (Woodman, 2013).

1830. Sorex talpoides Gapper, Zool. J. 5(31):202, pl. 8; June.

1837. Corsira (Blarina) talpoides: J. E. Gray, Proc. Zool. Soc. Lond. 5:124; 

November.

1837. Sorex dekayi Bachman, J. Acad. Nat. Sci. Phila. 7(2):377; November. Holotype: unknown, from “Queens county,” New York (De Kay, 

1842:18).

1842. [Sorex] concolor De Kay, Zool. New York, p. 18. Nomen nudum; 

working name for Sorex dekayi Bachman, 1837.

1848. G[alemys]. micrurus Pomel, Arch. Sci. Phys. Nat., 9:249; based on 

“Sorex de Kayi de Kay, 1842” = Sorex dekayi Bachman, 1837.

1857. Blarina angusticeps Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):47. Holotype: USNM 1318/2120, from “Burlington, Vermont.” The type and 

only specimen of this taxon was regarded by Merriam (1895a:10) 

as an individual with a deformed skull; see also Bole and Moulthrop 

(1942:111–112).

1895. Blarina telmalestes Merriam, N. Am. Fauna 10:15; 31 December. 

Holotype: USNM 71823, from “Lake Drummond, Dismal Swamp, 

Virginia.”

1899. Blarina simplicidens Cope, J. Acad. Nat. Sci. Phila. ser. 2, 11(2):219–

220. Holotype: ANSPVP 150, from mid-Pleistocene (late Irvingtonian) 

“carbonaceous layers of the Port Kennedy deposit,” Upper Merion 

Township, Montgomery County, Pennsylvania (Spamer et al., 1995; 

Bell et al., 2004). Jones et al. (1984) considered this fossil taxon “a representative of the talpoides semispecies.” Webster et al. (2011) treated it 

as a synonym of B. b. talpoides.

1902. Blarina brevicauda talpoides: Bangs, Proc. New Engl. Zool. Club

3:75; 31 March. First use of current name combination.

1908. Blarina brevicauda ozarkensis B. Brown, Mem. Am. Mus. Nat. Hist.

9(4):170; February. Holotype: AMNH 11794, from mid-Pleistocene 

(late Irvingtonian) sediments in Conard Fissure, “a mile north of the 

Buffalo River, four miles west of Willcockson and fifteen miles south of 

Harrison, near the northern line of Newton County, Arkansas.” Jones 

et al. (1984:81) treated this taxon “as a separate subspecies within the 

talpoides semispecies of B. brevicauda.” Webster et al. (2011) placed it 

in synonymy with B. b. talpoides.

1940. Blarina brevicauda pallida R. W. Smith, Am. Midl. Nat. 24(1):223; 

31 July. Holotype: MVZ 86682, from “Wolfville, Kings County, Nova 

Scotia.”

1942. Blarina brevicauda kirtlandi Bole and Moulthrop, Sci. Publ. Cleveland Mus. Nat. Hist. 5(6):99; 11 September. Holotype: CMNH 16895, 

from “The Holden Arboretum, Kirtland Township, Lake County, and 

Chardon Township, Geauga County, Ohio. (The county line bisects the 

type locality.)”

1942. Blarina brevicauda churchi Bole and Moulthrop, Sci. Publ. Cleveland 

Mus. Nat. Hist. 5(6):109; 11 September. Holotype: CMNH 3640, from 

“Roan Mountain, Mitchell County, North Carolina.” 

1942. Blarina brevicauda hooperi Bole and Moulthrop, Sci. Publ. Cleveland 

Mus. Nat. Hist. 5(6):110; 11 September. Holotype: UMMZ 77380, 

from “Lyndon, Caledonia County, Vermont.”

1943. Blarina brevicauda angusta Anderson, Ann. Rep. Provancher Soc. Nat.

Hist. 1942:52; 7 September. Holotype: CMN 11655, from “Kelly’s 

Camp, Berry Mountain Brook, near head of Grand Cascapedia River, 

Gaspé County, Quebec, attitude [sic], about 1,600 feet”.

Holotype: Natural History Museum Bristol, UK [now Bristol 

Museum and Art Gallery]; probably lost when the museum was destroyed by aerial bombing during World War II 

(Rhian Rowson, Bristol Museum and Art Gallery, email 

communication, 22 February 2017).

Type Locality: Canada: “the District of Upper Canada extending between York and Lake Simcoe,” southern Ontario. 

York is now within the city limits of Toronto. 

Distribution: Canada and USA: southernmost portions of 

Canada from western Ontario to Nova Scotia, south to the 

Ohio River in Illinois and western Indiana, eastern Kentucky, south along the Appalachian Mountains into northern Georgia and Alabama, and east through northern South 

Carolina to the eastern coast of the United States (Webster 

et al., 2011); elevational distribution from near sea level to 

1,950 m (USNM specimens).

Blarina carolinensis (Bachman, 1837)

FIGURE 20

Remarks: In general, the distributions of B. carolinensis and 

its constituent subspecies remain poorly circumscribed 

(Genoways and Choate, 1998; McCay, 2001). The karyotype of B. carolinensis is 2N = 37–46, FN = 44–45 (George 

et al., 1982). 

Blarina carolinensis carolinensis (Bachman, 1837) 

FIGURE 20

1837. Sorex carolinensis Bachman, J. Acad. Nat. Sci. Phila. 7(2):366; 

November.

1857. Blarina carolinensis: Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):45.

1895. Blarina brevicauda carolinensis: Merriam, N. Am. Fauna 10:13; 31 

December.

1984. Blarina carolinensis carolinensis: C. A. Jones et al., Spec. Publ. Carnegie Mus. 8:74; 11 September. First use of current name combination.

Neotype: USNM 574157, designated by Handley and Varn 

(1994).

Type Locality: USA: “beside Awendaw Creek, 3.2 km E 

Awendaw Post Office, Charleston County, South Carolina, 

in a thicket at the edge of a salt marsh” (Handley and Varn, 

1994). Bachman (1837:366) originally wrote, “It is found in 

various localities, both in the upper and maritime districts 

of South Carolina.” Merriam (1895a:13) restricted the type 

locality to “Eastern South Carolina.” 

Distribution: USA: Atlantic coastal plain and piedmont from 

south central and southeastern Virginia south to Alachua 

County, Florida, and west along the Gulf of Mexico coastal 

plain to central Mississippi, then north into the upland portion of western Tennessee, southern Illinois, central Arkansas, southeastern Oklahoma, and northeastern Texas. 
NUMBER 650 • 3 3

FIGURE 20. John Bachman’s (1837) shrews: from top to bottom, Sorex carolinensis

Bachman, 1837 (=Blarina carolinensis); Sorex longirostris Bachman, 1837; Sorex cinereus

Bachman, 1837 (=Blarina floridana Merriam, 1895 = Cryptotis parvus floridanus); Sorex 

dekayi Bachman, 1837 (=Sorex talpoides Gapper, 1830 = Blarina brevicauda talpoides). 

Image from Bachman (1837: pl. 23).
3 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Blarina carolinensis minima Lowery, 1943

1943. Blarina brevicauda minima Lowery, Occ. Pap. Mus. Zool. Louisiana 

State Univ. 13:218; 22 November.

1979. Blarina carolinensis minima: Sealander, A Guide to Arkansas Mammals, p. 44. First use of current name combination.

Holotype: LSU 2196.

Type Locality: USA: “Comite River, 13 mi. NE Baton Rouge, 

East Baton Rouge Parish, Louisiana.”

Distribution: USA: appears to occur in southeastern Texas, 

most of Louisiana, and the Mississippi Valley regions of 

western Mississippi, eastern Arkansas, and western Tennessee, possibly reaching the southeastern tip of Missouri, 

southern tip of Illinois, and western tip of Kentucky (Genoways and Choate, 1998; McCay, 2001). 

Remarks: Provisionally retained as a subspecies of B. carolinensis by Genoways and Choate (1998).

Blarina hylophaga Elliot, 1899

Remarks: The karyotype of B. hylophaga is 2N = 52, FN = 

60–62 (George et al., 1982).

Blarina hylophaga hylophaga Elliot, 1899

1899. Blarina brevicauda hulophaga Elliot, Field Col. Mus. Publ. 38, Zool.

Ser. 1(15):287; 24 May.

1905. Blarina brevicauda hylophaga: Elliot, Field Col. Mus. Publ. 105, Zool.

Ser. 6:461. Justified emendation of Blarina brevicauda hulophaga

Elliot, 1899. In a footnote, Elliot (1905:461) wrote, “Misprinted originally hulophaga.” The corrected spelling is a better transliteration of 

the Greek word ύλοφαγοσ, from which Elliot (1899b:288) derived the 

species name.

1939. Blarina brevicauda carolinensis: Blair, Am. Midl. Nat. 22:99; July. 

Part; not Sorex carolinensis Bachman, 1837 = Blarina carolinensis.

1972. Blarina carolinensis carolinensis: Genoways and Choate, Syst. Zool. 

21:114; March. Part; not Sorex carolinensis Bachman, 1837 = Blarina 

carolinensis.

1981. Blarina hylophaga hylophaga: George et al., Ann. Carnegie Mus.

50(21):504; 18 December. First use of current name combination.

Holotype: FMNH 6770.

Type Locality: USA: “Dougherty, Indian Territory”; Murray County, Oklahoma. Elliot (1905:461) subsequently 

provided a slightly more expanded locality: “Dougherty, 

Washita River, Chickasaw Nation, Indian Territory.”

Distribution: USA: southern Great Plains from southern Nebraska and southwest Iowa to east central Colorado, 

through Kansas and Missouri through northwestern Arkansas, central and eastern Oklahoma, to northeastern Louisiana (George et al., 1981).

Blarina hylophaga plumbea W. B. Davis, 1941

1941. Blarina brevicauda plumbea W. B. Davis, J. Mamm. 22(3):317; 14 

August.

1979. B[larina]. c[arolinensis]. plumbea: Schmidly and W. A. Brown, Southwest. Nat. 24(1):39; 30 March.

1981. Blarina hylophaga plumbea: George et al., Ann. Carnegie Mus. 

50(21):504; 18 December. First use of current name combination.

Holotype: TCWC 1541.

Type Locality: USA: “one-half mile west of Marano Mill, 

Aransas National Wildlife Refuge Aransas County, Texas.”

Distribution: USA: known only from two isolated populations 

in Aransas and Bastrop Counties, Texas (Reilly et al., 2005).

Blarina peninsulae Merriam, 1895

1895. Blarina carolinensis peninsulae Merriam, N. Am. Fauna 10:14; 31 

December.

1897. [Blarina brevicauda] peninsulae: Trouessart, Catalogus Mammalium,

Tam Viventium quam Fossilium 1:188.

2005. Blarina peninsulae: Hutterer, in MSW3, p. 270. First use of current 

name combination.

Holotype: USNM 70874.

Type Locality: USA: “Miami River, Dade County, Fla.”

Distribution: USA: most of peninsular Florida (Benedict et al., 

2006).

Remarks: Formerly treated as a subspecies of Blarina carolinensis (McCay, 2001; Benedict et al., 2006). Blarina peninsulae has a distinct karyotype (2N = 50–52, FN = 52) and 

morphology (George et al., 1982; Genoways and Choate, 

1998), leading Hutterer (2005:270) to recognize it as a distinct species. 

Blarina shermani Hamilton, 1955

1955. Blarina brevicauda shermani Hamilton, Proc. Biol. Soc. Wash. 68:37; 

20 May.

1992. Blarina carolinensis shermani: Layne, in Humphrey, Rare and Endangered Biota of Florida 1:328.

2006. Blarina shermani: Benedict et al., Occ. Pap. Mus. Texas Tech Univ. 

251:16; 23 January. First use of current name combination.

Holotype: CUMV 8026.

Type Locality: USA: “two miles north of Fort Myers, Lee 

County, Florida.”

Distribution: USA: restricted to southwestern coast of Charlotte, 

Lee, and Collier Counties, Florida (Benedict et al., 2006).

Remarks: McCay (2001) considered this taxon to be a subspecies of B. carolinensis, but Benedict et al. (2006) treated it 

as a distinct species.

Genus Cryptotis Pomel, 1848

1848. Cryptotis Pomel, Arch. Sci. Phys. Nat. 9:249; November. Proposed as 

a subgenus of Musaraneus Pomel, 1848. 

1848. Galemys Pomel, Arch. Sci. Phys. Nat. 9:249. Part; not Galemys Kaup, 

1829, a talpid.

1848. Brachysorex: Pomel, Arch. Sci. Phys. Nat. 9:249. Part; not Brachysorex Duvernoy, 1842.

1848. Musaraneus Pomel, Arch. Sci. Phys. Nat. 9:249. Part; not Musaraneus Brisson, 1762, which is unavailable; Musaraneus Pomel, 
NUMBER 650 • 3 5

1848 is a junior synonym of Crocidura Wagler, 1832 (Woodman, 

2004).

1857. Blarina: Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):48. Part; not Blarina

Gray, 1838.

1872. Cryptotus Milne-Edwards, Recherches des Mamm. 1:256. Incorrect 

subsequent spelling of Cryptotis Pomel, 1848.

1877. Soriciscus Coues, Bull. U.S. Geol. Geog. Surv. Terr. 3(3):649; 15 May. 

Type species: “Sorex parvus Say or S. cinereus Bachm” (=Cryptotis 

parvus parvus (Say, 1822) and Cryptotis parvus floridanus (Merriam, 

1895), respectively), by original designation. Proposed as a subgenus of 

Blarina Gray, 1838.

1901. Sorieiscus Elliot, Field Col. Mus. Publ. 45, Zool. Ser. 2(1):382; 

6 March. Incorrect subsequent spelling of Soriciscus Coues, 1877.

1911. Cryptotis: Miller, Proc. Biol. Soc. Wash. 24:221; 31 October (first use 

as a genus).

1958. Cryptotys Saban, in Piveteau, Traité de Paléontologie 6(2):846. Incorrect subsequent spelling of Cryptotis Pomel, 1848. 

1966. Xenosorex Schaldach, Säug. Mitteil. 14(4):289; October. Type species: 

Notiosorex phillipsii Schaldach, 1966 (=Cryptotis phillipsii (Schaldach, 

1966)), by original designation. Proposed as a subgenus of Notiosorex

Coues, 1877.

1984. Criptotis Durant and Péfaur, Revista de Ecología, Conservación y 

Ornitología Latinamericano 1(2):6. Incorrect subsequent spelling of 

Cryptotis Pomel, 1848.

Type Species: Musaraneus (Cryptotis) cinereus (Bachman, 

1837) (=Sorex cinereus Bachman, 1837 = Sorex parvus 

Say, 1822 = Cryptotis parvus), by monotypy. 

Remarks: Cryptotis was revised by Merriam (1895a) and reviewed by Hall and Kelson (1959) and Hall (1981); Middle 

American species were revised by Choate (1970). Species 

groups are based on Choate (1970) and Woodman and 

Timm (1993, 1999, 2017) and on molecular genetic analyses by Guevara and Cervantes (2013), He et al. (2015: 

tab. 1), and Baird et al. (2017). 

Although historically treated as feminine in gender 

(Miller, 1912a; Miller and Kellogg, 1955; Hall and Kelson, 

1959; Hall, 1981), a ruling by ICZN (2006) determined 

Cryptotis should be treated as a masculine noun.

Cryptotis alticola (Merriam, 1895)

1895. Blarina alticola Merriam, N. Am. Fauna 10:27; 31 December.

1912. Cryptotis alticola: Miller, Bull. U.S. Natl. Mus. 79:27; 31 December. 

First use of current name combination.

1967. Cryptotis euryrhynchis Genoways and Choate, Proc. Biol. Soc. Wash. 

80:203; 1 December. Holotype: KU107143, from “Volcán de Fuego 

(also called Volcán de Colima), 9800 ft, Jalisco.”

1970. Cryptotis goldmani alticola: Choate, Univ. Kansas Publ. Mus. Nat.

Hist. 19(3):245; 30 December.

Holotype: USNM 52047.

Type Locality: Mexico: “Mount Popocatepetl, Mexico (altitude, 11,500 feet).” 

Distribution: Transverse Volcanic Axis of central Mexico, 

from southern Jalisco and Colima to northern Hidalgo 

and northwestern Puebla; elevational distribution 1,060–

4,115 m (Woodman and Timm, 1999; Carraway, 2007).

Remarks: Molecular analyses (Guevara and Cervantes, 2014; 

He et al., 2015; Baird et al., 2017) position C. alticola as 

part of either the C. goldmani group or C. goodwini group. 

On biogeographical grounds, I tentatively place it in the former species group (Appendix A). 

Cryptotis aroensis Quiroga-Carmona and Molinari, 2012

2012. Cryptotis aroensis Quiroga-Carmona and Molinari, Zootaxa 3441:3; 

28 August.

Holotype: CVULA I-8548.

Type Locality: Venezuela: “Las Cumaraguas Sector, Sierra 

de Aroa, Municipio Cocorote, Estado Yaracuy, Venezuela 

(10°22′02.6′′ N, 68°49′20.4′′ W), elevation 1730 m.”

Distribution: Known only from the highlands of the Sierra de 

Aroa, northwestern Venezuela; elevational distribution is 

1,400–1,940 m (Quiroga-Carmona and Molinari, 2012; 

García et al., 2014).

Remarks: Cryptotis thomasi group.

Cryptotis berlandieri (Baird, 1857)

FIGURE 11

1857. Blarina berlandieri Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):53, 

pl. 18.

1895. Blarina parva: Merriam, N. Am. Fauna 10:17; 31 December. Part; not 

Sorex parvus Say, 1822.

1903. Blarina pergracilis Elliot, Field Col. Mus. Publ. 71, Zool. Ser. 3(8):149; 

20 March. Holotype: FMNH 8689, from “Ocotlan, state of Jalisco, 

Mexico.”

1903. B[larina]. b[erlandieri]. berlandieri: Elliot, Field Col. Mus. Publ. 71,

Zool. Ser. 3(8):149; 20 March. 

1911. Cryptotis pergracilis macer Miller, Proc. Biol. Soc. Wash. 24:223; 31 

October. Holotype: USNM 15565/38494, from “near Guanajuato 

City, Mexico”; state of Guanajuato.

1911. Cryptotis pergracilis pergracilis: Miller, Proc. Biol. Soc. Wash. 24:223; 

31 October.

1912. Cryptotis berlandieri: Miller, Bull. U.S. Natl. Mus. 79:25; 31 December. First use of current name combination.

1924. Cryptotis pergracilis macra: Miller, Bull. U.S. Natl. Mus. 128:31. 

Emendation of Cryptotis pergracilis macer Miller, 1911 for gender 

agreement.

1933. Cryptotis pergracilis nayaritensis Jackson, Proc. Biol. Soc. Wash. 

46:79; 27 April. Holotype: USNM 88015, from “Tepic, altitude 3000 

feet, state of Nayarit, Mexico.”

1941. Cryptotis parva berlandieri: W. B. Davis, J. Mamm. 22(4):413; 14 

November.

Lectotype: USNM 2159, designated by Lyon and Osgood 

(1909:237).

Type Locality: Mexico: “in the vicinity of Matamoros,” Tamaulipas (Baird, 1857:54). 

Distribution: USA and Mexico: southern Texas, Coahuila, 

Guanajuato, Jalisco, Michoacán, Nayarit, Nuevo León, San 
3 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Luis Potosí, and Tamaulipas; elevational distribution from 

near sea level to 2,715 m (Carraway, 2007).

Remarks: Cryptotis parvus group. Cryptotis berlandieri in 

Texas and northern Tamaulipas average larger than C. parvus from the central Great Plains. There is also preliminary 

evidence that populations of C. berlandieri in Texas and 

northern Tamaulipas may differ from those in more southern parts of the distribution in Mexico (Choate, 1970). A 

more comprehensive study of populations within C. berlandieri than has been undertaken so far may reveal more 

genetic variation than currently recognized.

Cryptotis brachyonyx Woodman, 2003

2003. Cryptotis brachyonyx Woodman, Proc. Biol. Soc. Wash. 116(4):855; 

31 December.

Holotype: BM 99.10.3.2.

Type Locality: “Colombia: Department of Cundinamarca: ‘La 

Selva, near Bogotá.’”

Distribution: Colombia: eastern range of the Andes in Departamento de Cundinamarca; known elevational distribution: 

1,300–2,715 m (Woodman and Péfaur, 2008).

Remarks: Cryptotis nigrescens group.

Cryptotis cavatorculus Woodman, 2015

2015. Cryptotis cavatorculus Woodman, Ann. Carnegie Mus. 83(2):114; 15 

November.

Holotype: FLMNH 27718.

Type Locality: Honduras: “desiccated cloud forest on a ridge 

above El Cedral, ca. 1900 m (ca. 14°54′ N, 88°06′ W), 

Parque National de Santa Barbara, Santa Barbara Dept.”

Distribution: Honduras: known only from the holotype.

Remarks: Cryptotis goodwini group.

Cryptotis celaque Woodman, 2015

2015. Cryptotis celaque Woodman, Ann. Carnegie Mus. 83(2):106; 

15 November.

Holotype: CM 112882.

Type Locality: Honduras: “Campamento Don Tomás, 2075 m, 

Celaque National Park, ca. 11 km SW of Gracias, Lempira 

Dept.”

Distribution: Honduras: known only from Sierra de Celaque, 

Departamento de Lempira; elevational distribution: 1,430–

2,560 m (Woodman, 2015).

Remarks: Cryptotis goodwini group.

Cryptotis colombianus Woodman and Timm, 1993

1993. Cryptotis colombiana Woodman and Timm, Field. Zool., n.s. 

74(1452):24; 30 September.

Holotype: FMNH 69816.

Type Locality: “Colombia; Central Cordillera; Antioquia 

Dept., Sonsón; 15 km E of Río Negrito; 1750 m.”

Distribution: Colombia: northern central range of the Andes 

in Antioquia Department; elevational distribution: 1,750–

2,150 m (Woodman, 2003).

Remarks: Cryptotis nigrescens group.

Cryptotis dinirensis Quiroga-Carmona and DoNascimiento, 2016

2016. Cryptotis dinirensis Quiroga-Carmona and DoNascimiento, Mamm.

Biol. 81(2016):1–12; 3 May. 

Holotype: EBRG 24723.

Type Locality: Venezuela: “Las Antenas sector of the Parque 

Nacional Dinira, along Carache–La Peña road, Trujillo 

State, Venezuela, 9°41′ N, 70°04′ W, elevation 2550 m.”

Distribution: Venezuela: northernmost portion of the eastern 

branch of the Venezuelan Andes Cordillera; elevational distribution: 2,430–3,100 m (Quiroga-Carmona and DoNascimiento, 2016).

Remarks: Cryptotis thomasi group.

Cryptotis endersi Setzer, 1950

1950. Cryptotis endersi Setzer, J. Wash. Acad. Sci. 40(9):300; 29 September.

Holotype: ANSP 20955.

Type Locality: Panama: “Cylindro, Province of Chiriquí.” The 

type locality is probably on the Caribbean slope in Bocas del 

Toro Province (Pine et al., 2002).

Distribution: Panama: known from two specimens taken in 

highlands near the boundary between Bocas del Toro and 

Chiriquí Provinces; known elevation: 1,800–1,856 m (Pine 

et al., 2002).

Remarks: Cryptotis thomasi group. 

Cryptotis equatoris (Thomas, 1912)

1912. Blarina equatoris Thomas, Ann. Mag. Nat. Hist., ser. 8, 9:409; April.

1913. Blarina meridensis: Lönnberg, Ark. Zool. 8(16):1. Part; not Blarina 

meridensis Thomas, 1898.

1921. Blarina aequatoris Lönnberg, Ark. Zool. 14(4):4. Incorrect subsequent 

spelling of Blarina equatoris Thomas, 1912.

1921. [Cryptotis] equatoris: Thomas, Ann. Mag. Nat. Hist., ser. 9, 8(45):354. 

First use of current name combination.

1958. Cryptotis thomasi equatoris: Cabrera, Rev. Mus. Argentino 4(1):47. 

Part.

1997. Cryptotis e[quatoris]. equatoris: Vivar et al., Am. Mus. Novit. 3202:6; 

29 August.

Holotype: BM 99.9.9.3.

Type Locality: Ecuador: “Sinche, Guabanda, 4000 m”; Hacienda 

de Sinche, northeast of Guaranda, Departamento de Bolívar.

Distribution: Ecuador: western Andean cordillera from northern Carchí Province south to central Bolivar Province; elevational distribution: 1,800–4,000 m (Moreno-Cárdenas and 

Albuja, 2014). Populations of C. equatoris reported from 

northern Peru (Pacheco et al., 2009) have since been reidentified as C. montivagus (Zeballos et al., 2018).

Remarks: Cryptotis thomasi group.
NUMBER 650 • 3 7

Cryptotis evaristoi Zeballos, Pino, Medina, Pari, and Ceballos, 

2018

2018. Cryptotis evaristoi Zeballos, Pino, Medina, Pari, and Ceballos, in Zeballos et al., Zootaxa 4377(1):61; 31 January.

Holotype: MUSA 7427.

Type Locality: Peru: “Miraflores, San Ignacio, Cajamarcas, 

Peru (5°12′ S and 79°12′ W) at 2,800 m above sea level.” 

Distribution: Peru: montane forest and páramo in Tabaconas 

Namballe National Sanctuary, Cajamarca Province, northern Peru; elevational distribution: 2,700–3,780 m (Zeballos 

et al., 2018).

Remarks: Cryptotis thomasi group. 

Cryptotis goldmani (Merriam, 1895)

Remarks: Cryptotis goldmani group. Cryptotis goldmani was 

once thought to be much more widespread (Choate, 1970), 

including populations near San Cristóbal de las Casas, Chiapas (C. griseoventris), and from Todos Santos Cuchumatán, Guatemala (C. mam). Carraway (2007:19) formally 

distinguished populations in Guerrero (C. goldmani goldmani) from those in Oaxaca (C. goldmani machetes) on the 

basis of the former’s “venter pelage with white-tipped hairs” 

rather than blond-tipped hairs. The two populations can be 

partially separated using discriminant function analysis of a 

suite of cranial measurements.

Cryptotis goldmani goldmani (Merriam, 1895) 

1895. Blarina mexicana goldmani Merriam, N. Am. Fauna 10:25; 31 

December.

1912. Cryptotis mexicana goldmani: Miller, Bull. U.S. Natl. Mus. 79:27; 31 

December.

1933. Cryptotis guerrerensis Jackson, Proc. Biol. Soc. Wash. 46:80; 27 April. 

Holotype: USNM 126895, from “Omilteme, altitude about 8000 feet, 

state of Guerrero, Mexico.”

1970. Cryptotis goldmani goldmani: Choate, Univ. Kansas Publ. Mus.

Nat. Hist. 19(3):247; 30 December. Part; first use of current name 

combination.

Holotype: USNM 70244.

Type Locality: Mexico: “mountains near Chilpancingo, Guerrero, Mexico (altitude, 10,000 feet).”

Distribution: Mexico: highland region around Omilteme, 

Guerrero; elevational distribution: 1,550–3,000 m (Woodman and Timm, 1999; Carraway, 2007).

Cryptotis goldmani machetes (Merriam, 1895) 

1895. Blarina mexicana machetes Merriam, N. Am. Fauna 10:26; 31 

December.

1895. Blarina fossor Merriam, N. Am. Fauna 10:28; 31 December. Holotype: USNM 68545, from “Mount Zempoaltepec, Oaxaca, Mexico 

(altitude, 10,500 feet).”

1911. Cryptotis frontalis Miller, Proc. Biol. Soc. Wash. 24:222; 31 October. Holotype: USNM 123429, from “near the City of Tehuantepec, 

Mexico”; state of Oaxaca.

1912. Cryptotis mexicana machetes: Miller, Bull. U.S. Natl. Mus. 79:27; 31 

December.

1912. Cryptotis fossor: Miller, Bull. U.S. Natl. Mus. 79:28; 31 December.

1967. Cryptotis mexicana mexicana: J. K. Jones and Genoways, J. Mamm. 

48(2):321; 20 May. Part; not Blarina mexicana Coues, 1877 (=Cryptotis mexicanus).

1970. Cryptotis goldmani goldmani: Choate, Univ. Kansas Publ. Mus. Nat.

Hist. 19:247; 30 December. Part; not Blarina goldmani Merriam, 1895 

(=Cryptotis goldmani goldmani).

2007. Cryptotis goldmani machetes: Carraway, Monogr. West. N. Am. Nat. 

3:20; 22 June. First use of current name combination.

Holotype: USNM 71456.

Type Locality: Mexico: “mountains near Ozolotepec, Oaxaca 

(altitude, 10,000 feet).”

Distribution: Mexico: highlands of Oaxaca; elevational distribution: 2,250–3,200 m (Woodman and Timm, 1999; Carraway, 2007).

Cryptotis goodwini Jackson, 1933

1933. Cryptotis goodwini Jackson, Proc. Biol. Soc. Wash. 46:81; 27 April.

Holotype: USNM 77074.

Type Locality: Guatemala: “Calel, altitude 10200 feet”; Quetzaltenango Department (see also Goldman, 1951).

Distribution: Guatemala: Cryptotis goodwini sensu stricto 

are known exclusively from highlands of southern Guatemala; elevational distribution: 1,200–3,350 m (Woodman, 

2015).

Remarks: Cryptotis goodwini group. Once considered a species widespread in highlands in Guatemala, El Salvador, 

and Honduras (Choate, 1970; Woodman et al., 2012), 

C. goodwini sensu latu included several cryptic species 

now recognized as distinct (C. cavatorculus, C. celaque, 

C. lacertosus, C. magnimanus, C. mam, C. mccarthyi, 

C. oreoryctes), and all are variously adapted for more fossorial behavior (Woodman and Gaffney, 2014). A record 

of Cryptotis “goodwini” from Chiapas (e.g., Guevara and 

Cervantes, 2014) represents a distinct species that is being 

described. 

Cryptotis gracilis Miller, 1911

1911. Cryptotis gracilis Miller, Proc. Biol. Soc. Wash. 24:221; 31 October.

1944. Cryptotis jacksoni Goodwin, Am. Mus. Novit. 1267:1; 10 December. 

Holotype: USNM 116649, from “Volcan Irazu”; Cartago Province, 

Costa Rica.

Holotype: USNM 12236/38471.

Type Locality: Costa Rica: “head of Rio Lari, near base of Pico 

Blanco, Talamanca, Costa Rica, by Wm. M. Gabb. Altitude 

about 6,000 feet”; Limón Province.

Distribution: Costa Rica and Panama: portions of the Cordillera Central and Cordillera de Talamanca; elevational distribution: 1,830–3,540 m.

Remarks: Cryptotis goodwini group.
3 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Cryptotis griseoventris Jackson, 1933

1933. Cryptotis griseoventris Jackson, Proc. Biol. Soc. Wash. 46:80; 27 April.

1970. Cryptotis goldmani goldmani: Choate, Univ. Kansas Publ. Mus. Nat.

Hist. 19:247; 30 December. Part; not Blarina goldmani Merriam, 1895 

(=Cryptotis goldmani).

Holotype: USNM 75894.

Type Locality: Mexico: “San Cristobal, altitude, 9500 feet, 

state of Chiapas.”

Distribution: Mexico: known only from the region of Los 

Altos de Chiapas, northern Chiapas; elevational distribution: 2,200–2,900 m (Guevara et al., 2014).

Remarks: Cryptotis goodwini group.

Cryptotis hondurensis Woodman and Timm, 1992

1992. Cryptotis hondurensis Woodman and Timm, Proc. Biol. Soc. Wash. 

105(1):2; 12 March.

Holotype: KU 106941.

Type Locality: Honduras: “Francisco Morazán Department; 

12 km WNW of El Zamorano, W slope of Cerro Uyuca 

[=Cerro Oyuca; ca. 14°05′ N, 87°06′ W], 1680 m.”

Distribution: Honduras: known only from forested highlands 

east of Tegucigalpa; elevational distribution: 1,680–1,700 m.

Remarks: Cryptotis nigrescens group.

Cryptotis lacandonensis Guevara, Sánchez-Cordero, LeónPaniagua, and Woodman, 2014

2014. Cryptotis lacandonensis Guevara, Sánchez-Cordero, León-Paniagua, 

and Woodman, J. Mamm. 95(4):743; August.

Holotype: MZFC 7168.

Type Locality: Mexico: “Yaxchilán Archaeological Site, 90-m elevation, municipality of Ocosingo, Chiapas, Mexico (. . . 16°54′

N, 90°58′ W). The type locality of the Lacandona shrew is in 

lowland tropical rain forest dominated by trees 40 m in height 

on the floodplain of the Lacantún–Usumacinta rivers.”

Distribution: Mexico: known only from the Lacandona rain 

forest, Chiapas; known elevation: 90 m.

Remarks: Cryptotis nigrescens group. 

Cryptotis lacertosus Woodman, 2010

2010. Cryptotis lacertosus Woodman, J. Mamm. 91(3):576; 16 June.

2017. Cryptotis lasertosus Matson and Ordóñez-Garza, Zootaxa 

4236(3):476; 24 February. Incorrect subsequent spelling.

Holotype: USNM 569443. 

Type Locality: Guatemala: “north-facing slope with abundant 

downed trees and mosses in a relatively closed-canopy cloud 

forest dominated by oaks, pines, and firs; 5 km SW San 

Mateo Ixtatán, 3,110 m, Huehuetenango.”

Distribution: Guatemala: known only from the northern Sierra 

de los Cuchumatanes near San Mateo Ixtatán, western Guatemala; elevational distribution: 2,680–3,110 m.

Remarks: Cryptotis goodwini group.

Cryptotis magnimanus Woodman and Timm, 1999

1999. Cryptotis goodwini magnimana Woodman and Timm, Field. Zool., 

n.s. 91(1497):11; 13 January.

2011. Cryptotis magnimanus: Woodman, Zool. J. Linn. Soc. 163:1268; December. First use of current name combination.

Holotype: KU 144611.

Type Locality: Honduras: “2.5 km N, 1.6 km E Cerro San Juanillo 

[14°30′ N, 87°53′ W], Reserva Biológica Cordillera de Montecillos, Comayagua Department, Honduras, 1750 m.”

Distribution: Honduras: known only from the type locality.

Remarks: Cryptotis goodwini group.

Cryptotis magnus (Merriam, 1895) 

1895. Blarina magna Merriam, N. Am. Fauna 10:28; 31 December.

1912. Cryptotis magna: Miller, Bull. U.S. Natl. Mus. 79:28; 31 December. 

First use of current name combination.

2010. Cryptotis magnus: Woodman, J. Mamm. 91(3):572; 16 June. 

Holotype: USNM 68575.

Type Locality: Mexico: “Totontepec, Oaxaca (altitude, 6,800 

feet).” 

Distribution: Mexico: known only from highlands in Oaxaca; 

elevational distribution: 1,500–2,850 m.

Remarks: Cryptotis mexicanus group.

Cryptotis mam Woodman, 2010

2010. Cryptotis mam Woodman, J. Mamm. 91(3):573; 16 June.

Holotype: USNM 77053.

Type Locality: Guatemala: “approximately 10,000 feet in cloud 

forest dominated by cypress, fir, and pine on the upper reaches 

of a spurlike ridge above Todos Santos Cuchumatán [approximately 15°36′ N, 91°37′ W], Huehuetenango, Guatemala.”

Distribution: Guatemala: Sierra de los Cuchumatanes, western 

Guatemala; elevational distribution: 2,740–3,350 m.

Remarks: Cryptotis goodwini group. Cryptotis mam previously was considered part of C. goldmani goldmani (Choate, 1970), C. goodwini (Choate, 1970; Woodman and 

Timm, 1999), and C. griseoventris (Woodman and Timm, 

1999), but it can be clearly distinguished from those species by cranial and postcranial morphology (Woodman and 

Stephens, 2010; Woodman, 2011a). 

Cryptotis mayensis (Merriam, 1901) 

1901. Blarina mayensis Merriam, Proc. Wash. Acad. Sci. 3:559; 

29 November.

1912. Cryptotis mayensis: Miller, Bull. U.S. Natl. Mus. 79:26; 31 December. 

First use of current name combination.

1917. Blarina mexicana: Gaumer, Monografía de los Mamíferos de Yucatán, p. 249. Part; not Blarina mexicana Coues, 1877 (=Cryptotis 

mexicanus).

1935. Cryptotis micrura: Murie, Misc. Publ. Mus. Zool., Univ. Mich. 26:17; 

15 July. Part; not Sorex micrurus Tomes, 1862 (=Blarina tropicalis Merriam, 1895 = Cryptotis tropicalis).
NUMBER 650 • 3 9

1970. Cryptotis nigrescens mayensis: Choate, Univ. Kansas Publ. Mus. Nat.

Hist. 19:275; 30 December.

Holotype: USNM 108087.

Type Locality: Mexico: “a Maya ruin at Chichenitza, Yucatan.”

Distribution: Central America: the Yucatan Peninsula in Belize, 

Guatemala, and Mexico, as well as a cryptic, isolated record 

from owl pellets in a cave in Guerrero, Mexico; elevational 

distribution from near sea level to 600 m (Woodman and 

Timm, 1993; Engilis et al., 2012).

Remarks: Cryptotis nigrescens group.

Cryptotis mccarthyi Woodman, 2015

2015. Cryptotis mccarthyi Woodman, Ann. Carnegie Mus. 83(2):110; 

15 November.

Holotype: CM 119693.

Type Locality: Honduras: “1500–1620 m; ca. 0.3 km SW of 

the Visitors Center, Cusuco National Park, Cortés Dept.”

Distribution: Honduras: known only from the Sierra de Omoa, 

which occupies parts of Cortés and northern Santa Barbara 

Departments; elevational distribution: ~1,500–1,620 m 

(Woodman, 2015).

Remarks: Cryptotis goodwini group.

Cryptotis medellinius Thomas, 1921

1921. Cryptotis medellinius Thomas, Ann. Mag. Nat. Hist., ser. 9, 8(45):354; 

September. 

1932. Cryptotis medellinus Tate, J. Mamm. 13(3):224; August. Incorrect 

subsequent spelling of Cryptotis medellinius Thomas, 1921.

1958. Cryptotis thomasi medellinius: Cabrera, Rev. Mus. Argentino 4(1):48.

2002. Cryptotis medellinia: Woodman, Proc. Biol. Soc. Wash. 115(2):252; 

2 July. Emendation for gender agreement.

Holotype: BM 21.7.1.9.

Type Locality: Colombia: “San Pedro, 30 km. north of Medellin”; Antioquia Province.

Distribution: Colombia: northern half of the central Andean 

Cordillera and the northern tip of the western Andean Cordillera; elevational distribution: 2,000–3,800 m (Woodman 

and Péfaur, 2008).

Remarks: Cryptotis thomasi group.

Cryptotis meridensis (Thomas, 1898)

1898. Blarina meridensis Thomas, Ann. Mag. Nat. Hist., ser. 7, 1(6):457; 

June.

1921. C[ryptotis]. meridensis: Thomas, Ann. Mag. Nat. Hist., ser. 9, 8:354. 

First use of current name combination.

1958. Cryptotis thomasi meridensis: Cabrera, Rev. Mus. Argentino 4(1):48. 

Part.

1997. Cryptotis thomasi thomasi: Díaz et al., in Wielgolaski (ed.), Polar and 

Alpine Tundra, p. 293, Ecosystems of the World 3. Part; not Blarina 

thomasi Merriam, 1897 (=Cryptotis thomasi).

1998. Cryptotis meridensis meridensis: Linares, Mamíferos de Venezuela, 

p. 106.

Holotype: BM 98.5.15.5.

Type Locality: Venezuela: “Merida, alt. 2165 m,” modified to 

“Montes del Valle Merida 2165 m” by Woodman (2002) 

on the basis of information recorded on the skin label of 

the holotype.

Distribution: Venezuela: found in the Cordillera de Mérida 

in eastern Táchira, Mérida, and possibly southern Trujillo 

states; elevational distribution: 1,640–3,950 m (Woodman 

and Díaz de Pascual, 2004; Woodman and Péfaur, 2008).

Remarks: Cryptotis thomasi group.

Cryptotis merriami Choate, 1970

1970. Cryptotis nigrescens merriami Choate, Univ. Kansas Publ. Mus. Nat.

Hist. 19:277; 30 December.

1970. Cryptotis nigrescens nigrescens: Choate, Univ. Kansas Publ.

Mus. Nat. Hist. 19:281; 30 December. Part; not Blarina nigrescens

J. A. Allen, 1895 (=Cryptotis nigrescens).

1993. Cryptotis merriami: Woodman and Timm, Field. Zool., n.s. 

74(1452):14; 30 September. First use of current name combination.

Holotype: USNM 77050.

Type Locality: Guatemala: “Jacaltenango, 5400 ft., 

Huehuetenango.” 

Distribution: Central America: occurs at middle elevations 

from western Guatemala to the Tilarán Highlands of Costa 

Rica; elevational distribution: 600–1,665 m (Woodman and 

Timm, 1993; Woodman et al., 2012).

Remarks: Cryptotis nigrescens group. Genetic analyses (He et 

al., 2015; Baird et al., 2017) suggest that C. merriami may 

be a complex of at least two cryptic species. 

Cryptotis merus Goldman, 1912

1912. Cryptotis merus Goldman, Smithson. Misc. Coll. 60(2):17; 20 

September.

1959. Cryptotis mera: Hall and Kelson, Mammals of North America, p. 61; 

31 March. Emendation for gender agreement.

1966. C[ryptotis]. n[igrescens]. mera: Handley, in Wenzel and Tipton (eds.), 

Ectoparasites of Panama, p. 756; 22 November.

1970. Cryptotis nigrescens nigrescens: Choate, Univ. Kansas Publ. Mus. Nat. 

Hist. 19:279; 30 December. Part; not Blarina nigrescens J. A. Allen, 

1895 (=Cryptotis nigrescens).

Holotype: USNM 178976.

Type Locality: Panama: “near head of Rio Limon (altitude 

4,500 feet), Mount Pirri, eastern Panama.” 

Distribution: Panama and Colombia: Known only from Serranía de Pirre, Cerro Tacarcuna, and Serranía de Darién, 

Darién Province, along the Panama/Colombia border 

(Woodman and Timm, 1993; Woodman and Péfaur, 

2008).

Remarks: Cryptotis nigrescens group.

Cryptotis mexicanus (Coues, 1877) 

1877. Blarina (Soriciscus) mexicana Coues, Bull. U.S. Geol. Geog. Surv.

Terr. 3:652; 15 May. Not Sorex mexicanus Kerr, 1792, a geomyid; not 
4 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Blaria Mexicana Gray, 1847 from “Coban, South America,” a nomen 

nudum. 

1911. Cryptotis mexicana: Miller, Proc. Biol. Soc. Wash. 24:221; 31 October. 

First use of current name combination.

1912. Cryptotis mexicana mexicana: Miller, Bull. U.S. Natl. Mus. 79:26; 31 

December.

1969. Cryptotis mexicana peregrina: Goodwin, Bull. Am. Mus. Nat. Hist. 

141(1):39; 30 April. Part; not Blarina peregrina Merriam, 1895 

(=Cryptotis peregrinus).

Holotype: USNM 3525/4438.

Type Locality: Mexico: “Xalapa, Mexico”; Jalapa, state of 

Veracruz. 

Distribution: Mexico: highland regions from east central Hidalgo and northern Puebla south through west central 

Oaxaca and Chiapas; elevational distribution: 520–3,200 m 

(Choate, 1970; Carraway, 2007).

Remarks: Cryptotis mexicanus group.

Cryptotis monteverdensis Woodman and Timm, 2016 

2016. Cryptotis monteverdensis Woodman and Timm, Mamm. Res.

62(2017):93, published online 27 August 2016.

Holotype: KU 134852.

Type Locality: Costa Rica: “Puntarenas Province, Monteverde, 

Monteverde Cloud Forest Reserve, in ‘cloud forest at [the 

continental] divide’ (ca. 10°18′ N, 84°47′ W).”

Distribution: Costa Rica: Known only from the type locality; 

this species is probably restricted to the highest elevations of 

Cordillera de Tilarán (Woodman and Timm, 2017).

Remarks: Cryptotis thomasi group.

Cryptotis montivagus (Anthony, 1921)

1921. Blarina montivaga Anthony, Am. Mus. Novit. 20:5; 3 November.

1925. [Cryptotis] montivaga: Cabrera, Gen. Mamm. Insect. Galeop., p. 134. 

First use of current name combination.

1958. Cryptotis montivagus: Cabrera, Rev. Mus. Argentino 4(1):47. 

1997. Cryptotis peruviensis: Vivar, Pacheco, and Valqui, Am. Mus. Novit. 

3202:7; 29 August. Part, not Cryptotis peruviensis Vivar et al., 1997.

2009. Cryptotis equatoris: Pacheco et al., Rev. Peruana Biol. 16:11; August. 

Part, not Cryptotis equatoris (Thomas, 1912).

Holotype: AMNH 47200.

Type Locality: Ecuador: “Bestion, Prov. del Azuay, Ecuador; 

altitude 10,000 ft.”

Distribution: Ecuador and Peru: Andean highlands of southern 

Chimborazo, Azuay, eastern Loja, and western ZamoraChinchipe Provinces of Ecuador and Piura Province, 

northern Peru; elevational distribution: 2,500–3,910 m 

(Woodman and Péfaur, 2008; Moreno-Cárdenas and Albuja, 2014; Zeballos et al., 2018). Zeballos et al. (2018) 

listed (but did not map) two localities for C. montivagus

in Imbabura Province, northern Ecuador, which would 

represent a large northward extension of the geographical 

distribution of this species. One locality (Laguna Atillo) 

appears to be more appropriately located in Chimborazo, 

and the second locality (Páramo Angochagua) needs to be 

substantiated.

Remarks: Cryptotis thomasi group. Small-eared shrews from 

Piura Department, Peru, earlier identified as C. peruviensis

(Vivar et al., 1997) and later as C. equatoris (Pacheco et 

al., 2009), were more recently identified as C. montivagus

(Zeballos et al., 2018).

Cryptotis nelsoni (Merriam, 1895)

1895. Blarina nelsoni Merriam, N. Am. Fauna 10:26; 31 December.

1912. Cryptotis nelsoni: Miller: Bull. U.S. Natl. Mus. 79:27; 31 December. 

First use of current name combination.

1970. Cryptotis mexicana nelsoni: Choate, Univ. Kansas Publ. Mus. Nat.

Hist. 19:234; 30 December.

Holotype: USNM 65437.

Type Locality: Mexico: “Volcano of Tuxtla, Vera Cruz, Mexico (altitude, 4,800 feet).”

Distribution: Mexico: known only from the geographically 

isolated Volcán San Martín Tuxtla, Veracruz; elevational 

distribution: 1,300–1,500 m (Choate, 1970; Cervantes and 

Guevara, 2010).

Remarks: Cryptotis mexicanus group. 

Cryptotis niausa Moreno-Cárdenas and Albuja, 2014

2014. Cryptotis niausa Moreno-Cárdenas and Albuja, Papéis Avulsos de 

Zoologia 54(28):408.

Holotype: MEPN 9627.

Type Locality: Ecuador: “provincia de Napo: cantón de Quijos: 

Papallacta (Páramos de la Virgen), 00°20′49.2′′ S, 78°12′0′′

W, 3700 m, a 64 km al oriente de la ciudad de Quito.”

Distribution: Ecuador: Páramos and montane forests, from the 

Andes of northern Carchí Province south along the eastern 

cordillera to western Napo and eastern Pichincha Provinces; 

elevational distribution: 2,800–3,900 m.

Remarks: Cryptotis thomasi group. Specimens identified by 

Voss (2003:18) as Cryptotis cf. montivagus are referable to 

this species. The name niausa is used as a noun in apposition. It is adapted from ñausa, which means “blind” in 

the Kichwa language of Colombia, Ecuador, and Peru, with 

ñausa ucucha (blind mouse) being a local name for a shrew 

(Moreno-Cárdenas and Albuja, 2014).

Cryptotis nigrescens (J. A. Allen, 1895) 

1895. Blarina (Soriciscus) nigrescens J. A. Allen, Bull. Am. Mus. Nat. Hist.

7(10):339; 8 November.

1911. C[ryptotis] nigrescens: Miller, Proc. Biol. Soc. Wash. 24:222; 31 October. First use of current name combination.

1950. Cryptotis zeteki Setzer, J. Wash. Acad. Sci. 40(9):299; 29 September. 

Holotype: USNM 290466, from “Cerro Punta (lat. 8°42′ N., long. 

82°48′ W.), 6,500 feet, Chiriquí Province, Republic of Panama.”
NUMBER 650 • 4 1

1954. Cryptotis tersus Goodwin, Am. Mus. Novit. 1677:1; 28 June. Holotype: AMNH 164695, from “Santa Clara, 4200 feet elevation, on 

the Pan American Highway, 15 miles from the border of Costa Rica, 

Chiriqui Province, Republic of Panama.”

1966. C[ryptotis]. n[igrescens]. zeteki: Handley, in Wenzel and Tipton (eds.), 

Ectoparasites of Panama, p. 756; 22 November.

1970. Cryptotis parva orophila: Choate, Univ. Kansas Publ. Mus. Nat. Hist. 

19(3):264; 30 December. Part; not Blarina orophila J. A. Allen, 1895 

(=Cryptotis orophilus). 

Holotype: AMNH 9591/7952.

Type Locality: Costa Rica: “San Isidro (San José).”

Distribution: Central America: highlands in Costa Rica and 

western Panama; elevational distribution: 870–2,865 m 

(Woodman and Timm, 1993).

Remarks: Cryptotis nigrescens group.

Cryptotis obscurus (Merriam, 1895) 

1895. Blarina obscura Merriam, N. Am. Fauna 10:23; 31 December.

1912. Cryptotis obscura: Miller, Bull. U.S. Natl. Mus. 79:26; 31 December. 

First use of current name combination.

1954. Cryptotis mexicana madrea Goodwin, Am. Mus. Novit. 1670:1; 28 

June. Holotype: AMNH 147901, from “Rancho del Cielo, 3500 feet 

elevation, 5 miles northwest of Gómez Farías, Sierra Madre Oriental, 

Tamaulipas, Mexico.”

1970. Cryptotis mexicana obscura: Choate, Univ. Kansas Publ. Mus. Nat.

Hist. 19(3):235; 30 December.

2010. Cryptotis obscurus: Woodman, J. Mamm. 91(3):572; 16 June. 

Holotype: USNM 55634.

Type Locality: Mexico: “Tulancingo, Hidalgo, Mexico (altitude, 8,500 feet).” 

Distribution: Mexico: highlands in the states of Hidalgo, 

México, San Luis Potosí, Tamaulipas, and Veracruz, 1,040–

2,900 m (Choate, 1970; Carraway, 2007). 

Remarks: Cryptotis mexicanus group.

Cryptotis oreoryctes Woodman, 2011

2011. Cryptotis oreoryctes Woodman, Zool. J. Linn. Soc. 163(4):1280; 

December.

Holotype: USNM 569877.

Type Locality: Guatemala: “Chelemhá Cloud Forest Reserve (c. 15°23′ N, 90°04′ W), c. 2090 m.a.m.s.l., Alta 

Verapaz.”

Distribution: Guatemala: Sierra de Yalijux, departments of 

Alta Verapaz, El Progreso, and Zacapa; elevational distribution: 2,040–2,700 m (Woodman, 2011a, 2011b, 

2015:119).

Remarks: Cryptotis goodwini group.

Cryptotis orophilus (J. A. Allen, 1895) 

1895. Blarina (Soriciscus) orophila J. A. Allen, Bull. Am. Mus. Nat. Hist. 

7:340; 8 November.

1908. Blarina olivaceus J. A. Allen, Bull. Am. Mus. Nat. Hist. 24:669; 13 October. Holotype: AMNH 28356, from “San Rafael del Norte (altitude 

about 5000 feet), Nicaragua”; Jinotega Department.

1911. C[ryptotis]. orophila: Miller, Proc. Biol. Soc. Wash. 24:221; 31 October. First use of current name combination.

1970. Cryptotis parva orophila: Choate, Univ. Kansas Publ. Mus. Nat. Hist. 

19(3):262; 30 December.

2010. Cryptotis orophilus: Woodman, J. Mamm. 91(3):566; 16 June. 

Holotype: AMNH 9640/9558.

Type Locality: Costa Rica: “Volcan de Irazú” (Cartago Province). 

Distribution: Central America: middle elevations from northern El Salvador and western Honduras to the Valle Central of Costa Rica; elevational distribution: 1,150–1,985 m 

(Woodman and Timm, 1992; Woodman et al., 2012). Panamanian specimens identified by Choate (1970:264) as C.

parvus orophilus were subsequently determined to be C. nigrescens (Woodman and Timm, 1993:21, 30).

Remarks: Cryptotis parvus group.

Cryptotis osgoodi (Stone, 1914)

1914. Blarina osgoodi Stone, Proc. Acad. Nat. Sci. Phila. 66:16; 31 March.

1932. C[ryptotis]. osgoodi: Tate, J. Mamm. 13(3):225; August. First use of 

current name combination.

1958. Cryptotis thomasi equatoris: Cabrera, Rev. Mus. Argentino 4(1):47. 

Part; not Blarina equatoris Thomas, 1912 (=Cryptotis equatoris).

1997. Cryptotis equatoris osgoodi: Vivar et al., Am. Mus. Novit. 3202:7; 

29 August.

Holotype: ANSP 12732.

Type Locality: Ecuador: “Hacienda Garzon, Mt. Pichincha, 

10,500 ft. altitude;” Pichincha Province.

Distribution: Ecuador: eastern Pichincha, eastern Cotopaxi, 

eastern Tungurahua, and western Napo provinces, from the 

lower eastern flanks of Volcán Pichincha through the eastern 

Andean Cordillera to Volcán Sumaco; elevational distribution: 1,700–3,710 m (Moreno-Cárdenas and Albuja, 2014).

Remarks: Cryptotis thomasi group. 

Cryptotis parvus (Say, 1822)

FIGURES 11, 16, 20, 21

Cryptotis parvus floridanus (Merriam, 1895)

FIGURE 20

1837. Sorex cinereus Bachman, J. Acad. Nat. Sci. Phila. 7(2):373. Syntypes: 

ANSP 477, 478; USNM 94/1771 from “Goose Creek about twentytwo miles from Charleston”; Berkeley County, South Carolina (not 

Sorex cinereus Kerr, 1792; see Handley and Varn, 1994).

1837. Sorex cinereous Bachman, J. Acad. Nat. Sci. Phila. 7(2): pl. 23, fig. 3. 

Incorrect subsequent spelling of Sorex cinereus Bachman, 1837.

1895. Blarina floridana Merriam, N. Am. Fauna 10:19; 31 December.

1912. Cryptotis floridana: Miller, Bull. U.S. Natl. Mus. 79:25; 31 December.

1927. Cryptotis parva floridana: Harper, Proc. Boston Soc. Nat. Hist. 38(7): 

270; March. First use of current name combination.
4 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Holotype: USNM 16510/23937.

Type Locality: USA: “Chester Shoal, 11 miles north of Cape 

Canaveral, Brevard County, Fla.” 

Distribution: USA: coastal Carolinas through peninsular Florida (Hall, 1981; Handley and Varn, 1994).

Remarks: Cryptotis parvus group. Sorex cinereus Bachman, 

1837 has typically been placed in synonymy with Cryptotis 

parvus parvus (Say, 1822), but Handley and Varn (1994) 

indicated that on the basis of body size, coastal populations in the Carolinas are more appropriately treated as 

C. parvus floridanus. Some preliminary data suggest, however, that throughout the distribution of C. parvus, coastal 

populations have larger size than nearby inland populations. Rather than a genetically distinctive subspecies, this 

variation, together with latitudinal clinal variation in body 

size, may result in seemingly complex geographic patterns of 

body size variation. 

Cryptotis parvus parvus (Say, 1822) 

FIGURES 11, 16, 21

1822. Sorex parvus Say, in James, Acct. Exped. Pittsb. Rocky Mtns. 1:163; 

31 December.

1842. [Sorex] Brachysorex harlani Duvernoy, Mag. Zool. Anat. Comp. Pal., 

ser. 2, 2:40, pl. 53. Holotype: unknown, from “New-Harmony, dans 

l’Etat d’Indiana (Amérique du Nord)” [Posey County, southwestern 

Indiana]. No specimen matching the holotype was found in a recent 

search of the collection of the MNHN (Cécile Callou, MNHN, email 

personal communication, 1 September 2017).

1857. Blarina exilipes Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):51. Lectotype: USNM 2157, from “Washington, Miss[issippi],” designated by 

Lyon and Osgood, 1909:237.

1857. Blarina eximius Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):52. Syntypes: 

USNM 639/1790 from “St. Louis, M[iss]o[uri].” and 2166/3109 from 

“De Kalb county, Ill[inois].”

1884. Blarina cinerea: True, U.S. Natl. Mus. Circ. 29:22; 29 November. Part; 

not Sorex cinereus Bachman, 1837 = Blarina floridana Merriam, 1895 

(=Cryptotis parvus floridanus).

1912. Cryptotis parva: Miller, Bull. U.S. Natl. Mus. 79:24; 31 December.

1927. [Cryptotis parva] parva: Harper, Proc. Boston Soc. Nat. Hist. 

38(7):270; March. First use of current name combination.

1942. Cryptotis parva elasson Bole and Moulthrop, Sci. Publ. Cleveland 

Mus. Nat. Hist. 5(6):97; 11 September. Holotype: CMNH 14025, from 

“Bettsville, Seneca County, Ohio.”

1942. Cryptotis parva harlani Bole and Moulthrop, Sci. Publ. Cleveland 

Mus. Nat. Hist. 5(6):99.

Holotype: The holotype (Figure 16) in the Peale Museum, 

Philadelphia (see Godman, 1826:79–80 and accompanying 

plate) was lost or destroyed (Woodman, 2009).

Type Locality: USA: “Engineer cantonment . . . taken near 

our cabins” (west bank of the Missouri River, Washington 

County, Nebraska). The location of Engineer Cantonment 

was rediscovered by Carlson et al. (2004) and described by 

Genoways and Ratcliff (2008:6–7): “Engineer Cantonment is 

located in the extreme southeastern corner of modern Washington County, NE, at a place three miles south and four 

FIGURE 21. Sorex harlani Duvernoy, 1842 (=Cryptotis parvus (Say, 1822)). Image from Duvernoy (1842: pl. 53), courtesy of Biodiversity 

Heritage Library (https://www.biodiversitylibrary.org/), contributed by the Ernst Mayer Library, Museum of Comparative Zoology, Harvard 

University, Cambridge.
NUMBER 650 • 4 3

miles east of the town of Fort Calhoun (41°24′43′′ N latitude, 

95°57′00.6′′ W longitude, SE 1⁄4 NW 1⁄4 SE 1⁄4, Section 28, 

Township 17 N, Range 13 E). The site is 305 m in elevation.”

Distribution: Canada and USA: the eastern United States, 

from western South Dakota, east central Wyoming, eastern 

Colorado, and east central New Mexico east through the 

southern tip of Ontario to Connecticut and Florida. There is 

evidence that the species is expanding its distribution to the 

west and to the north (e.g., Backlund, 2002; Geluso et al., 

2004; Seimers et al., 2006). 

Remarks: Cryptotis parvus group. Hall (1981) recognized the 

subspecies C. p. elasson (in Ohio) and C. p. harlani (in portions of eastern Illinois and Indiana). In contrast, Hoffmeister (1989:77) found “no basis for regarding Cryptotis parva 

harlani as a distinct species,” and Whitaker and Mumford 

(2009:158) wrote that “all Indiana specimens should be referred to C. p. parva.” 

Preliminary genetic evidence indicates that C. parvus

may include two cryptic species, possibly distributed to the 

west (C. parvus) and to the east (C. harlani) of the Mississippi River (He et al., 2015).

Cryptotis peregrinus (Merriam, 1895) 

1895. Blarina mexicana peregrina Merriam, N. Am. Fauna 10:24; 31 

December.

1911. C[ryptotis]. mexicana peregrina: Miller, Proc. Biol. Soc. Wash. 

24:222; 31 October.

1999. Cryptotis peregrina: Woodman and Timm, Field. Zool., n.s. 

91(1497):3; 13 January. First use of current name combination.

2010. Cryptotis peregrinus: Woodman, J. Mamm. 91(3):572; 16 June. 

Holotype: USNM 68317.

Type Locality: Mexico: “mountains 15 miles west of city of 

Oaxaca, Mexico (altitude, 9,500 feet).” 

Distribution: Mexico: Sierra de Miahuatlán, southern Oaxaca; 

elevational distribution: 1,060–2,745 m (Woodman and 

Timm, 2000).

Remarks: Cryptotis goldmani group.

Cryptotis perijensis Quiroga-Carmona and Woodman, 2015

2015. Cryptotis perijensis Quiroga-Carmona and Woodman, J. Mamm. 

96(4):804, published online 3 August.

Holotype: MBLUZ 105.

Type Locality: Colombia: “near Finca el Suspiro, Departamento 

del Cesar, Colombia, 2,000 m (10°21′ N, 72°57′ W . . .).”

Distribution: Colombia and Venezuela: “known solely from the 

northern half of the Sierra de Perijá in eastern Colombia and 

western Venezuela” (Quiroga-Carmona and Woodman, 2015).

Remarks: Cryptotis thomasi group.

Cryptotis peruviensis Vivar, Pacheco, and Valqui, 1997

1993. Cryptotis thomasi: Hutterer, MSW2, p. 109. Part; not Blarina thomasi

Merriam, 1897 (=Cryptotis thomasi).

1997. Cryptotis peruviensis Vivar, Pacheco, and Valqui, Am. Mus. Novit. 

3202:7; 29 August.

Holotype: MUSM 8373.

Type Locality: “Peru, Department Cajamarca, Las Ashitas, 

3150 m, about 42 km W of Jaén (05°42′ S, 79°08′ W).”

Distribution: Peru: known only from Cajamarca Department 

in the northern Peruvian Andes (Pacheco et al., 2009; Zeballos et al., 2018). 

Remarks: Cryptotis thomasi group. Cryptotis peruviensis was 

described on the basis of two specimens: the holotype from 

Cajamarca Department (3,150 m) in the northern Peruvian 

Andes and a paratype (LSU 26887) from adjoining Piura 

Department (2,050 m). The population of Cryptotis in 

Piura was subsequently reidentified, first as C. equatoris by 

Pacheco et al. (2009) and then as C. montivagus by Zeballos 

et al. (2018).

Cryptotis phillipsii (Schaldach, 1966)

1966. Notiosorex [Xenosorex] phillipsii Schaldach, Säug. Mitteil. 14(4):289; 

October.

1969. Cryptotis mexicana machetes: Goodwin, Bull. Am. Mus. Nat. Hist. 

141:40; 30 April. Part; not Blarina machetes Merriam, 1895 (=Cryptotis goldmani machetes).

1970. Cryptotis mexicana peregrina: Choate, Univ. Kansas Publ. Mus. Nat.

Hist. 19:237; 30 December. Part; not Blarina peregrina Merriam, 1895 

(=Cryptotis peregrinus).

2000. Cryptotis phillipsii: Woodman and Timm, Proc. Biol. Soc. Wash. 

113(2):351; 31 July. First use of current name combination.

Holotype: CNMA 8445.

Type Locality: Mexico: “Río Molino, 3 kilometers S. W. San 

Miguel Suchixtepec, altitude 2250 meters, southern Oaxaca.”

Distribution: Mexico: southern Oaxaca; elevational distribution: 1,060–2,600 m.

Remarks: Cryptotis mexicanus group. This is the type species 

of subgenus Xenosorex Schaldach, 1966.

Cryptotis pueblensis Jackson, 1933

1933. Cryptotis pergracilis pueblensis Jackson, Proc. Biol. Soc. Wash. 46:79; 

27 April.

1956. Cryptotis celatus Goodwin, Am. Mus. Novit. 1791:1; 28 September. 

Holotype: AMNH 145838, from “Las Cuevas, Santiago Lachiguiri, 

District of Tehuantepec, Oaxaca, México.”

1970. Cryptotis parva pueblensis: Choate, Univ. Kansas Publ. Mus. Nat.

Hist. 19:264; 30 December.

Holotype: USNM 92720.

Type Locality: Mexico: “Huauchinango, altitude 5,000 feet, 

state of Puebla.”

Distribution: Mexico: from the southeastern corner of San Luis 

Potosí southeast to Oaxaca and central Chiapas; elevational 

distribution: 10–2,300 m (Choate, 1970; Carraway, 2007). 

Remarks: Cryptotis parvus group.
4 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Cryptotis soricinus (Merriam, 1895) 

1895. Blarina soricina Merriam, N. Am. Fauna 10:22; 31 December.

1911. C[ryptotis]. soricina: Miller, Proc. Biol. Soc. Wash. 24:221; 31 October. First use of current name combination.

1970. Cryptotis parva soricina: Choate, Univ. Kansas Publ. Mus. Nat. Hist. 

19:267; 30 December.

Holotype: USNM 50762.

Type Locality: Mexico: “Tlalpam, Valley of Mexico (altitude, 

7,600 feet)”; Tlalpan, Distrito Federal. 

Distribution: Mexico: the Valley of Mexico in the Distrito Federal and state of México; elevational distribution: 1,250–

2,370 m (Choate, 1970; Carraway, 2007).

Remarks: Cryptotis parvus group.

Cryptotis squamipes (J. A. Allen, 1912)

1912. Blarina (Cryptotis) squamipes J. A. Allen, Bull. Am. Mus. Nat. Hist. 

31(7):93; 19 April.

1932. C[ryptotis]. squamipes: Tate, J. Mamm. 13(3):225; August. First use of 

current name combination.

Holotype: AMNH 32378.

Type Locality: Colombia: “crest of Western Andes (alt. 10,340 

ft.), 40 miles west of Popayan, Cauca.”

Distribution: Colombia: the southern cordillera and the southern parts of the western and central cordilleras of the Andes; 

elevational distribution: 1,500–3,375 m (Woodman and 

Péfaur, 2008).

Remarks: Cryptotis thomasi group.

Cryptotis tamensis Woodman, 2002

2002. Cryptotis tamensis Woodman, Proc. Biol. Soc. Wash. 115(2):254; 

2 July.

Holotype: USNM 418567.

Type Locality: Venezuela: “state of Táchira: Buena Vista, 7°27′

N, 72°26′ W, 2415 m; near Páramo de Tamá; 35 km S, 22 

km W of San Cristóbal.”

Distribution: Colombia and Venezuela: Tamá highlands in the 

north central portion of the eastern range (in Colombia) 

of the Andes Cordillera, from northeastern Santander and 

southeastern Norte de Santander Departments in Colombia to western Táchira, Venezuela; elevational distribution: 

2,385–3,330 m.

Remarks: Cryptotis thomasi group.

Cryptotis thomasi (Merriam, 1897)

1897. Blarina thomasi Merriam, Proc. Biol. Soc. Wash. 11:227; 15 July.

1904. [Blarina (Cryptotis)] thomasi: Trouessart, Cat. Mamm. Suppl. 

1904:138.

1921. C[ryptotis]. thomasi: Thomas, Ann. Mag. Nat. Hist., 9th ser., 

8(45):354; September. First use of current name combination.

1923. Cryptotis avia G. M. Allen, Proc. New Engl. Zool. Club 8:37; 12 

February. Holotype: MCZ 20091, from “El Verjón, in the Andes east 

of Bogotá, Colombia” [Distrito Capital].

1958. Cryptotis avius: Cabrera, Rev. Mus. Argentino 4(1):47. 

1958. Cryptotis thomasi thomasi: Cabrera, Rev. Mus. Argentino 4(1):48.

Holotype: BM 97.5.21.2.

Type Locality: Colombia: “Plains of Bogota, Colombia (on 

G. O. Child’s estate near City of Bogota, alt. about 9000 

feet)” [Distrito Capital].

Distribution: Colombia: south central portion of the eastern 

range of the Andes Cordillera; elevational distribution: 

2,800–3,500 m.

Remarks: Cryptotis thomasi group.

Cryptotis tropicalis (Merriam, 1895)

FIGURE 22

1843. Corsira tropicalis Gray, Proc. Zool. Soc. Lond. 11:79; December. Type 

locality: “Coban, in Central America”; Alta Verapaz Department, Guatemala (nomen nudum).

1862. Sorex micrurus Tomes, Proc. Zool. Soc. Lond. 1861:279; April. Lectotype: BM 7.1.1.33, designated by Choate (1970:268), but see also 

Handley and Choate (1970); from “Dueñas, Guatemala”; Coban, Alta 

Verapaz Department, Guatemala. Preoccupied by Galemys micrurus

Pomel, 1848 = Sorex talpoides Gapper, 1830 (=Blarina brevicauda 

talpoides).

1877. Blarina micrura: Alston, Proc. Zool. Soc. Lond. 1877:446; 1 October.

1895. Blarina tropicalis Merriam, N. Am. Fauna 10:21; 31 December. Replacement name for Sorex micrurus Tomes, 1862, preoccupied by 

Galemys micrurus Pomel, 1848 (=Sorex talpoides Gapper, 1830 = 

Blarina brevicauda talpoides).

1911. C[ryptotis]. tropicalis: Miller, Proc. Biol. Soc. Wash. 24:221; 31 October. First use of current name combination.

1924. Cryptotis micrura: Miller, Bull. U.S. Natl. Mus. 128:32. Part.

1970. Cryptotis parva soricina: Choate, Univ. Kansas Publ. Mus. Nat.

Hist. 19:268; 30 December. Part; not Blarina soricina Merriam, 1895 

(=Cryptotis soricinus).

1970. Cryptotis parva tropicalis: Choate, Univ. Kansas Publ. Mus. Nat. Hist. 

19:268; 30 December.

Holotype: BM 7.1.1.33.

Type Locality: Guatemala: “Coban, Guatemala (altitude about 

4,400 feet)”; Alta Verapaz Department.

Distribution: Central America: eastern Chiapas through 

Guatemala; elevational distribution: 975–1,580 m (Choate, 1970; Woodman and Timm, 1993; Woodman et al., 

2012). 

Remarks: Cryptotis parvus group. See Handley and Choate 

(1970) for an account of the complex nomenclatural history 

of this species.

Cryptotis venezuelensis Quiroga-Carmona, 2013 

2013. Cryptotis venezuelensis Quiroga-Carmona, Mast. Neotr. 20(1):125; 

23 March.

Holotype: EBRG 27336. 

Type Locality: Venezuela: “Sector Cerro Geremba del Monumento Natural Pico Codazzi, Municipio Tovar, Estado 
NUMBER 650 • 4 5

Aragua, Venezuela (coordenadas geográficas: 10°24′36′′ N, 

67°13′16′′ W), 2238 m de elevación.”

Distribution: Venezuela: cloud forests of the Serranía del Litoral; elevational distribution: 2,100–2,238 m.

Remarks: Cryptotis thomasi group. 

Tribe Notiosoricini Reumer, 1984

1984. Notiosoricini Reumer, Scripta Geologica, 73:18.

Genus Megasorex Hibbard, 1950

1950. Megasorex Hibbard; Contr. Mus. Paleo. Univ. Mich. 8(6):127; 

29 June. 

Type Species: Notiosorex gigas Merriam, 1897, by original 

designation. The genus was reviewed by Hall and Kelson 

(1959) and Hall (1981).

Megasorex gigas (Merriam, 1897) 

1897. Notiosorex gigas Merriam, Proc. Biol. Soc. Wash. 11:227; 15 July.

1950. Megasorex gigas: Hibbard, Contr. Mus. Paleo. Univ. Mich. 8(6):128; 

29 June. First use of current name combination.

Holotype: USNM 88012.

Type Locality: Mexico: “Mts. at Milpillas, near San Sebastian, 

Jalisco.”

Distribution: Mexico: west central states of Nayarit, Jalisco, 

Colima, Michoacán, and Guerrero; elevational distribution: 

80–1,800 m (Carraway, 2007).

Remarks: Megasorex gigas is the type species of the genus 

Megasorex Hibbard, 1950.

Genus Notiosorex Coues, 1877

1877. Notiosorex Coues, Bull. U.S. Geol. Geog. Surv. Terr. 3:646; 15 May. 

Proposed as a subgenus of Sorex. 

Type Species: Sorex (Notiosorex) crawfordi Coues, 1877, by 

original designation.

Remarks: Notiosorex was revised by Merriam (1895a), Carraway and Timm (2000), and Carraway (2007), and it was 

reviewed by Hall and Kelson (1959) and Hall (1981).

Notiosorex cockrumi R. J. Baker, O’Neill, and McAliley, 2003

2003. Notiosorex cockrumi R. J. Baker, O’Neill, and McAliley, Occas. Pap.

Mus. Texas Tech Univ. 222:2; 20 June. 

FIGURE 22. Sorex veraepacis Alston, 1877 (left) and Blarina micrura Alston, 1877 (=Blarina tropicalis Merriam, 1895 = Cryptotis tropicalis) 

(right). Image from Alston (1879–1882: tab. 5), courtesy of Biodiversity Heritage Library (https://www.biodiversitylibrary.org/), contributed by 

Smithsonian Institution Libraries, Washington.
4 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

2003. N[otiosorex]. crockrumi R. J. Baker, O’Neill, and McAliley, Occas.

Pap. Mus. Texas Tech Univ. 222:4; 20 June. Incorrect subsequent spelling of Notiosorex cockrumi R. J. Baker et al., 2003. 

Holotype: TTU 100000.

Type Locality: USA, “Arizona, Cochise County, Leslie Canyon 

National Wildlife Refuge, T21S, R28E, Section NW 1⁄4 20, 

Elevation 4460.” 

Distribution: USA and Mexico: south central and southeastern 

Arizona to central Sonora.

Remarks: This species represents a hypothesis in need of 

comprehensive testing. It was originally identified and described on the basis of differences in the sequences of the 

mitochondrial cytochrome b gene and a portion of intron 

7 of the nuclear beta fibrinogen gene between individuals 

of Notiosorex crawfordi in southern Arizona and Sonora 

and individuals from throughout the rest of the geographic 

range of the species. Neither Notiosorex evotis nor Notiosorex villai was included in the analysis as an outgroup, 

as a gauge of interspecific variation, or to test the distinctiveness of the new species. No morphological or morphometric study was done, making it difficult to identify 

existing specimens in systematic collections, although the 

authors noted that Carraway and Timm (2000) “detected 

no geographically discernable patterns in skin and skull 

characteristics and measurements across the range of 

N. crawfordi” (Baker et al., 2003:4). This cryptic species is 

sympatric with N. crawfordi in Cochise County, Arizona, 

where one might expect to detect morphological character 

displacement between two closely related species. Similar 

molecular data suggest the presence of another morphologically cryptic mitochondrial species in Baja California, 

Mexico (Baker et al., 2003; Ohdachi et al., 2006; McAliley 

et al., 2007).

Notiosorex crawfordi (Coues, 1877) 

1877. Sorex (Notiosorex) crawfordi Coues, Bull. U.S. Geol. Geog. Surv.

Terr. 3:651; 15 May.

1895. Notiosorex crawfordi: Merriam, N. Am. Fauna 10:32; 31 December. 

First use of current name combination.

Holotype: USNM 2653/4437.

Type Locality: USA: “Fort Bliss, New Mexico, or vicinity.” 

This locality is now in the vicinity of the city of El Paso, El 

Paso County, Texas. 

Distribution: USA and Mexico: the southern United States from 

southern California, southern Colorado, southern Kansas, 

and northwestern Arkansas south to northern Jalisco and 

Hidalgo in central Mexico and to the southern tip of Baja 

California; elevational distribution 640–2,040 m (Carraway and Timm, 2000). Notiosorex crawfordi appears to 

be expanding its geographic range northward, southward, 

and eastward (Manning et al., 2014; Rojas-Martínez et al., 

2014; Dreier et al., 2015).

Remarks: Notiosorex crawfordi is the type species of the genus 

Notiosorex Coues, 1877. 

Notiosorex evotis (Coues, 1877) 

1877. Sorex (Notiosorex) evotis Coues, Bull. U.S. Geol. Geog. Surv. Terr. 

3:652; 15 May.

1895. Notiosorex crawfordi evotis: Merriam, N. Am. Fauna 10:34; 

31 December.

1962. Notiosorex evotis: J. K. Jones et al., Univ. Kansas Publ. Mus. Nat.

Hist. 14(12):148; 18 May. First use of current name combination. 

Holotype: USNM 9066.

Type Locality: Mexico: “Mazatlan”; state of Sinaloa. 

Distribution: Mexico: Sinaloa south to Michoacán; elevational 

distribution from near sea level to 550 m (Carraway and 

Timm, 2000).

Notiosorex villai Carraway and Timm, 2000

2000. Notiosorex villai Carraway and Timm, Proc. Biol. Soc. Wash. 

113:307; 6 April. 

Holotype: KU 54932.

Type Locality: Mexico: “Jaumave, Tamaulipas, Mexico, 

2400 ft.”; 23°34′N, 99°23′W (Carraway and Timm, 

(2000).

Distribution: Mexico: known only from the central mountains of Tamaulipas; elevational distribution 575–1,340 m 

(Carraway and Timm, 2000).

Tribe Soricini G. Fischer, 1814

1814. Soricinorum G. Fischer, Zoognosia 3:x. Proposed as a family-level 

taxon.

Genus Sorex Linnaeus, 1758

1758. Sorex Linnaeus, Systema Naturae, 10th ed., 1:53. 

1762. Musaraneus Brisson, Regnum Animale, p. 126. Type species: “Musaraneus” (=Sorex araneus Linnaeus, 1758), by original designation.

1829. Oxyrhin Kaup, Skizzirte Entwickelungs-Geschichte und natürliches System der europäischen Thierwelt, p. 120. Type species: Sorex 

tetragonurus Hermann, 1780 (=Sorex araneus Linnaeus, 1758), designated by Miller (1912b:29).

1835. Amphi-Sorex Duvernoy, Mém. Soc. Mus. d’Hist. Nat. Strasbourg

2:23. Type species: skull of Sorex hermanni Duvernoy, 1835 (=Neomys 

fodiens (Pennant, 1771)) and skin of Sorex araneus tetragonurus Hermann, 1780 (see Hall, 1981:25).

1838. Corsira Gray, Proc. Zool. Soc. Lond. 1837:123; May. Type species: 

Sorex vulgaris Linnaeus, 1754 (=Sorex araneus Linnaeus, 1758; not 

Sorex vulgaris Nathusius, 1838 = Sorex araneus Linnaeus, 1758), by 

monotypy.

1842. Otisorex De Kay, Zool. New York, p. 22 + pl. 5, fig. 1. Type species: 

Otisorex platyrhinus De Kay, 1842 (=Sorex cinereus Kerr, 1792), designated by Baird (1857:9).
NUMBER 650 • 4 7

1848. Hydrogale Pomel, Arch. Sci. Phys. Nat. 9:248; November. Type species: Sorex fimbripes Bachman, 1837, by original designation. The 

type of S. fimbripes is lost, and the species is unidentifiable (Coues, 

1877:641; Hollister, 1911:380). Preoccupied by Hydrogale Kaup, 1829 

(=Neomys Kaup, 1829).

1868. Crocidura: Fitzinger, Sitzungsber. Math.-natürwissensch. Classe Kaiserl. Akad. Wissensch. 57(1):513 (in synonymy). Part; not Crocidura

Wagler, 1832.

1868. Crossopus: Fitzinger, Sitzungsber. Math.-natürwissensch. Classe Kaiserl. Akad. Wissensch. 57(1):513. Part; not Crossopus Wagler, 1832.

1857. Neosorex Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):11; 14 July. Type 

species: Neosorex navigator Baird, 1857, by original designation.

1884. Atophyrax Merriam, Trans. Linn. Soc. New York 2:221; August. Type 

species: Atophyrax bendirii Merriam, 1884, by original designation.

1877. Microsorex Coues, Bull. U.S. Geol. Geog. Surv. Terr. 3(3):646; 

15 May. Type species: Sorex hoyi Baird, 1857, by original designation. 

Proposed as a subgenus of Sorex Linnaeus, 1758.

1890. Homalurus Schulze, Schriften des Naturwissenschaftlichen Vereins des 

Harzes in Wernigerode 5:28. Type species: Sorex alpinus Schinz, 1837, 

by original designation. Proposed as a subspecies of Sorex Linnaeus, 

1758; treated as a genus by Brandes (1896). Preoccupied by Homalura

Meigen, 1826, a genus of Diptera (Palmer, 1904:330).

1911. Sorax Hollister, Proc. U.S. Natl. Mus. 40(1825):378; 17 April. Incorrect subsequent spelling of Sorex Linnaeus, 1758.

1927. Soricidus Altobello, Rev. Franc. Mamm. 1:6. Type species: Soricidus 

monsvairani Altobello, 1927 (=Sorex araneus Linnaeus, 1758), by 

original designation. 

1952. Eurosorex Stroganov, Byull. Moscov. Ovshch. Ispyt. Pryr. Otd. Biol. 

57(5):22. Type species: Sorex buchariensis Ognev, 1921, by original 

designation. Proposed as a subgenus of Sorex Linnaeus, 1758. 

1959. Amphiosorex Hall and Kelson, Mammals of North America, p. 23; 

31 March. Incorrect subsequent spelling of Amphisorex Duvernoy, 

1835.

1967. Ognevia Dolgov and Geptner, in Geptner and Dolgov, Zool. Zh. 

46(9):1422. Type species: Sorex mirabilis Ognev, 1937, by original designation. Proposed as a subgenus of Sorex Linnaeus, 1758.

1989. Strogonovia Yudin, Nasek. Mlekop. Sibiri, p. 121. Type species: Sorex 

daphaenodon Thomas, 1907, by monotypy. Proposed as a subgenus of 

Sorex Linnaeus, 1758.

2012. Sore Hope, Speer, Demboski, Talbot, and Cook, Molecular Phylogenetics and Evolution 64:675. Incorrect subsequent spelling of Sorex

Linnaeus, 1758.

Type Species: Sorex araneus Linnaeus, 1758.

Remarks: North American taxa of Sorex were revised by 

Bachman (1837), Baird (1857), Merriam (1895b), Miller 

(1895), and Jackson (1928), and they were reviewed by 

Hall and Kelson (1959) and Hall (1981). Subfamilies and 

species groups are based on molecular genetic analyses by 

George (1988), Demboski and Cook (2001, 2003), Maldonado et al. (2001, 2004), Shafer and Stewart (2007), Esteva et al. (2010), and Hope et al. (2010, 2012, 2014). My 

interpretation of this body of work is that it indicates the 

existence of three subgenera in North America: Otisorex, 

which is equivalent to Esteva et al.’s (2010) clade B (Sorex

dispar, S. oreopolus, S. veraecrucis, S. cinereus, S. hoyi, 

S. vagrans, S. palustris, S. sonomae, and S. monticola species groups); Sorex (S. arcticus and S. minutissimus species 

groups); and an unnamed subgenus (George, 1988), equivalent to Esteva et al.’s (2010) clade A (S. merriami, S. salvini, S. saussurei, S. trowbridgii, and S. veraepacis species 

groups). See Appendix A.

Sorex albibarbis (Cope, 1862)

1862. Neosorex albibarbis Cope, Proc. Acad. Nat. Sci. Phila. 14:188.

1892. Sorex albibarbis: Merriam, Proc. Biol. Soc. Wash. 7:25; April. First use 

of current name combination.

1903. Sorex palustris albibarbis: Rhoads, The Mammals of Pennsylvania and 

New Jersey, p. 191.

1915. Neosorex palustris acadicus G. M. Allen, Proc. Biol. Soc. Wash. 28:15; 

12 February. Holotype: MCZ BANGS 2046, from “Digby, Nova 

Scotia”; Digby County. Preoccupied by Sorex acadicus Gilpin, 1865 

(=Sorex cinereus Kerr, 1792).

1915. Neosorex palustris albibarbis G. M. Allen, Proc. Biol. Soc. Wash. 

28:17; 12 February.

1926. Sorex palustris gloveralleni Jackson, J. Mamm. 7(1):57; 15 February. 

Replacement name for Neosorex palustris acadicus G. M. Allen, 1915, 

preoccupied by Sorex acadicus Gilpin, 1865. 

1930. Sorex palustris fimbripes: Green, Contr. Mamm. N. Mtn. Reg., p. 11; 

31 March. Part; not Sorex fimbripes Bachman, 1837, an unidentifiable 

species.

1938. Sorex palustris labradorensis Burt, Occ. Pap. Mus. Zool. Univ. Mich. 

383:1; 27 August. Holotype: UMMZ 68109, from “Red Bay, Straits of 

Belle Isle, Labrador”; Newfoundland and Labrador. 

1942. Sorex palustris punctulatus Hooper, Occ. Pap. Mus. Zool. Univ. Mich. 

463:1; 15 September. Holotype: UMMZ 85498, from “West Virginia, 

Randolph County, six miles northwest of Durbin, Shavers Fork of the 

Cheat River, 3600 feet elevation.”

1951. Sorex palustris turneri D. H. Johnson, Proc. Biol. Soc. Wash. 64:110; 

24 August. Holotype: USNM 282879, from “Fort Chimo (on the eastern bank of Koksoak River, lat. 58°8′ N., long. 68°15′ W.), Ungava 

district, Quebec, Canada.”

Syntype: USNM 11239/38743. Cope (1862) described S. albibarbis from two specimens but did not designate a holotype. The location of the second specimen, if it still exists, 

is unknown.

Type Locality: USA: “Profile Lake, in the Franconia Mountains, New Hampshire”; Grafton County.

Distribution: Canada and USA: eastern Ontario through southern Quebec to the Atlantic Coast; north to Ungava Bay and 

south to central Pennsylvania.

Remarks: Subgenus Otisorex; Sorex palustris group. Hope et 

al. (2014) recognized three species-level molecular clades 

among the subspecies formerly united as Sorex palustris:

eastern S. albibarbis, cordilleran S. navigator, and boreal 

S. palustris. Their study recognized S. albibarbis as comprising the former subspecies S. palustris albibarbis, S. p.
4 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

gloveralleni, S. p. labradorensis, S. p. punctulatus, and S. p.

turneri. The latter two subspecies were included, however, 

on the basis of geography alone because no sequences from 

them were analyzed (Hope et al., 2014). 

Green (1930:11) noted that Neosorex albibarbis Cope, 

1862 could be antedated by Sorex fimbripes Bachman, 

1837. Unfortunately, the type of S. fimbripes is lost, and the 

species is unidentifiable (Hollister, 1911:380). 

Sorex altoensis Carraway, 2007

2007. Sorex veraecrucis altoensis Carraway, Monogr. West. N. Am. Nat. 

3:58; 22 June.

2010. S[orex]. veraecrucis (5): Esteva et al., Zootaxa 2615:55, fig. 1; 17 September. Part; not Sorex veraecrucis Jackson, 1925.

2010. S[orex]. veraecrucis (7): Esteva et al., Zootaxa 2615:55, fig. 1; 17 September. Part; not Sorex veraecrucis Jackson, 1925.

2014. Sorex salvini altoensis: Ramírez-Pulido et al., Spec. Publ. Mus. Texas 

Tech Univ. 63:6; 18 September.

Holotype: KU 107135.

Type Locality: Mexico: “‘Volcán de Fuego, 9800 ft., Jalisco,’ 

latitude 19.55° N, longitude 103.63° W.” 

Distribution: Mexico: highlands in western Durango, southern Coahuila, Nuevo León, and southwestern Tamaulipas, 

Jalisco, Guanajuato, Querétaro, Hidalgo, Colima, Michoacán, México, Morelos, Distrito Federal, Puebla, Guerrero, and Oaxaca; elevational distribution: 1,820–3,500 m 

(Carraway, 2007).

Remarks: Unnamed subgenus; Sorex saussurei group; part of Esteva et al.’s (2010) clade B1 (as S. veraecrucis localities 5, 7), 

in which it is sister to S. saussurei (as S. saussurei locality 6).

Sorex arcticus Kerr, 1772

FIGURES 4, 23

Remarks: Subgenus Sorex; Sorex araneus group (Fumagalli et 

al., 1999). 

Sorex arcticus arcticus Kerr, 1772

FIGURE 4

1792. Sorex arcticus Kerr, Animal Kingdom, p. 206.

1829. Sorex parvus: Richardson, Fauna Bor.-Am., p. 8. Based on MZSL 89, 

listed in Waterhouse (1838:18) as specimen 160. Part; not Sorex parvus

Say, 1822 (=Cryptotis parvus). 

1837. Sorex Richardsonii Bachman, J. Acad. Nat. Sci. Phila. 7(2):383; 

November. Lectotype: BM 1855.12.24.92 (previously MZSL 89), 

from “North America”; designated by Jackson (1928:68) on the 

basis of information from Oldfield Thomas. Bachman (1837) based 

his description of Sorex richardsonii on Richardson’s (1829) description of Sorex parvus, which was based on MZSL 89, listed in 

Waterhouse (1838:18) as specimen 160. Neither Bachman (1837) 

nor Richardson (1829) provided a locality, and information associated with the specimen indicates only that it is from North America. 

Waterhouse (1838:18) similarly states only that it is from “North 

America.” Merriam (1895b:63) listed the type locality as “unknown; probably plains of Saskatchewan.”

1843. S[orex]. Richardsoni: Sundevall, Kung. Svenska Vet. Hand., ser. 3, 

30(1842):182. Incorrect subsequent spelling of Sorex richardsonii

Bachman, 1837.

1877. Sorex sphagnicola Coues, Bull. U.S. Geol. Geog. Surv. Terr. 3:650; 

15 May. Holotype: USNM 6361, from “Fort Liard, H[udson’s]. B[ay]. 

T[erritory]., or vicinity,” Dehcho Region, southwestern Northwest Territories, Canada.

1890. Sorex vulgaris: Dobson, Monogr. Insectiv., pt. 3, fasc. 1, pl. 23, fig. 4. 

Part; not Sorex vulgaris Linnaeus, 1754 or Sorex vulgaris Nathusius, 

1838.

1892. Sorex belli Merriam, Proc. Biol. Soc. Wash. 7:25; 13 April. Holotype: 

CMN 46, from “Shamattawa River, Kenora District, Ontario.” Nomen 

nudum based on an 1885 manuscript name by Dobson (see Jackson, 

1928:69; Youngman, 1973:1).

1892. [Sorex] sphagnicolus: Merriam, Proc. Biol. Soc. Wash. 7:25; April. 

Unjustified emendation of Sorex sphagnicola Coues, 1877. 

1925. Sorex arcticus arcticus Jackson, Proc. Biol. Soc. Wash. 38:127; 13 November. First use of current name combination.

1983. S[orex]. arctucus: Okhotina, Zool. Zh. 62(3):417; March. Incorrect 

subsequent spelling of Sorex arcticus Kerr, 1792.

Holotype: “None known to exist” (Jackson, 1928:69). The 

holotype was originally part of the collection of the Royal 

Society of London (Forster, 1772), which was transferred 

to the British Museum (Natural History), now The Natural 

History Museum, London, in 1781 (Chambers, 2016:23). 

Kerr (1792) based his “Labradore shrew,” Sorex arcticus, 

mainly on Pennant’s (1784:139) description of a specimen 

of the “foetid shrew.” This was the same specimen Forster 

(1772:380) reported as the foetid shrew, Sorex araneus (see 

Jackson, 1928:69–70). 

Type Locality: Canada: “Inhabits Hudson’s Bay and Labradore.” An editor’s footnote to Forster’s (1772:370) account of the quadrupeds from Hudson’s Bay indicates 

that the specimens on which he based his report were sent 

from the Hudson’s Bay Company factory at Hudson Bay 

by Mr. Graham, “a gentlemen belonging to the settlement 

on Severn River.” On the basis of this information, Jackson (1925:55) restricted the type locality of S. a. arcticus

to “Severn Settlement (now Fort Severn, mouth of Severn 

River, Ontario)” (see also Jackson, 1928:69). The specimens 

documented by Forster (1772) came from a variety of localities, however, including “Severn River,” “Albany Fort” (on 

the Albany River), and “Churchill River”—all near Hudson 

Bay—but also from “Musquash [River]” and “York Fork” 

in southern Ontario. Unfortunately, Forster (1772) did not 

provide a specific locality for his foetid shrew, so it may have 

been obtained from anywhere in the region of east central to 

southeastern Ontario.

Distribution: Canada and USA: southeastern Yukon Territory and southwestern Northwest Territories south through 

northwestern corner of British Columbia, Alberta, and Saskatchewan to northwestern North Dakota and east through 

Manitoba, Ontario, and central Quebec to the west bank of 

the St. Lawrence River (Hall, 1981; Naughton, 2012).
NUMBER 650 • 4 9

Sorex arcticus laricorum Jackson, 1925

FIGURE 23

1857. Sorex pachyurus Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):20, pl. 27; 

14 July. Lectotype USNM 1674/38820 from “Pembina, Minnesota 

[Territory]”; this is now in Pembina County, North Dakota, USA. 

Designated by Lyon and Osgood (1909:247). Preoccupied by Sorex 

pachyurus Küster, 1835 (=Sorex etruscus Savi, 1822 = Suncus etruscus) 

(Contoli et al., 2004; Hutterer, 2005).

1925. Sorex arcticus laricorum Jackson, Proc. Biol. Soc. Wash. 38:127; 13 

November.

Holotype: USNM 186837.

Type Locality: USA: “Elk River, Sherburne County, Minnesota.” 

Distribution: Canada and USA: southern Manitoba, eastern 

North Dakota and northeastern South Dakota east through 

Wisconsin and the northern peninsula of Michigan (Hall, 

1981; Naughton, 2012).

Sorex arizonae Diersing and Hoffmeister, 1977

1977. Sorex arizonae Diersing and Hoffmeister, J. Mamm. 58(3):329; 

20 August.

Holotype: MSB 275328 (originally UIMNH 3809).

Type Locality: USA: “upper end of Miller Canyon, 15 mi S 

[=10 mi S, 43⁄4 mi E] Fort Huachuca [near spring at lower 

edge of Douglas fir zone, Huachuca Mts.], Cochise County, 

Arizona” (brackets in the original).

Distribution: USA and Mexico: disjunct mountains in SE Arizona, SW New Mexico, and the Sierra Madre Occidental of 

Chihuahua (Carraway, 2007).

Remarks: Unnamed subgenus (George, 1988); Sorex merriami

group. Originally referred to subgenus Sorex by Diersing 

and Hoffmeister (1977).

Sorex bendirii (Merriam, 1884)

Remarks: Subgenus Otisorex; Sorex palustris group.

Sorex bendirii albiventer Merriam, 1895

1895. Sorex (Atophyrax) bendirii albiventer Merriam, N. Am. Fauna 10:97; 

31 December.

1901. [Atophyrax bendirii] albiventer: Elliot, Field Col. Mus. Publ. 45, Zool.

Ser. 2(1):381; 6 March.

1912. Neosorex bendirii albiventer: Miller, Bull. U.S. Natl. Mus. 79:22; 31 

December.

1926. Sorex bendirii albiventer: Jackson, J. Mamm. 7:58; 15 February. First 

use of current name combination. 

Holotype: USNM 66198.

Type Locality: USA: “Lake Cushman, Olympic Mountains, 

Washington”; Mason County.

Distribution: USA: Olympic Peninsula of Washington (Hall, 1981).

Sorex bendirii bendirii (Merriam, 1884) 

1884. Atophyrax bendirii Merriam, Trans. Linn. Soc. New York 2:217; 28 

August.

1884. Atophyrax Bendirei: True, U.S. Natl. Mus. Circ. 29:22; 29 November. 

Incorrect subsequent spelling of Atophyrax bendirii Merriam, 1884 

(=Sorex bendirii).

1890. Sorex bendirii: Dobson, Monogr. Insectiv., pt. 3, pl. 23, fig. 17 and 

legend.

1904. [Sorex (Atophyrax)] bendiri: Trouessart, Cat. Mamm. Suppl. 1:135. 

Incorrect subsequent spelling of Atophyrax bendirii Merriam, 1884 

(=Sorex bendirii).

1906. Sorex bendirei: Stephens, California Mammals, p. 255; June. Incorrect subsequent spelling of Atophyrax bendirii Merriam, 1884 (=Sorex 

bendirii).

1912. Neosorex bendirii bendirii: Miller, Bull. U.S. Natl. Mus. 79:22; 31 

December.

1926. Sorex bendirii bendirii: Jackson, J. Mamm. 7:58; 15 February. First use 

of current name combination. 

Holotype: USNM 186442 (by monotypy; Fisher and Ludwig, 

2015).

Type Locality: USA: “about a mile from Williamson’s River, 

and some eighteen miles southeast of Fort Klamath Basin”; 

Klamath County, Oregon. 

Distribution: Canada and USA: extreme southwestern British Columbia south through the Cascades and lowlands 

of western Washington (excluding the Olympic Peninsula), 

west central Oregon to coastal California as far south as 

southern Mendocino County (Hall, 1981; Verts and Carraway, 1998).

Sorex bendirii palmeri Merriam, 1895

1895. Sorex (Atophyrax) bendirii palmeri Merriam, N. Am. Fauna 10:97; 

31 December.

1901. [Atophyrax bendirii] palmeri: Elliot, Field Col. Mus. Publ. 45, Zool.

Ser. 2(1):381; 6 March.

1912. Neosorex bendirii palmeri: Miller, Bull. U.S. Natl. Mus. 79:22; 31 

December.

1926. Sorex bendirii palmeri: Jackson, J. Mamm. 7:58; 15 February. First use 

of current name combination. 

Holotype: USNM 17338/24263.

Type Locality: USA: “Astoria, Oregon”; Clatsop County. 

Distribution: USA: Oregon west of the Cascade Mountains 

to extreme northwestern California, north of the Klamath 

River (Hall, 1981).

Sorex chiapensis Jackson, 1925

1925. Sorex veraepacis chiapensis Jackson, Proc. Biol. Soc. Wash. 38:129; 

13 November.

2012. Sorex veraepacis veraepacis: Woodman et al., Ann. Carnegie Mus. 

80(3):217; 15 July. Part; not Sorex veraepacis Alston, 1877.

2017. Sorex chiapensis: Matson and Ordóñez-Garza, Zootaxa 4236(3):477; 

24 February. First use of current name combination.

Holotype: USNM 75877.

Type Locality: Mexico: “San Cristobal, altitude 9,500 feet, 

state of Chiapas.”
5 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 23. Spencer Baird’s shrews: from top to bottom, USNM 1686, Sorex thompsoni Baird, 1857 (=Sorex 

hoyi thompsoni ); USNM 1685, Sorex haydeni Baird, 1857; Sorex suckleyi Baird, 1857 (=Sorex vagrans Baird, 

1857); USNM 1674, Sorex pachyurus Baird, 1857 (=Sorex arcticus laricorum Jackson, 1925). Image from Baird 

(1857: pl. 17).
NUMBER 650 • 5 1

Distribution: Mexico and Guatemala: highlands in Chiapas, 

Mexico, and in adjacent northwestern Guatemala, 2,895–

3,160 m (Carraway, 2007; Matson and Ordóñez-Garza, 

2017).

Remarks: Subgenus Otisorex; Sorex veraepacis group. 

Sorex cinereus Kerr, 1792

FIGURES 4, 11, 24–27 

Remarks: Type species of subgenus Otisorex; southern clade 

of the Sorex cinereus group (Hope et al., 2012); clade A2 of 

Esteva et al. (2010). 

Molecular analyses of the Sorex cinereus group (Demboski and Cook, 2003; Hope et al., 2012) have provided 

intriguing insight into relationships among species and populations, including low genetic divergence among a number 

of recognized arctic species and the presence of unrecognized cryptic species. As currently understood, S. cinereus

is almost certainly paraphyletic (Stewart et al., 1993; Hope 

et al., 2012). Unfortunately, sampling for genetic studies remains insufficient to completely elucidate the relationships 

of genetic clades to named subspecies, the geographic distributions of clades, or the taxonomy of the S. cinereus group. 

Sorex cinereus cinereus Kerr, 1792

FIGURES 4, 11, 24–26 

1792. Sorex arcticus cinereus Kerr, Animal Kingdom, p. 206.

1827. Sorex personatus I. Geoffroy Saint-Hilaire, Dict. Class. d’Hist. Nat. 

11:319; January. Holotype: unknown, from the “États-Unis.” According to I. Geoffroy Saint-Hilaire (1827a, 1827b), the single specimen of 

the species then in the Muséum National d’Histoire Naturelle, Paris 

(MNHN), was obtained by Jacques-Gérard Milbert (1766–1840), a 

French artist who lived in the United States from 1815 to 1823. Beginning about 1817, Milbert collected animals, plants, minerals, and rocks 

for the MNHN and Jardin des Plantes. Although he resided mostly in 

New York City, Milbert traveled extensively in the middle Atlantic and 

northeastern states, going as far south as Virginia and visiting Maryland, Pennsylvania, New Jersey, and much of New England as well as 

upstate New York. Milbert eventually sent 58 shipments containing 

7,868 specimens back to France (Weimerskirch, 1998). The holotype 

of Sorex personatus most likely would have been obtained somewhere 

in the northeastern to Mid-Atlantic regions of the United States. No 

specimen matching the holotype was found in a recent search of the 

collection of the MNHN (Cécile Callou, MNHN, email communication, 1 September 2017).

1827. Sorex personatus: I. Geoffroy Saint-Hilaire, Mém. Mus. Hist. Nat. 

15:122; December. This work is often incorrectly cited as the original 

description of the species. 

1828. Sorex forsteri Richardson, Zool. J. 3:516; April. Holotype: BM 

1842.10.7.2, from “Capt. Franklin’s late Expedition,” “common 

throughout the Hudson’s Bay countries.” John Richardson participated 

in John Franklin’s 1825 Canadian expedition, which traveled north 

down the Mackenzie River, then east (Richardson) and west (Franklin) along the coast of the Beaufort Sea. The expedition thereby passed 

through portions of the Northwest Territories, Yukon Territory, Nunavut and Alaska (Figures 4, 24).

1837. Sorex cooperi Bachman, J. Acad. Nat. Sci. Phila. 7(2):388. Holotype: not known to exist; from “the North Western Territory.” The 

“Northwest Territory” or “Territory Northwest of the Ohio River” 

was incorporated as a U.S. territory in 1787 and encompassed what 

are now the states of Ohio (statehood in 1803), Indiana (1816), Illinois (1818), Michigan (1837), Wisconsin (1848), and parts of Minnesota (1858). Once Ohio achieved statehood, the territory no longer 

existed officially, which may be why Merriam (1895b:60) speculated 

that the holotype was “probably from northern part of Mississippi 

Valley” (Figures 4, 25).

1837. Corsira Forsteri: Gray, Proc. Zool. Soc. London 5(1837):124.

1842. Otisorex platyrhinus De Kay, Zool. New York, 1:22, pl. 5, fig. 1. Holotype: unknown, from “Tappan, Rockland county,” New York.

1842. Otisorex platyrhincus De Kay, Zool. New York, 1:143. Incorrect subsequent spelling of Otisorex platyrhinus De Kay, 1842.

1842. S[orex]. platyrrhinchus Linsley, Am. J. Sci. 43(2):346; October. Incorrect subsequent spelling of Otisorex platyrhinus De Kay, 1842. 

1855. S[orex]. platyrrhinus Wagner, Die Säugthiere Suppl. 5:547. Incorrect 

subsequent spelling of Otisorex platyrhinus De Kay, 1842. 

1857. Sorex platyrhinus: Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):25.

1865. Sorex, Acadicus Gilpin, Proc. Trans. Nova Scot. Inst. Nat. Sci. 1(2):2. 

Neotype: USNM 59647, designated by Woodman (2018) from “Canada: Nova Scotia, Halifax County, Halifax.” The date for the publication of the description of this species is typically given as 1867, but see 

Woodman (2018).

1866. Sorex fosteri Packard, Proc. Boston Soc. Nat. Hist. 10:244. Incorrect 

subsequent spelling of Sorex forsteri Richardson, 1828.

1868. Amphisorex forsteri: Fitzinger, Sitzungsber. Math.-natürwissensch.

Classe Kaiserl. Akad. Wissensch. 57(1):509 (in synonymy).

1868. Crocidura cooperi: Fitzinger, Sitzungsber. Math.-natürwissensch.

Classe Kaiserl. Akad. Wissensch. 57(1):513 (in synonymy).

1868. Otisorex platyrrhinus Fitzinger, Sitzungsber. Math.-natürwissensch.

Classe Kaiserl. Akad. Wissensch. 57(1):584 (in synonymy). Incorrect 

subsequent spelling of Otisorex platyrhinus De Kay, 1842.

1868. Sorex platyrhinchus Fitzinger, Sitzungsber. Math.-natürwissensch.

Classe Kaiserl. Akad. Wissensch. 57(1):584 (in synonymy). Incorrect 

subsequent spelling of Otisorex platyrhinus De Kay, 1842.

1868. Crocidura platyrhyimncha Fitzinger, Sitzungsber. Math.-

natürwissensch. Classe Kaiserl. Akad. Wissensch. 57(1):585 (in 

synonymy). Incorrect subsequent spelling of Otisorex platyrhinus

De Kay, 1842.

1869. Sorex platyrinus Gilpin, Proc. Trans. Nova Scot. Inst. Nat. Sci. 2(2):59. 

Incorrect subsequent spelling of Otisorex platyrhinus De Kay, 1842.

1869. Sorex Acadica: Gilpin, Proc. Trans. Nova Scot. Inst. Nat. Sci. 2(2):59. 

Unjustified emendation of Sorex acadicus Gilpin, 1865.

1890. Sorex richardsoni Dobson, Monogr. Insectiv., pt. 3, fasc. 1, pl. 23, fig. 

9. Part; not Sorex richardsonii Bachman, 1837 (=Sorex arcticus Kerr, 

1772).

1891. Sorex idahoensis Merriam, N. Am. Fauna 5:32; 30 July. Holotype: 

USNM 23527/30945, from “Timber Creek, Salmon River Mountains, 

Idaho, . . . Altitude about 2,500 meters (8,200 feet)”; Lemhi Mountains, Lemhi County.

1901. [Sorex] platyrhnchus Elliot, Field Col. Mus. Publ. 45, Zool. Ser. 

2(1):366; 6 March. Incorrect subsequent spelling of Otisorex platyrhinus De Kay, 1842.
5 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1913. Sorex foresteri Fleming, in J. H. Faull, Natural History of the Toronto 

Region (Toronto: The Canadian Institute), p. 209; November. Incorrect 

subsequent spelling of Sorex forsteri Richardson, 1828.

1925. Sorex cinereus cinereus Jackson, J. Mamm. 6(1):56; 9 February. First 

use of current name combination.

1926. Sorex frankstounensis Peterson, Ann. Carnegie Mus. 16(2):292; 

March. Holotype: CMVP 11159a, a Pleistocene (Rancholabrean) 

fossil from “Frankstown Cave, near Hollidaysburg, Blair County, 

Pennsylvania.”

1940. Sorex cinereus acadicus: R. W. Smith, Am. Midl. Nat. 24(1):219; 

31 July.

1942. Sorex cinereus ohionensis Bole and Moulthrop, Sci. Publ. Cleveland 

Mus. Nat. Hist. 5(6):89; 11 September. Holotype: CMNH 16901, from 

“Hunting Valley, Cuyahoga County, Ohio,” USA.

Holotype: “None now known to exist” (Jackson, 1928:41). The 

holotype was originally part of the collection of the Royal 

Society of London (Forster, 1772), which was transferred 

to the British Museum (Natural History), now The Natural 

History Museum, London, in 1781 (Chambers, 2016:23). 

Kerr (1792) based his description of the “grey Labradore 

shrew,” Sorex arcticus cinereus, on Pennant’s (1784:139) 

description of a smaller variety of what he called the “foetid 

shrew,” which, in turn, was based on Forster’s (1772:381) 

description of two specimens sent to the Royal Society by 

Andrew Graham, “long a resident in Hudson’s Bay” (Pennant, 1784: fifth page of unpaginated “Advertisement”). 

Type Locality: Canada: “Hudson’s Bay.” An editor’s footnote 

to Forster’s (1772:370) account of the quadrupeds from 

Hudson’s Bay indicates that the specimens on which he 

based his report were sent from the Hudson’s Bay Company 

factory at Hudson Bay by Mr. Graham, “a gentlemen belonging to the settlement on Severn River.” On the basis of 

this information, Jackson (1925:55) restricted the type locality of S. c. cinereus to: “Severn Settlement,” which he previously noted was “now Fort Severn, mouth of Severn River, 

FIGURE 24. Sorex forsteri Richardson, 1828 (=Sorex cinereus Kerr, 1792). Image from Gapper (1830: pl. 7), courtesy of Biodiversity Heritage 

Library (https://www.biodiversitylibrary.org/), contributed by The Natural History Museum Library, London.
NUMBER 650 • 5 3

FIGURE 25. Spencer Baird’s shrews: from top to bottom, USNM 2047, Sorex cooperi Bachman, 1837 (=Sorex 

cinereus Kerr, 1792); USNM 1675, Sorex vagrans Baird, 1857; Sorex trowbridgii Baird, 1857; USNM 629/1780 

(not 629/1760), Neosorex navigator Baird, 1857 (=Sorex navigator). Image from Baird (1857: pl. 16).
5 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ontario,” Canada (see also Jackson, 1928:41). The various 

species noted by Forster came, however, from a variety of 

localities, including “Severn River,” “Albany Fort” (on the 

Albany River), and “Churchill River”—all near Hudson 

Bay—and also from “Musquash [River]” and “York Fork” 

in southern Ontario. Unfortunately, Forster (1772) did not 

provide a specific locality for these two shrews, so they may 

have been obtained from anywhere in the region, from east 

central to southeastern Ontario.

Distribution: Canada and USA: In western North America,

S. c. cinereus occurs from north central Alaska, northern 

Yukon Territory, northern Northwest Territories, central 

and southeastern Nunavut, and the southern edge of Hudson’s Bay south to central Washington (but excluding most 

of the coastal region), west central and southeastern Idaho, 

south central Utah (but not eastern Utah or western Colorado), central Colorado, north central New Mexico, southern and north central Wyoming, central Montana, across 

central Saskatchewan, parts of southern Manitoba, to central and southern Minnesota. In eastern North America, it 

occupies most of western and central Quebec (except the 

Ungava Peninsula) and parts of westernmost Labrador 

south through New England (excluding Nova Scotia) to 

central New Jersey and Pennsylvania in and west of the Appalachians, south along the Appalachians to northern Georgia, southern Ohio, southern Indiana, to central Illinois 

(Hall, 1981); elevational distribution from near sea level to 

3,625 m (USNM specimens).

Remarks: On the basis of multigene molecular analysis, S. c.

cinereus as currently conceived comprises populations from 

the West, Midwest, Southwest, and East molecular clades 

of Hope et al. (2012) and includes at least two undescribed 

species. It also appears to be paraphyletic with respect to 

S. milleri.

Sorex cinereus acadicus in Nova Scotia and S. c. ohioensis in Ohio have generally been recognized as valid taxa 

(e.g., Hall, 1981; Whitaker, 2004; Hutterer, 2005). Sorex

c. acadicus differs insubstantially from adjacent populations 

of S. c. cinereus, and I recently placed the former in synonymy with the latter (Woodman, 2018). Van Zyll de Jong 

and Kirkland (1989) considered S. c. ohioensis morphologically and morphometrically transitional between S. c. cinereus and S. c. lesueurii. In fact, a review of their evidence 

suggests it is little differentiated at all from S. c. cinereus, 

and for this reason, I synonymize S. c. ohioensis with S. c. 

cinereus herein.

FIGURE 26. Otisorex platyrhinus De Kay, 1842 (=Sorex cinereus Kerr, 1792). Image from De Kay (1842: pl. 5, fig. 1), courtesy of Biodiversity 

Heritage Library (https://www.biodiversitylibrary.org/), contributed by the Marine Biology Laboratory, Woods Hole Oceanographic Institution, Woods Hole.
NUMBER 650 • 5 5

Sorex cinereus hollisteri Jackson, 1925

1900. Sorex personatus arcticus Merriam, Proc. Wash. Acad. Sci. 2:17; 14 

March. Preoccupied by Sorex arcticus Kerr, 1792.

1925. Sorex cinereus hollisteri Jackson, J. Mamm. 6(1):55; 9 February. Replacement name for Sorex personatus arcticus Merriam, 1900, preoccupied by Sorex arcticus Kerr, 1792.

Holotype: USNM 99305.

Type Locality: USA: “St. Michaels, Alaska.” 

Distribution: USA: western Alaska from Franklin Point south 

to Anchorage (Miller and Kellogg, 1955; Hall, 1981). 

Remarks: On the basis of multigene molecular analysis, S. c.

hollisteri comprises a part of the West clade of S. cinereus

(Hope et al., 2012).

Sorex cinereus lesueurii Duvernoy, 1842

FIGURE 27

1842. [Sorex] Amphisorex lesueurii Duvernoy, Mag. Zool. Anat. Comp. Pal. 

25:33, pl. 50; November.

1843. S[orex]. lesueurii: Sundevall, Kung. Svenska Vet. Hand., ser. 3, 

30(1842):182.

1847. [Sorex] lesneurii A. B. Reichenbach, Praktische Naturgeschichte Menschen und der Säugethiere für Gebildete aller Stände (Leipzig: Gebhardt 

and Reisland), p. 165 [reference not seen]. Incorrect subsequent spelling 

of Sorex lesueurii Duvernoy, 1842.

1857. Amphisorex lesuerii Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):27 (in 

synonymy). Incorrect subsequent spelling of Sorex lesueurii Duvernoy, 

1842.

1857. Sorex lesueri Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):27 (in synonymy). Incorrect subsequent spelling of Sorex lesueurii Duvernoy, 1842.

1892. Amphisorex lesueri Herrick, Bull. Geol. Nat. Hist. Surv. 7:48. Incorrect subsequent spelling of Sorex lesueurii Duvernoy, 1842.

1892. Amphisorex leseurii Butler, Proc. Indiana Acad. Sci. 1891:163. Incorrect subsequent spelling of Sorex lesueurii Duvernoy, 1842.

1895. Amphisorex leseueri Miller, N. Am. Fauna 10:53; 31 December. Incorrect subsequent spelling of Sorex lesueurii Duvernoy, 1842.

1895. Amphisorex lesueuri Merriam, N. Am. Fauna 10:60; 31 December (in 

synonymy). Incorrect subsequent spelling of Sorex lesueurii Duvernoy, 

1842.

1895. [Sorex personatus] lesueuri Merriam, N. Am. Fauna 10:61; 31 December. Incorrect subsequent spelling of Sorex lesueurii Duvernoy, 1842.

1901. Sorex personatus lesueurii: Miller and Rehn, Proc. Boston Soc. Nat.

Hist. 30:25; 27 December.

1911. Sorex longirostris lesueurii: Hollister, Proc. U.S. Natl. Mus. 40:380; 

17 April.

1942. Sorex cinereus lesueurii Bole and Moulthrop, Sci. Publ. Cleveland 

Mus. Nat. Hist. 5(6):95; 11 September. First use of current name 

combination. 

Holotype: None known to exist (Diersing and Hoffmeister, 

1981). No specimen matching the type of Amphisorex 

lesueurii Duvernoy, 1842 was found in a recent search of 

the collection of the MNHN (Cécile Callou, MNHN, email 

communication, 1 September 2017).

Type Locality: USA: “la vallée de la rivière de Wabasch, qui arrose l’Indiana, l’un des États unis de l’Amérique septrentrionale” [the valley of the Wabash River, which waters Indiana, 

one of the United States of North America]; restricted to 

“New Harmony, Posey County, Indiana,” by Diersing and 

Hoffmeister (1981). 

Distribution: USA: Diersing and Hoffmeister (1981:6) considered the distribution of S. c. lesueurii to be mostly restricted 

to the lower Wabash, Ohio, and Mississippi river valleys 

in southern and southeastern Illinois and southwestern Indiana, possibly extending into adjacent parts of the Mississippi River Valley in Kentucky, Missouri, Tennessee, and 

Arkansas. In contrast, Whitaker (2004) indicated a more 

widespread distribution that also included portions of 

southern Wisconsin, northern Illinois, northern and west 

central Indiana, and possibly the southern half of the lower 

peninsula of Michigan.

Remarks: The taxonomic status of this subspecies has not been 

evaluated using molecular phylogenetic techniques (e.g., 

Hope et al., 2012).

Sorex cinereus miscix Bangs, 1899

1899. Sorex personatus miscix Bangs, Proc. New Engl. Zool. Club 1:15; 28 

February.

1901. [Sorex merriami] miscix: Elliot, Field Col. Mus. Publ. 45, Zool. Ser. 

2(1):376; 6 March.

1925. Sorex cinereus miscix: Jackson, J. Mamm. 6:56; February. First use of 

current name combination.

Holotype: MCZ BANGS 8651.

Type Locality: Canada: “Black Bay, Labrador”; Newfoundland and Labrador Province.

Distribution: Canada: eastern Labrador and eastern Quebec to 

the north shore of Gulf of St. Lawrence at Moisie River and 

Bay of Seven Islands (Miller and Kellogg, 1955; Hall, 1981).

Remarks: On the basis of multigene molecular analysis, S. c. miscix is a member of the Midwest clade of S. cinereus, which is 

otherwise composed of populations from the upper Midwest, 

upper Great Plains, and Rocky Mountains (Hope et al., 2012).

Sorex cinereus nigriculus Green, 1932

1932. Sorex cinereus nigriculus Green, Univ. Calif. Publ. Zool. 38(7):387; 

9 June. 

Holotype: MVZ 51413.

Type Locality: USA: “alluvial tidewater marsh on Tuckahoe 

River, east of Tuckahoe, Cape May County, southern New 

Jersey.”

Distribution: USA: this is an isolated subspecies known only 

from the vicinity of the type locality (Hall, 1981; Whitaker, 

2004).

Remarks: The taxonomic status of this subspecies has not 

been evaluated using molecular phylogenetic techniques 

(e.g., Hope et al., 2012).
5 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sorex cinereus streatori Merriam, 1895

1895. Sorex personatus streatori Merriam, N. Am. Fauna 10:62; 

31 December.

1925. Sorex cinereus streatori: Jackson, J. Mamm. 6(1):56; 9 February. First 

use of current name combination.

Holotype: USNM 73537.

Type Locality: USA: “Yakutat, Alaska (about latitude 59°35′).” 

Distribution: Canada and USA: Pacific coastal region from the 

Kenai Peninsula, Alaska, along the coast of British Columbia west of Cascades to the north bank of the lower Frasier 

River (Hall, 1981; Woodman and Fisher, 2016).

Remarks: Subgenus Otisorex. On the basis of multigene molecular analysis, S. c. streatori is a part of the West clade of 

S. cinereus (Hope et al., 2012).

Sorex dispar Batchelder, 1911

1896. Sorex macrurus Batchelder, Proc. Biol. Soc. Wash. 10:133; 8 December. Preoccupied by Sorex macrourus Lehmann, 1822 (=Sorex fodiens

Pennant, 1771 = Neomys fodiens); not Sorex macrurus Hodgson, 

1863:9, a nomen nudum; not Soriculus macrurus Blanford, 1888:231 

(=Episoriculus macrurus).

1911. Sorex dispar Batchelder, Proc. Biol. Soc. Wash. 24:97; 15 May. Replacement name for Sorex macrurus Batchelder, 1896, preoccupied by 

Sorex macrourus Lehmann, 1822.

1924. Sorex gaspensis Anthony and Goodwin, Am. Mus. Novit. 109:1; 

10 March. Holotype: AMNH 64190, from “Mount Albert, Gaspé 

Peninsula, Quebec, 2000 feet elevation.”

1956. Sorex dispar blitchi Schwartz, J. Elisha Mitchell Sci. Soc. 72(1):26; 

May. Holotype: USNM 301993, from “2 mi. NE Wagon Road Gap, 

elevation 4525 feet, Haywood County, North Carolina.”

1956. Sorex d[ispar]. dispar: Schwartz, J. Elisha Mitchell Sci. Soc. 

72(1):26; May. 

2004. S[orex]. d[ispar]. gaspensis: Rhymer et al., J. Mamm. 85(2):336; 

12 April.

Holotype: MCZ 41744.

Type Locality: USA: “Beede’s [sometimes called Keene 

Heights], in the township of Keene, Essex county, New 

York”; restricted by Martin (1966) to “0.6 mile south and 

0.5 mile east of Saint Huberts, Essex County, New York, lat. 

44°09′, long. 73°46′.”

Distribution: Canada and USA: occurs in a narrow band from 

central Maine to the southern border between Tennessee and North Carolina, with disjunct populations in the 

Gaspé Peninsula of Quebec, New Brunswick, Nova Scotia, 

and Cape Breton Island; elevational distribution: ~150–

2,025 m (Conaway and Pfitzer, 1952; Kirkland, 1981; Scott 

and van Zyll de Jong, 1989; Shafer and Stewart, 2006; 

Shafer et al., 2008).

FIGURE 27. Amphisorex lesueurii Duvernoy, 1842 (=Sorex cinereus Kerr, 1792). Image from Duvernoy (1842: pl. 50), courtesy of Biodiversity 

Heritage Library (https://www.biodiversitylibrary.org/), contributed by the Ernst Mayer Library, Museum of Comparative Zoology, Harvard 

University, Cambridge.
NUMBER 650 • 5 7

Remarks: Subgenus Otisorex; Sorex dispar group. Molecular analyses indicate that S. dispar may be the most basal 

branch within Otisorex (Shafer and Stewart, 2007). 

Analyses of mitochondrial and nuclear genetic markers 

among populations of S. dispar and S. gaspensis have indicated that they represent a single monophyletic group lacking 

any genetic structuring sufficient for either species or subspecies recognition (Rhymer et al., 2004; Shafer et al., 2008). Instead, the smaller body size of S. gaspensis appears to reflect a 

continuation of a cline of gradually decreasing body size from 

south to north (Rhymer et al., 2004), a conclusion consistent 

with those of Kirkland and Van Deusen (1979) and Scott and 

van Zyll de Jong (1989). Because the morphological relationships among S. gaspensis, S. dispar blitchi, and S. d. dispar

appear to represent a continuous cline and because the boundary between S. d. blitchi and S. d. dispar is clearly arbitrary 

(Schwartz, 1956; Kirkland and Van Deusen, 1979; Scott and 

van Zyll de Jong, 1989) and unsupported by genetic analyses 

(Shafer et al., 2008), I recognize no subspecies of S. dispar. 

Sorex emarginatus Jackson, 1925

1925. Sorex emarginatus Jackson, Proc. Biol. Soc. Wash. 38:129; 

13 November.

1955. Sorex oreopolus emarginatus: Findley, Univ. Kansas Publ. Mus. Nat.

Hist. 7(14):616; 10 June.

Holotype: USNM 90847.

Type Locality: Mexico: “Sierra Madre near Bolanos, altitude, 

7,600 feet, state of Jalisco.” 

Distribution: Mexico: highlands in southern Durango, southern Zacatecas, and northern Jalisco, 2,300–2,900 m (Carraway, 2007).

Remarks: Subgenus Otisorex; southern clade of the Sorex cinereus group (Hope et al., 2012); clade A2 of Esteva et al. 

(2010). Esteva et al. (2010) showed S. emarginatus embedded within S. milleri, and Hope et al. (2012) showed S. milleri embedded within S. cinereus. The relationships of these 

taxa require further careful investigation.

Sorex fontinalis Hollister, 1911

1911. Sorex fontinalis Hollister, Proc. U.S. Natl. Mus. 40(1825):378; 

17 April. 

1937. Sorex cinereus fontinalis: E. L. Poole, J. Mamm. 18(1):96; 11 February.

1977. S[orex]. c[inereus]. fonitnalis Kirkland, Proc. Penna. Acad. Sci. 51:46. 

Incorrect subsequent spelling of Sorex fontinalis Hollister, 1911.

Holotype: USNM 85439.

Type Locality: USA: “Cold Spring Swamp, near Beltsville, 

Maryland”; Prince George’s County.

Distribution: USA: southeastern and south central Pennsylvania, Delaware, and Maryland, extending into Accomack 

County, Virginia, and into Hampshire County, West Virginia (Kirkland and Levengood, 1987). 

Remarks: Subgenus Otisorex; Southern clade of the Sorex cinereus group (Hope et al., 2012). Apparently, the taxonomic 

status of S. fontinalis is still questionable (e.g., Hutterer, 

2005) despite the fact that Kirkland (1977) and Kirkland 

and Levengood (1987) showed clear size differences between 

S. cinereus and S. fontinalis in Pennsylvania and Kirkland 

(1977) reported the two taxa in sympatry in Lancaster and 

York Counties, Pennsylvania.

Sorex fumeus Miller, 1895

Remarks: Subgenus Otisorex; Sorex oreopolus group, part of 

Esteva et al.’s (2010) clade A3.

The validity of the two recognized subspecies warrants investigation as they may simply represent samples 

from either end of a latitudinal size cline, as seen in Sorex 

dispar, or some form of local variation. Sorex fumeus umbrosus was originally differentiated by its having grayer 

winter pelage and averaging slightly larger than S. f.

fumeus. Huggins and Kennedy (1989) reported a more 

complex pattern of size variation, however, with larger individuals generally in the central portion of the species’ 

distribution (southern Pennsylvania, Maryland, West Virginia, and northern Virginia) and smaller individuals to the 

north and to the south.

Sorex fumeus fumeus Miller, 1895

1857. Sorex forsteri: Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):22. Part; not 

Sorex forsteri Richardson, 1819 (=Sorex cinereus Kerr, 1792).

1857. Sorex richardsonii: Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):24. Part; 

not Sorex richardsonii Bachman, 1837 (=Sorex arcticus Kerr, 1772).

1890. Sorex platyrhinus: Dobson, Monogr. Insectiv., pt. 3, fasc. 1, pl. 23, 

fig. 5. Part; not Otisorex platyrhinus De Kay, 1842 (=Sorex cinereus

Kerr, 1792).

1895. Sorex fumeus Miller, N. Am. Fauna 10:50; 31 December.

1917. Sorex fumeus fumeus: Jackson, Proc. Biol. Soc. Wash. 30:149; 27 July. 

First use of current name combination.

Holotype: BM 1907.7.7.2582.

Type Locality: USA: “Peterboro, N. Y.”; Madison County.

Distribution: Canada and USA: from the northeastern tip of 

Minnesota northeast into southern Ontario (south of Lake 

Nipigon) and southern Quebec (Val-Jalbert; Saint-Nicolas) 

to the St. Lawrence River. In the west, the distribution is 

southwestward through the southeastern half of Ohio to 

central Kentucky and central Tennessee to the northeastern 

corner of Alabama and northern Georgia. In the east, the 

distribution passes through southern and central Vermont, 

New Hampshire, and the southernmost tip of Maine, then 

south through northern and western New Jersey, Pennsylvania, western Maryland, western Virginia, and western 

North Carolina to the northwestern corner of South Carolina (Hall, 1981; Owen, 1984; Jannett and Oehlenschlager, 

1994; Desroches and Picard, 2004; Felix et al., 2009). A 

single individual collected in 1883 is evidence of an isolated 

population at Racine, Racine County, Wisconsin (Jackson, 1928); elevational distribution: 209–1,981 m (USNM 

specimens).
5 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sorex fumeus umbrosus Jackson, 1917

1896. S[orex]. fumens Cox, Canadian Record of Science 7(1–2):118; 

24 June. Incorrect subsequent spelling of Sorex fumeus Miller, 1895.

1917. Sorex fumeus umbrosus Jackson, Proc. Biol. Soc. Wash. 30:149; 

27 July.

Holotype: USNM 150065.

Type Locality: Canada: “James River, Antigonish County, 

Nova Scotia.” 

Distribution: Canada and USA: Quebec south of the St. Lawrence River, New Brunswick, Prince Edward Island, and 

Nova Scotia (and probably Cape Breton Island), south to the 

northeastern corner of New York, northern Vermont, northern New Hampshire, and southern Maine (Hall, 1981; Owen, 

1984); elevational distribution: 6–457 m (USNM specimens).

Sorex haydeni Baird, 1857

FIGURE 23

1857. Sorex haydeni Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):29.

1896. Sorex personatus haydeni: J. A. Allen, Bull. Am. Mus. Nat. Hist. 

8(15):257; 25 November.

1925. Sorex cinereus haydeni: Jackson, J. Mamm. 6(1):56; 9 February.

Lectotype: USNM 1685, designated by Lyon and Osgood 

(1909:244).

Type Locality: USA: “Fort Union, Nebraska.” This locality is 

now near Fort Buford, Williams County, North Dakota. 

Distribution: Canada and USA: the approximate boundary of 

its distribution is from east central Alberta through southern Saskatchewan to southwestern Manitoba, then south 

through western Minnesota and Iowa to northern Missouri and northern Kansas, then north through Nebraska, 

northeastern Colorado, and eastern Montana, northwestern Iowa, northern Nebraska, southwestern South Dakota, 

southeastern Wyoming, and eastern Montana (Hall, 1981); 

elevational distribution: 785–2,682 m (USNM specimens).

Remarks: Subgenus Otisorex; Beringian clade of the Sorex cinereus group. The distribution of Sorex haydeni may overlap with the distributions of the Southwest and Midwest 

clades of Sorex cinereus (Hope et al., 2012).

Sorex hoyi Baird, 1857

FIGURES 11, 23

Remarks: Subgenus Otisorex; the Sorex hoyi group part of Esteva et al.’s (2010) clade A1. Sorex hoyi is the type species 

of Microsorex Coues, 1877. Sorex hoyi was revised by Long 

(1971) and Diersing (1980). 

Long (1971) noted a clinal increase in size in S. hoyi

from south to north. This variation is particularly noticeable in Long’s (1971: fig. 5) data, both among populations 

within eastern North American (South Carolina to through 

Quebec) and among populations within western North 

America (Idaho through Alaska). On the basis of morphology and morphometrics, Long (1971, 1974) recognized two 

species: Microsorex hoyi (Alaska, all of Canada north of 

the St. Lawrence River plus the Gaspé Peninsula, the western and midwestern United States) and M. thompsoni (the 

northeastern United States east of Lake Michigan and eastern Canada south of the St. Lawrence River). In contrast, 

van Zyll de Jong (1976) determined that the morphological 

differences used to differentiate the two taxa were size related and resulted from latitudinal clinal variation. Analysis 

of partial sequences of the mitochondrial cytochrome b gene 

by Stewart et al. (2002b) supported separation of S. hoyi

(western clade) and S. thompsoni (eastern clade), but with 

different distributions than those suggested by Long (1974), 

and they provided no clear definition of where the geographic boundary between the two taxa may occur. Drawn 

boundaries of putative subspecies vary greatly (Long, 1971, 

1974; Diersing, 1980; Hall, 1981; Kirkland et al., 1987; 

Whitaker and Hamilton, 1998; Stewart et al., 2002b), making it difficult to clearly define operational taxonomic units. 

Regardless, it appears extremely likely that S. hoyi includes 

at least two species. 

Sorex hoyi eximius (Osgood, 1901) 

1901. Sorex (Microsorex) eximius Osgood, N. Am. Fauna 21:71; 26 

September.

1905. Microsorex eximius: Elliot, Field Col. Mus. Publ. 105, Zool. Ser. 

6:457.

1925. Microsorex hoyi eximius: Jackson, Proc. Biol. Soc. Wash. 38:125; 13 

November.

1980. Sorex hoyi eximius: Diersing, J. Mamm. 61(1):89; 20 February. First 

use of current name combination.

Holotype: USNM 107126.

Type Locality: USA: “Tyonek, Cook Inlet, Alaska.” 

Distribution: USA: Alaska south of the Brooks Range (Diersing, 1980). 

Sorex hoyi hoyi Baird, 1857

FIGURE 11

1857. Sorex hoyi Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):32, pl. 28.

1901. [Microsorex] hoyi: Elliot, Field Col. Mus. Publ. 45, Zool. Ser. 2(1):377; 

6 March.

1902. Sorex (Microsorex) alnorum Preble, N. Am. Fauna 22:72; 31 October. Holotype: USNM 107014, from “Robinson Portage, Keewatin, 

Canada”; redefined by Miller and Kellogg (1955:34) as “upper Hayes 

River, about 35 miles southwest of Oxford Lake, at about lat. 54°30′

N., long. 96° W.), Manitoba.”

1905. Microsorex alnorum: Elliot, Field Col. Mus. Publ. 105, Zool. Ser. 

6:457.

1925. Microsorex hoyi intervectus Jackson, Proc. Biol. Soc. Wash. 38:125; 

13 November. Holotype: USNM 226979, from “Lakewood, Oconto 

County, Wisconsin.”

1925. Microsorex hoyi washingtoni Jackson, Proc. Biol. Soc. Wash. 38:125; 

13 November. Holotype: USNM 91007, from “Loon Lake, Stevens 

County, Washington.”
NUMBER 650 • 5 9

1925. Microsorex hoyi hoyi: Jackson, Proc. Biol. Soc. Wash. 38:126; 

13 November.

1925. Microsorex hoyi alnorum: Jackson, Proc. Biol. Soc. Wash. 38:126; 

13 November.

1980. Sorex hoyi hoyi: Diersing, J. Mamm. 61(1):90; 20 February. First use 

of current name combination.

Lectotype: USNM 632/1783, designated by Jackson (1928: 

203).

Type Locality: USA: “Racine”; Racine County, Wisconsin. 

Distribution: Canada and USA: southern Yukon, southern 

Northwest Territories, and northern Manitoba, south to 

northeastern Washington; Idaho County, Idaho; across 

western and northern Montana; east through northern 

North Dakota; south through central South Dakota east of 

the Missouri River to Clay and Union Counties; east across 

northern Iowa, Wisconsin, northern Illinois, and Michigan; 

north of Lake Erie, Lake Ontario, and the St. Lawrence 

River, but then extending through the Gaspé Peninsula 

(Diersing, 1980; Foresman, 1999; King et al., 1999; Murphey et al., 2007; Hendricks and Lenard, 2014).

Remarks: Long (1971, 1974) and Hall (1981) recognized S. h.

alnorum as a distinct subspecies; Diersing (1980) did not. 

On the basis of preliminary molecular analysis, Stewart et 

al. (2002b) indicated that eastern populations of S. h. hoyi

and S. h. thompsoni may be distinct from populations west 

of James Bay and Lake Michigan.

Sorex hoyi montanus L. N. Brown, 1966

1966. Microsorex hoyi montanus L. N. Brown, Proc. Biol. Soc. Wash. 79:50; 

23 May. Not Sorex montanus Kelaart, 1850:211 (=Suncus montanus); 

not Sorex isodon princeps montanus Skalon and Rajevsky, 1940:199; 

not Sorex isodon montanus Pavlinov, Borisenko, Kruskop, and 

Yahontov, 1995:36 (see Hutterer and Zaitsev, 2004).

1980. Sorex hoyi montanus: Diersing, J. Mamm. 61(1):99; 20 February. First 

use of current name combination.

Holotype: UWYMV 1449.

Type Locality: USA: “from edge of Trail’s Divide, 1⁄4 mi. S of 

Univ. Wyoming summer Science Camp on state Highway 

130, Centennial, Albany Co., Wyoming.”

Distribution: USA: isolated distribution in the Medicine Bow 

Range of southcentral Wyoming and the central Rocky 

Mountains of Colorado (Diersing, 1980). 

Sorex hoyi thompsoni Baird, 1857

FIGURE 23

1857. Sorex thompsoni Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):34, pl. 27.

1865. Sorex, Thomsoni Gilpin, Proc. Trans. Nova Scot. Inst. Nat. Sci. 1(2):1. 

Incorrect subsequent spelling of Sorex thompsoni Baird, 1857.

1925. Microsorex hoyi thompsoni: Jackson, Proc. Biol. Soc. Wash. 38:126; 

13 November.

1971. Microsorex thompsoni thompsoni: Long, Trans. Kansas Acad. Sci. 

74(2):192; 7 April.

1980. Sorex hoyi thompsoni: Diersing, J. Mamm. 61(1):94; 20 February. 

First use of current name combination.

Lectotype: USNM 1686/38838, designated by Lyon and Osgood (1909:250).

Type Locality: USA: “Burlington, V[ermon]t”; Chittenden County.

Distribution: Canada and USA: south of the Gaspé Peninsula 

and the St. Lawrence River in southernmost Quebec, New 

Brunswick, Prince Edward Island, and Nova Scotia; south 

through Maine, Vermont, New Hampshire, and New York 

through all but southeastern Pennsylvania to eastern Ohio 

and extending into Maryland as far as Baltimore (Diersing, 

1980; Kirkland et al., 1987; Laerm et al., 1994; Whitaker 

and Hamilton, 1998; Kirkland and Hart, 1999).

Remarks: Long (1971, 1974) recognized Microsorex thompsoni as a distinct species with two subspecies: M. t. thompsoni and M. t. winnemana; Diersing (1980) considered both 

taxa to be subspecies of S. hoyi. On the basis of preliminary 

molecular analysis Stewart et al. (2002b) indicated that eastern populations of S. hoyi, including S. h. thompsoni, may 

be distinct from populations west of James Bay and Lake 

Michigan.

Sorex hoyi winnemana Preble, 1910

1910. Microsorex winnemana Preble, Proc. Biol. Soc. Wash. 23:101; 

24 June.

1925. Microsorex hoyi winnemana: Jackson, Proc. Biol. Soc. Wash. 38:126; 

13 November. 

1971. Microsorex thompsoni winnemana: Long, Trans. Kansas Acad. Sci. 

74(2):187; 7 April.

1980. Sorex hoyi winnemana: Diersing, J. Mamm. 61(1):98; 20 February. 

First use of current name combination.

Holotype: USNM 126320.

Type Locality: USA: “Fairfax County (bank of Potomac River 

near Stubblefield Falls), Virginia”; Jackson (1928:206) 

added that this locality is “4 miles below Great Falls of the 

Potomac” River. 

Distribution: USA: in the east, from Prince George’s and Garrett Counties, Maryland, south through central Virginia, to 

the northwestern corner of South Carolina and northern 

Georgia; in the west, from Greenup County, Kentucky, and 

Owen, Monroe, Brown, and Bartholomew Counties, Indiana, south through central and eastern Kentucky and eastern Tennessee to northeastern Alabama and northwestern 

Georgia. A possibly isolated population was reported from 

Camden and Gates Counties, North Carolina (Diersing, 

1980; Caldwell and Bryan, 1982; Cudmore and Whitaker, 

1984; Kirkland et al., 1987; Laerm et al., 1994; Padget and 

Rose, 1994; Whitaker and Hamilton, 1998; Czech et al., 

2017).

Remarks: Long (1971, 1974) considered this taxon to be a subspecies of Microsorex thompsoni; Diersing (1980) considered it to be a subspecies of S. hoyi. 
6 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sorex ibarrai Matson and McCarthy, 2005

2005. Sorex veraepacis ibarrai Matson and McCarthy, Special Publication of 

the International Society of Shrew Biologists 1:68; December.

2017. Sorex ibarrai: Matson and Ordóñez-Garza, Zootaxa 4236(3):478; 

24 February. First use of current name combination.

Holotype: CM 113283.

Type Locality: “Guatemala, Departamento El Progreso, Municipio San Augustín Acasaguastlan, Reserva Biosfera Sierra 

de las Minas, Cerro Pinalón, Camino de las Torres, 2700 m; 

15°04′54′′ N, 89°55′59′′ W.”

Distribution: Guatemala: Sierra de las Minas in El Progreso 

and Zacapa Departments; elevational distribution: 1,475–

2,800 m (Matson and McCarthy, 2005).

Remarks: Unnamed subgenus; S. veraepacis group (Matson 

and Ordóñez-Garza, 2017). Originally described as a subspecies of S. veraepacis by Matson and McCarthy (2005).

Sorex ixtlanensis Carraway, 2007

2007. Sorex ixtlanensis Carraway, Monogr. West. N. Am. Nat. 3:52; 

22 June.

Holotype: KU 124320.

Type Locality: Mexico: “‘N slope Cerro Pélon, 31.6 kms. S (by 

road) Vista Hermosa, 2650 m., Oaxaca,’ latitude 17.36° N, 

longitude 95.25° W.” 

Distribution: Mexico: Guerrero and Oaxaca, 1,920–3,000 m 

(Carraway, 2007).

Remarks: Subgenus Otisorex; S. veraecrucis group. Populations now recognized as S. ixtlanensis were previously part 

of S. veraepacis mutabilis (Jackson, 1928).

Sorex jacksoni Hall and Gilmore, 1932

1932. Sorex jacksoni Hall and Gilmore, Univ. Calif. Publ. Zool. 38(9):392; 

17 September.

Holotype: MVZ 51142

Type Locality: USA: “Sevoonga, 2 miles east of North Cape, St. 

Lawrence Island, Bering Sea, Alaska.” 

Distribution: Known only from St. Lawrence Island.

Remarks: Subgenus Otisorex; Beringian clade of the Sorex cinereus group (Hope et al., 2012). Hope et al. (2012) found 

low levels of genetic divergence among high-latitude tundra/

taiga species in the Beringian clade of the Sorex cinereus

group (S. camtschatica Yudin, 1972; S. jacksoni; S. leucogaster Kuroda, 1933; S. portenkoi Stroganov, 1956; S. pribilofensis Merriam, 1895; S. ugyunak Anderson and Rand, 

1945) that they interpreted as evidence of recent speciation. 

In that analysis, S. jacksoni was sister to the eastern Siberian 

species S. camtschatica. Despite the relative lack of differentiation and lack of evidence of paraphyly and polyphyly in 

their cytochrome b tree, they urged caution in revising the 

taxonomic status of these species.

Sorex longirostris Bachman, 1837

FIGURE 20

Remarks: Subgenus Otisorex. Sorex longirostris may be either part of the southern clade of the Sorex cinereus group 

(Esteva et al., 2010) or one of the two basal branches in 

the Sorex cinereus group (Hope et al., 2012). Sorex longirostris was reviewed by Jones et al. (1991) and Webster et 

al. (2009).

Sorex longirostris eionis J. A. Davis, 1957

1957. Sorex longirostris eionis J. A. Davis, Am. Mus. Novit. 1844:3; 

10 October.

Holotype: AMNH 163855.

Type Locality: USA: “Homosassa Springs, Citrus County, 

Florida.”

Distribution: USA: northern two-thirds of the Florida peninsula (Jones et al., 1991).

Sorex longirostris fisheri Merriam, 1895

1895. Sorex fisheri Merriam, N. Am. Fauna 10:86; 31 December.

1928. Sorex longirostris fisheri: Jackson, N. Am. Fauna 51:87; 24 July. First 

use of current name combination.

Holotype: USNM 75166.

Type Locality: USA: “Lake Drummond, Dismal Swamp, Virginia”; Norfolk County.

Distribution: USA: originally restricted to the vicinity of the 

historical Great Dismal Swamp in Virginia and North Carolina (Jones et al., 1991), the distribution is now considered 

to encompass much of the Atlantic Coastal Plain from the 

mouth of the James River on the eastern coastline of Virginia to the northeastern corner of South Carolina (Webster 

et al., 2009).

Sorex longirostris longirostris Bachman, 1837

FIGURE 20

1837. Sorex longirostris Bachman, J. Acad. Nat. Sci. Phila. 7(2):370, pl. 23: 

fig. 2.

1842. O[tisorex]. longirostris: De Kay, Zool. New York, p. 23.

1848. [Musaraneus (Crocidura)] Bachmani Pomel, Arch. Sci. Phys. Nat., 

9:249. Renaming of Sorex longirostris Bachman, 1837.

1857. Sorex personatus: Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):30. Part; 

not Sorex personatus I. Geoffroy Saint-Hilaire, 1827.

1868. Sorex wagneri Fitzinger, Sitzungsber. Math.-natürwissensch. Classe 

Kaiserl. Akad. Wissensch. 57(1):512. Holotype: not known to exist, from “Nord-Amerika.” The holotype was a specimen in the 

“Münchener Museum” that Wagner (1855:547) previously referred to 

S. longirostris. The specimen presumably was destroyed during World 

War II (Anneke van Heteren, Zoologische Staatssammlung München, 

Munich, Germany, email communication, 6 July 2017). 

1928. Sorex longirostris longirostris: Jackson, N. Am. Fauna 51:28; 24 July. 

First use of current name combination.
NUMBER 650 • 6 1

2009. Sorex logirostris: Whitaker and Mumford, Mammals of Indiana, 2nd 

ed., p. 124. Incorrect subsequent spelling of Sorex longirostris Bachman, 1837.

Holotype: ANSP 479.

Type Locality: USA: “swamps of Santee.” Restricted by Jackson (1928:85) to “Hume Plantation, swamps of the Santee 

River [Cat Island, mouth of Santee River], S. C.” Handley 

and Varn (1994:395) noted that “Cat Island is on the north 

side of the mouth of the Santee River.” The type locality is 

now part of the Tom Yawkey Wildlife Center, Georgetown 

County, South Carolina.

Distribution: USA: western limit from north central Missouri, 

central Illinois, Indiana, and northern Kentucky, south 

through southwestern Missouri, eastern Oklahoma, northern and southeastern Louisiana to the coast; eastern limit 

from central West Virginia and southern Maryland east 

of the Chesapeake Bay, south through Virginia, except the 

southeastern corner, the piedmont of North Carolina, all 

but the northeastern corner of South Carolina to southeastern Georgia, and the panhandle of Florida; elevational distribution: near sea level to 366 m (Pagels et al., 1982; Jones 

et al., 1991; Webster et al., 2009).

Sorex lyelli Merriam, 1902

1902. Sorex tenellus lyelli Merriam, Proc. Biol. Soc. Wash. 15:75; 

22 March.

1928. Sorex lyelli: Jackson, N. Am. Fauna 51:57; 24 July. First use of current 

name combination.

Holotype: USNM 109530.

Type Locality: USA: “Mount Lyell, Tuolumne Co., California.” 

Distribution: USA: central Sierra Nevada in Mariposa, Mono, 

Tuolumne, and Fresno Counties, California; elevational distribution: 2,100–3,630 m (Epachin and Engilis, 2009).

Remarks: Subgenus Otisorex; Sorex cinereus group. Within 

the Beringian clade of the Sorex cinereus group, S. lyelli is a 

member of a subclade that also includes S. preblei and two 

undescribed western North American species (Hope et al., 

2012).

Sorex macrodon Merriam, 1895

1895. Sorex macrodon Merriam, N. Am. Fauna 10:82, pl. 7, figs. 2, 2a; pl. 

12, figs. 12, 13; 31 December.

Holotype: USNM 58272.

Type Locality: Mexico: “Orizaba, Veracruz, Mexico (altitude, 

4,200 feet).” 

Distribution: Mexico: highlands in Puebla, Veracruz, and 

Oaxaca; elevational distribution: 1,280–2,590 m (Carraway, 2007).

Remarks: Unnamed subgenus; Sorex veraepacis group (clade 

B2 of Esteva et al., 2010). 

Sorex madrensis Matson and Ordóñez-Garza, 2017

1928. Sorex veraepacis chiapensis Jackson, N. Am. Fauna 51:150; 24 July. 

Part.

2017. Sorex madrensis Matson and Ordóñez-Garza, Zootaxa 4236(3):479; 

24 February.

Holotype: USNM 569709.

Type Locality: Guatemala: “coniferous-hardwood cloud forest 

approximately 5 km ENE Cabricán, Bosque Ojo de Agua, 

3100 m, Quetzaltenango, Guatemala, latitude 15°05′ N, 

longitude 91°36′ W.”

Distribution: Guatemala: Sierra Madre of southern Guatemala; elevational distribution: 2,300–3,100 m (Matson and 

Ordóñez-Garza, 2017).

Remarks: Unnamed subgenus; S. veraepacis group. The populations now recognized as S. madrensis previously formed 

part of Sorex veraepacis chiapensis (Jackson, 1928; Woodman et al., 2012).

Sorex maritimensis R. W. Smith, 1939

1939. Sorex arcticus maritimensis R. W. Smith, J. Mamm. 20(2):244; 

14 May.

2002. Sorex maritimensis: Stewart et al., Can. J. Zool. 80:94; published online 7 February. First use of current name combination.

Holotype: MVZ 84479.

Type Locality: Canada: “Wolfville, Kings County, Nova 

Scotia.” 

Distribution: Canada: Nova Scotia and New Brunswick 

(McAlpine et al., 2012).

Remarks: Subgenus Sorex; Sorex araneus group. Sorex maritimensis was formerly treated as a subspecies of S. arcticus

but was recognized as a distinct species by Stewart et al. 

(2002a) and Hutterer (2005) on the basis of morphometric 

work by van Zyll de Jong (1983), karyotypes analyzed by 

Volobouev and van Zyll de Jong (1988), and genetic evidence provided Stewart et al. (2002a).

Sorex mccarthyi Matson and Ordóñez-Garza, 2017

2017. Sorex mccarthyi Matson and Ordóñez-Garza, Zootaxa 4236(3):475; 

24 February.

Holotype: CM 119732.

Type Locality: Honduras: “in cloud forest, approximately 

11 km SW Gracias, 2560 m, Campamento El Naranjo, 

Parque Nacional Celaque, Lempira, Honduras, latitude 

14°33′ N, longitude 88°40′ W.” 

Distribution: Honduras: known only from the type locality 

(Matson and Ordóñez-Garza, 2017).

Remarks: Unnamed subgenus; Sorex saussurei group. The holotype was originally identified as S. salvini (Woodman et 

al., 2012). Matson and Ordóñez-Garza (2017) considered 

S. mccarthyi to be closely related to S. salvini.
6 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sorex mediopua Carraway, 2007

2007. Sorex mediopua Carraway, Monogr. West. N. Am. Nat. 3:54; 22 June.

Holotype: KU 112032.

Type Locality: Mexico: “‘12 mi SW Cd. Guzmán, 10000 ft., 

Jalisco,’ latitude 19.56° N, longitude 103.61° W.” 

Distribution: Mexico: highlands in Jalisco, Michoacán, 

México, and Guerrero; elevational distribution: 1,875–

3,050 m (Carraway, 2007).

Remarks: Unnamed subgenus: Sorex saussurei group. This 

species previously formed part of Sorex saussurei saussurei

Merriam, 1892.

Sorex merriami Dobson, 1890

1890. Sorex merriami Dobson, Monogr. Insectiv., pt. 3, fasc. 1, pl. 23, fig. 

6; May.

1909. Sorex leucogenys Osgood, Proc. Biol. Soc. Wash. 22:52; 17 April. Holotype: USNM 157952, from “mouth of the canyon of Beaver River, 

about 3 miles east of Beaver, Beaver Co., Utah,” USA. 

1939. Sorex merriami merriami: Benson and Bond, J. Mamm. 20(3):348; 

14 August.

1939. Sorex merriami leucogenys: Benson and Bond, J. Mamm. 20(3):348; 

14 August.

Holotype: USNM 186441.

Type Locality: USA: “Fort Custer, Montana.” Restricted by 

Miller and Kellogg (1955:17) to “Little Bighorn River, 

about a mile and a half above Fort Custer, Bighorn County, 

Mont.” 

Distribution: Canada and USA: from southernmost British 

Columbia, north central Montana, and west central North 

Dakota south to southern California and southern New 

Mexico; elevational distribution 195–2,900 m (Diersing and Hoffmeister, 1977; Shaughnessy and Woodman, 

2015).

Remarks: Unnamed subgenus (George, 1988:456); Sorex merriami group. Sorex merriami is considered to be monotypic 

(Diersing and Hoffmeister, 1977:328).

Sorex milleri Jackson, 1947

1947. Sorex milleri Jackson, Proc. Biol. Soc. Wash. 60:131; 9 October.

Holotype: USNM 274950.

Type Locality: Mexico: “Madera Camp, altitude 8,000 feet, 

Carmen Mountains, Coahuila.” 

Distribution: Mexico: highlands of Coahuila and Nuevo León; 

elevational distribution: 2,440–3,660 m.

Remarks: Subgenus Otisorex; southern clade of the Sorex cinereus group (Hope et al., 2012); clade A2 of Esteva et al. 

(2010). Esteva et al. (2010) showed S. emarginatus embedded within S. milleri, and Hope et al. (2012) showed S. milleri embedded within S. cinereus. The relationships of these 

taxa require further careful investigation.

Sorex minutissimus Zimmermann, 1780

1780. Sorex minutissimus Zimmermann, Geog. Gesch. Mensch. Vierf. 

Thiere 2:385. 

Holotype: Not known to exist. This species was described by 

Zimmermann (1780) based on earlier descriptions by Pallas 

(1773:664) and Schreber (1778).

Type Locality: Russia: “Herr Pallas fand sie am Ienisen” (Mr. 

Pallas found them at the Yenisei [River]). This locality description is a shorter version of Schreber’s (1778:577): “Der 

Herr Professor Pallas hat sie in Sibirien am Ienisei entdeckt” 

(Professor Pallas discovered it in Siberia at the Yenisei 

[River]). The type locality was restricted by Pavlinov and 

Rossolimo (1987:23) to “Красноярский кр., Красноярск” 

(Krasnoyarskii krai, Krasnoyarsk), Russia.

Remarks: Subgenus Sorex; Sorex minutissimus group. Only 

one subspecies of Sorex minutissimus is known to occur in 

North America.

Sorex minutissimus yukonicus Dokuchaev, 1997

1997. Sorex yukonicus Dokuchaev, J. Mamm. 78(3):814; August. 

2012. Sorex minutissimus: Hope et al., Northwest. Nat. 93(2):104; Autumn.

Holotype: UAM 19268.

Type Locality: USA: “Crow Creek, 1 3⁄4 miles N, 2 1⁄4 miles W 

Beaver Creek (64°44′ N, 156°50′ W) near Galena, Alaska.”

Distribution: Canada and USA: through much of Alaska and 

into the northern Yukon Territory (Hope et al., 2010; Cook 

et al., 2016).

Remarks: The taxon Sorex yukonicus Dokuchaev, 1997 

was described for the North American representatives of 

S. minutissimus, but Hope et al. (2010) determined that 

S. yukonicus and Siberian S. minutissimus are conspecific. 

Despite this, North American S. minutissimus appears to 

represent a weakly differentiated clade (Hope et al., 2010), 

which is why I recognize it herein as a subspecies.

Sorex monticola Merriam, 1890

Remarks: Subgenus Otisorex; S. monticola group. Phylogenetic 

analysis of the mitochondrial cytochrome b gene indicated 

that S. monticola, as considered by Alexander (1996), is not 

monophyletic but includes three genetically divergent clades 

(Demboski and Cook, 2001; Shafer and Stewart, 2007) that 

I treat herein as three species using the oldest available name 

for each clade. Sorex monticola (sensu stricto) thus refers to 

Demboski and Cook’s (2001) “southern continental clade”; 

S. obscurus refers to their “northern continental clade,” 

and S. pacificus refers to their “coastal clade.” Subspecies 

comprising these clades are mostly those recognized in taxonomic revisions by Carraway (1990) and Alexander (1996). 

Most species-group names are adjectives that must 

agree with the genus-level name in gender and number 

(ICZN, 1999: Article 31.2.1; see also Gardner and Hayssen, 
NUMBER 650 • 6 3

2004). The name monticola, however, like any species-group 

name ending in -cola, is an invariable noun in apposition, 

which does not change gender (Woodman, 2012a). The suffix -cola means “dweller” or “inhabitant”; hence, monticola = “mountain dweller.” Merriam (1890:43) made this 

error when he first described the species as S. monticolus, 

but he subsequently corrected the spelling to S. monticola

(Merriam, 1895b:69).

Sorex monticola monticola Merriam, 1890

1890. Sorex monticolus Merriam, N. Am. Fauna 3:43; 11 September.

1895. Sorex vagrans monticola: Merriam, N. Am. Fauna 10:69; 31 December. Justified emendation for noun in apposition.

1912. Sorex obscurus obscurus: Miller, Bull. U.S. Natl. Mus. 79:15; 31 December. Part; not Sorex obscurus Merriam, 1895. 

1925. Sorex durangae Jackson, Proc. Biol. Soc. Wash. 38:127; 13 November. 

Holotype: USNM 94540, from “El Salto, Durango, Mexico.” 

1932. Sorex melanogenys Hall, J. Mamm. 13(3):260; 9 August. Holotype: 

MVZ 50247, from “Marijilda Canyon, 8600 feet altitude, Graham 

Mountains [=Pinaleno Mountains], Graham County, Arizona.” 

1955. Sorex vagrans obscurus: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):43; 10 December. Part; not Sorex obscurus Merriam, 1895.

1977. S[orex]. m[onticolus]. monticolus: Hennings and Hoffmann, Occ.

Pap. Mus. Nat. Hist. Univ. Kansas 68:4; 15 July. First use of current 

name combination.

1977. S[orex]. monticolus obscurus: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:5; 15 July. Part; not Sorex obscurus

Merriam, 1895.

Holotype: USNM 17599/24535.

Type Locality: USA: “San Francisco Mountain, Arizona . . . 

Altitude 3,500 meters (11,500 feet)” [Coconino County]. 

Distribution: USA and Mexico: discontinuously distributed 

in montane regions from east central California, Utah, and 

Wyoming south to southern Durango; elevational distribution: 1,830–3,810 m (Hennings and Hoffmann, 1977; Alexander, 1996; Carraway, 2007). 

Remarks: Includes Alexander’s (1996) S. m. monticola and 

southern populations that she referred to S. m. obscurus

(Demboski and Cook, 2001). 

Sorex monticola neomexicanus Bailey, 1913

1913. Sorex obscurus neomexicanus Bailey, Proc. Biol. Soc. Wash. 26:133; 

21 May.

1955. Sorex vagrans neomexicanus: Findley, Univ. Kansas Publ. Mus. Nat.

Hist. 9(1):50; 10 December.

1977. S[orex]. m[onticolus]. neomexicanus: Hennings and Hoffmann, Occ.

Pap. Mus. Nat. Hist. Univ. Kansas 68:4; 15 July. First use of current 

name combination.

1996. Sorex neomexicanus: Alexander, Univ. Kansas Mus. Nat. Hist., Misc.

Publ. 88:21; 21 February. 

Holotype: USNM 100440.

Type Locality: USA: “Cloudcroft, New Mexico (alt. 9000 feet), 

in the Sacramento Mountains” [Otero County].

Distribution: USA: the Capitan and Sacramento Mountains of 

south central New Mexico; elevational distribution: 2,055–

2,750 m [Hennings and Hoffmann, 1977; Alexander, 1996; 

VertNet records (http://www.vertnet.org; accessed 18 June 

2017)].

Remarks: Subgenus Otisorex. Sorex monticola group. Molecular analysis indicates this taxon is embedded within the southern clade of Demboski and Cook (2001; Esteva et al., 2010).

Sorex mutabilis Merriam, 1898

1895. Sorex saussurei caudatus Merriam, N. Am. Fauna 10:84; 31 December. Preoccupied by Sorex caudatus Horsfield, 1851 (=Episoriculus caudatus); not Sorex caudatus Hodgson, 1849, a nomen nudum.

1898. Sorex saussurei mutabilis Merriam, Science 8:782; 2 December. Renaming of Sorex saussurei caudatus Merriam, 1895, preoccupied by 

Sorex caudatus Horsfield, 1851.

1925. Sorex veraepacis mutabilis: Jackson, Proc. Biol. Soc. Wash. 38:130; 

13 November.

2010. S[orex]. veraepacis (10): Esteva et al., Zootaxa 2615:55, fig. 1; 17 September. Part; not Sorex veraepacis Alston, 1877.

Holotype: USNM 69600.

Type Locality: Mexico: “Reyes, Oaxaca, Mexico (altitude 

10,200 feet).”

Distribution: Mexico: highlands in Guerrero and Oaxaca; 

elevational distribution: 1,120–3,110 m (Carraway, 2007).

Remarks: Unnamed subgenus; Sorex veraepacis group (Matson 

and Ordóñez-Garza, 2017); clade B2 of Esteva et al. (2010; 

as S. veraepacis locality 10). Sorex chiapensis, S. ibarrai, 

and S. mutabilis previously were treated as subspecies of 

S. veraepacis (Carraway, 2007; Woodman et al., 2012). 

Matson and Ordóñez-Garza (2017) showed S. chiapensis, 

S. ibarrai, and S. veraepacis to be distinct species. Given the 

disjunct distributions of S. mutabilis to the west of the Isthmus of Tehuantepec and S. veraepacis to the east and given 

the strong genetic divergence between them (Esteva et al., 

2010), it seems most appropriate to treat S. mutabilis and 

S. veraepacis as distinct species.

Sorex nanus Merriam, 1928

1895. Sorex tenellus nanus Merriam, N. Am. Fauna 10:81; 31 December.

1928. Sorex nanus: Jackson, N. Am. Fauna 51:174; 24 July.

Holotype: USNM 73773.

Type Locality: USA: “Estes Park, Colorado”; Larimer County.

Distribution: USA: from north central Montana south to western Wyoming and central South Dakota to east central 

Arizona and south central New Mexico; elevational distribution: 2,130–3,505 m [Hall, 1981; Backlund, 1995; VertNet, http://www.vertnet.org; accessed 21 June 2017].

Remarks: Subgenus Otisorex; Sorex oreopolus group. 
6 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sorex navigator (Baird, 1857) 

FIGURE 25

Remarks: Subgenus Otisorex; Sorex palustris group. Sorex navigator is the type species of Neosorex Baird, 1857. Within 

the S. palustris group, S. navigator is sister to S. bendirii

(Hope et al., 2014).

Hope et al.’s (2014) investigation of the molecular phylogenetics of North American water shrews revealed three 

species-level clades among the subspecies formerly united 

as Sorex palustris: eastern S. albibarbis, western cordilleran 

S. navigator, and boreal S. palustris (sensu stricto). Nagorsen 

et al. (2017) demonstrated that parapatric S. navigator and 

S. palustris can be distinguished morphometrically.

Sorex navigator includes two subspecies: S. navigator

navigator on the continent and S. n. brooksi from Vancouver 

Island. The northeastern distributional extent of S. navigator navigator is incompletely understood (Hope et al., 2014).

Sorex navigator brooksi Anderson, 1934

1934. Sorex palustris brooksi Anderson, Can. Field-Nat. 48(8):134; 

1 November.

2014. S[orex]. navigator brooksi: Hope et al., J. Mamm. 95(4):730; 

22 August. 

Holotype: CMN 12370.

Type Locality: Canada: “Black Creek, 150 feet altitude, Comox 

district, east coast of Vancouver Island, British Columbia, 

latitude 49°50′ north, longitude 125°08′ west.” 

Distribution: Canada: Vancouver Island (Hope et al., 2014).

Sorex navigator navigator (Baird, 1857)

FIGURE 25

1857. Neosorex navigator Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):11, pl. 

26; 14 July.

1863. Sorex navigator: Verrill, Proc. Boston Soc. Nat. Hist. 9:167; February.

1891. Sorex palustris: Merriam, N. Am. Fauna 5:35; August.

1895. Sorex (Neosorex) palustris navigator: Merriam, N. Am. Fauna 10:92; 

31 December.

1900. Sorex navigator alaskanus Merriam, Proc. Wash. Acad. Sci. 2:18; 

14 March. Holotype: USNM 97713, from “Point Gustavus, Glacier 

Bay, Alaska,” USA.

1901. [Neosorex palustris] navigator: Elliot, Field Col. Mus. Publ. 45, Zool.

Ser. 2(1):379; 6 March.

1906. Sorex palustris navigator: Stephens, California Mammals, p. 254. 

1912. Neosorex navigator navigator: Miller, Bull. U.S. Natl. Mus., 79:21; 

31 December.

2014. Sorex navigator: Hope et al., J. Mamm. 95(4):730; 22 August.

Holotype: USNM 629/1780 (not 629/1760, as indicated in 

Baird, 1857: pl. 26).

Type Locality: USA: “Fort Vancouver, W[estern]. T[erritories],” 

along the Columbia River, Vancouver, Clark County, Washington. In describing Neosorex navigator, Baird (1857) apparently based the type locality on faulty information. The 

original collector, J. G. Cooper (1860b:73), later wrote, 

“But one specimen of this species was obtained during the 

expedition; this, according to the label now attached, was 

found at Fort Vancouver, but I am inclined to consider this a 

mistake, and that it was really taken while swimming under 

water in a lake near the summit of the Cascade mountains; 

August 31, 1853.” Further supporting his memory of the 

capture of this animal, Cooper (1860a:36) noted, “Aquatic 

mammalia . . . abound in the fresh waters; and one seems 

to be peculiar to the Territory, the water shrew, (Neosorex 

navigator,) caught while swimming a foot below the surface 

of one of the lakes at the head of the Yakima river, and at 

least 2,500 feet above the ocean.”

Cooper’s accounts caused Jackson (1928:184) to fix the 

type locality as “near head of Yakima River, Cascade Mountains, Wash.”; Kittitas County.

Distribution: Canada and USA: from central Alaska and central Yukon Territory south through British Columbia; in 

and east of the Cascades in Washington and Oregon; western and central Wyoming; to east central Arizona and west 

central New Mexico (Hope et al., 2014); elevational distribution: 90–3,415 m (USNM specimens).

Sorex obscurus Merriam, 1895

Remarks: Subgenus Otisorex; Sorex monticola group. Phylogenetic analysis of the mitochondrial cytochrome b gene 

indicated that S. monticola, as conceived by Alexander 

(1996), is not monophyletic but comprises three genetically 

divergent species-level clades for which the oldest available 

names are (Demboski and Cook, 2001; Shafer and Stewart, 

2007) S. monticola for their “southern continental clade,” 

S. obscurus for the “northern continental clade,” and 

S. pacificus for the “coastal clade.” The subspecies of S. obscurus recognized here are based primarily on Alexander’s 

(1996) review of S. monticola. 

Sorex obscurus alascensis Merriam, 1895

1895. Sorex obscurus alascensis Merriam, N. Am. Fauna 10:76; 

31 December.

1900. Sorex glacialis Merriam, Proc. Wash. Acad. Sci. 2:16; 14 March. 

Holotype: USNM 97709, from “Point Gustavus, on east side of entrance to Glacier Bay, Alaska,” USA.

1900. S[orex]. alascensis: Merriam, Proc. Wash. Acad. Sci. 2:18; 14 March.

1901. [Sorex glacialis] alascensis: Elliott, Field Col. Mus. Publ. 45, Zool. Ser. 

2(1):372; 6 March.

1912. Sorex alascensis alascensis: Miller, Bull. U.S. Natl. Mus. 79:16; 

31 December. 

1955. Sorex vagrans alascensis: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):41; 10 December.

1977. S[orex]. m[onticolus]. alascensis: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

Holotype: USNM 73539.

Type Locality: USA: “Yakutat Bay, Alaska.” 
NUMBER 650 • 6 5

Distribution: Canada and USA: coastal southeastern Alaska 

from western Prince William Sound to Juneau and Admiralty Island and adjacent parts of northwestern British Columbia (Alexander, 1996).

Sorex obscurus obscurus Merriam, 1895

1891. Sorex vagrans similis Merriam, N. Am. Fauna 5:34; 30 July. Preoccupied by Sorex similis Hensel, 1855 (=Asoriculus similis), an extinct 

subfossil shrew from Cagliari, Sardinia.

1895. Sorex obscurus Merriam, N. Am. Fauna 10:72; 31 December. Replacement name for Sorex vagrans similis Merriam, 1891, preoccupied 

by Sorex similis Hensel, 1855.

1912. Sorex obscurus obscurus: Miller, Bull. U.S. Natl. Mus. 79:15; 31 December. First use of current name combination. 

1955. Sorex vagrans obscurus: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):43; 10 December.

1955. Sorex vagrans longiquus Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):49; 10 December. Holotype: UMMZ 87332, from “25 mi. ESE Big 

Sandy, Eagle Creek, Choteau Co., Montana,” USA. 

1955. Sorex vagrans obscuroides Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):58; 10 December. Holotype: USNM 30064/42074, from “Bishop 

Creek, 6600 ft., Inyo Co., California,” USA.

1977. S[orex]. m[onticolus]. obscurus: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

1996. S[orex]. m[onticola]. obscuroides: Alexander, Univ. Kansas Mus. Nat.

Hist., Misc. Publ. 88:6; 21 February. 

Holotype: USNM 23525/30943.

Type Locality: USA: “Timber Creek, Salmon River Mountains, 

Idaho (altitude, 8,200 feet)”; Lemhi Mountains, 16 km west 

of Junction, Lemhi County (Jackson, 1928:117).

Distribution: Canada and USA: from central northern Alaska 

and northwestern corner of the Northwest Territory south 

through western Canada (excluding the coastal region) 

to western Oregon, Idaho, and northwestern Wyoming; 

elevational distribution: 120–2,500 m (Hennings and 

Hoffmann, 1977; Alexander, 1996; Demboski and Cook, 

2001).

Remarks: Hennings and Hoffmann (1977:15) and Alexander 

(1996:30) agreed on morphological grounds that S. vagrans 

obscuroides Findley, 1955 does not warrant recognition as 

a subspecies distinct from S. o. obscurus. The relationships 

of this seemingly isolated population warrant molecular investigation, particularly with respect to S. monticola, S. obscurus, and S. ornatus. 

Sorex obscurus shumaginensis Merriam, 1900

1900. Sorex alascensis shumaginensis Merriam, Proc. Wash. Acad. Sci. 2:18; 

14 March.

1901. S[orex]. shumaginensis: Osgood, N. Am. Fauna 21:71; 26 September.

1901. [Sorex glacialis] shumaginensis: Elliot, Field Col. Mus. Publ. 45, Zool.

Ser. 2(1):373; 6 March.

1902. Sorex obscurus shumaginensis: J. A. Allen, Bull. Am. Mus. Nat. Hist. 

16:228; 12 July. First use of current name combination.

1955. Sorex vagrans shumaginensis: Findley, Univ. Kansas Publ. Mus. Nat.

Hist. 9(1):42; 10 December.

1977. S[orex]. m[onticolus]. shumaginensis: Hennings and Hoffmann, Occ.

Pap. Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

Holotype: USNM 97993.

Type Locality: USA: “Popof I[slan]d., Shumagin Islands, 

Alaska.” 

Distribution: USA: western Alaska, from the Bering Sea east 

to western Prince William Sound, including the Seward, 

Alaska, and Kenai peninsulas (Alexander, 1996).

Sorex obscurus soperi Anderson and Rand, 1945

1945. Sorex obscurus soperi Anderson and Rand, Can. Field-Nat. 59(2):47; 

16 October.

1955. Sorex vagrans soperi: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):48; 10 December.

1977. S[orex]. m[onticolus]. soperi: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July. 

Holotype: CMN 18249.

Type Locality: Canada: “2 1⁄2 miles northwest of Lake Audy, 

Riding Mountain National Park, southwestern Manitoba, 

Canada; altitude about 1740 feet.” 

Distribution: Canada: eastern Saskatchewan and Manitoba 

(Wrigley et al., 1979; Alexander, 1996).

Sorex oreopolus Merriam, 1892

1892. Sorex oreopolus Merriam, Proc. Biol. Soc. Wash. 7:173; 29 

September.

1959. Sorex oreopolus oreopolus: Hall and Kelson, Mammals of North 

America, p. 49.

Holotype: USNM 33663/45698.

Type Locality: Mexico: “the Sierra de Colima, Jalisco, Mexico 

(altitude, 10,000 feet).” 

Distribution: Mexico: discontinuous distribution in highlands 

of Jalisco, México, Distrito Federal, Morelos, Tlaxcala, and 

Puebla; elevational distribution: 2,440–3,965 m (Carraway, 

2007).

Remarks: Subgenus Otisorex; Sorex oreopolus group (Esteva 

et al., 2010, clade A3). Within the S. oreopolus group, 

S. oreopolus is sister to S. ventralis (Esteva et al., 2010).

Sorex orizabae Merriam, 1895

1895. Sorex orizabae Merriam, N. Am. Fauna 10:71; 31 December.

1928. Sorex vagrans orizabae: Jackson, N. Am. Fauna 51:113; 24 July.

Holotype: USNM 53633.

Type Locality: Mexico: “Mount Orizaba, state of Puebla, 

Mexico (altitude, 9,500 ft).”

Distribution: Mexico: highlands in Michoacán, México, Distrito Federal, Morelos, Tlaxcala, Puebla, and Veracruz; elevational distribution: 2,060–4,175 m (Carraway, 2007).

Remarks: Subgenus Otisorex; S. oreopolus group. 
6 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sorex ornatus Merriam, 1895

Remarks: Subgenus Otisorex; Sorex vagrans group. Molecular 

analyses by Maldonado et al. (2001, 2004) demonstrated 

the existence of three genetic clades (northern, central, and 

southern) whose geographical distributions do not correspond entirely to those of morphologically defined subspecies of S. ornatus. They showed that S. ornatus sinuosus and 

some northern populations of S. o. californicus are more 

closely related to S. vagrans and that S. monticola parvidens

is more closely related to S. ornatus. The latter point was 

also demonstrated morphologically by Woodman (2012a). 

Sorex ornatus californicus Merriam, 1895

1895. Sorex californicus Merriam, N. Am. Fauna 10:80; 31 December.

1912. Sorex californicus californicus: Miller, Bull. U.S. Natl. Mus. 79:18; 31 

December.

1922. Sorex ornatus californicus: Jackson, J. Wash. Acad. Sci. 12:264; 

4 June. First use of current name combination.

1928. Sorex ornatus ornatus: Jackson, N. Am. Fauna 51:164; 24 July. Part; 

not S. ornatus ornatus Merriam, 1895.

Holotype: USNM 32578/44426.

Type Locality: USA: “Walnut Creek, Contra Costa County, Calif.”

Distribution: USA: west central California south of San Pablo 

Bay, Suisun Bay, and the San Joaquin River to central Santa 

Barbara and southern Kern Counties (Maldonado et al., 

2001, 2004); elevational range: from near sea level to 950 m 

(Grinnell, 1933).

Remarks: Sorex ornatus californicus comprises the bulk of 

Maldonado et al.’s (2001, 2004) central clade, which also 

includes S. o. relictus and S. o. salarius. Its distribution now 

excludes a large northern population of the former S. o.

californicus, which was demonstrated to be more closely 

related to S. v. vagrans. The geographic distribution of S. o.

californicus now extends farther south to include populations previously considered to be S. o. ornatus. 

Sorex ornatus juncensis Nelson and Goldman, 1909

1909. Sorex californicus juncensis Nelson and Goldman, Proc. Biol. Soc.

Wash. 22:27; 10 March.

1917. Sorex californicus jucensis: Elliot, Checkl. Mamm. Suppl., p. 146. Incorrect subsequent spelling of Sorex californicus juncensis Nelson and 

Goldman, 1909.

1928. Sorex juncensis: Jackson, N. Am. Fauna 51:172; 24 July.

1981. Sorex o[rnatus]. juncensis: Junge and Hoffmann, Occ. Pap. Mus.

Nat. Hist. Univ. Kansas 94:34; 5 November. First use of current name 

combination.

Holotype: USNM 139594.

Type Locality: Mexico: “Socorro, 15 miles south of San Quintin, Lower California, Mexico.” 

Distribution: Mexico: known only from marshes along a short 

section of the Pacific Coast of northern Baja California 

(Hall, 1981; Maldonado et al., 2001, 2004).

Remarks: This taxon forms part of the southern clade of Sorex 

ornatus; hence, it is closely allied with S. ornatus ornatus

(Maldonado et al., 2001, 2004).

Sorex ornatus lagunae Nelson and Goldman, 1909

1909. Sorex lagunae Nelson and Goldman, Proc. Biol. Soc. Wash. 22:27; 

10 March.

1928. Sorex ornatus lagunae: Jackson, N. Am. Fauna 51:169; 24 July. First 

use of current name combination.

Holotype: USNM 147119.

Type Locality: Mexico: “La Laguna, Sierra Laguna, Lower 

California, Mexico (5,500 ft.).”

Distribution: Mexico: known only from extreme southern Baja 

California.

Remarks: This taxon is part of Maldonado et al.’s (2001, 2004) 

southern clade of Sorex ornatus, which includes S. ornatus 

ornatus.

Sorex ornatus ornatus Merriam, 1895

1895. Sorex ornatus Merriam, N. Am. Fauna 10:79; 31 December.

1903. Sorex oreinus Elliot, Field Col. Mus. Publ. 74, Zool. Ser. 3(10):172; 7 

May. Holotype: FMNH 10842, from “Aguaje de las Fresas, San Pedro 

Martir mountains, Lower California, Mexico, 6,000 feet elevation.”

1903. Sorex orinus Elliot, Field Col. Mus. Publ. 74, Zool. Ser. 3(12):228; 

June. Justified emendation of Sorex oreinus Elliot, 1903.

1928. Sorex ornatus ornatus: Jackson, N. Am. Fauna 51:166; 24 July. First 

use of current name combination.

Holotype: USNM 31333/43198.

Type Locality: USA: “head of San Emigdio Canyon, Mount 

Piños, California”; Kern County. 

Distribution: USA and Mexico: from about central Santa Barbara County and southern Kern County in west central California to northwestern Baja California (Maldonado et al., 

2001, 2004); elevational distribution: from near sea level to 

2,750 m (Grinnell, 1933).

Remarks: Sorex ornatus ornatus comprises the bulk of Maldonado et al.’s (2001, 2004) southern clade, which also 

includes S. o. lagunae, S. o. parvidens, S. o. salicornicus, 

and S. o. willetti. The distribution of S. o. ornatus excludes 

a large northern population that is now referred to S. o.

californicus.

Sorex ornatus parvidens Jackson, 1921

1921. Sorex obscurus parvidens Jackson, J. Mamm. 2(3):161; 19 August.

1895. Sorex ornatus: Merriam, N. Am. Fauna 10:79; 31 December. Part; not 

Sorex ornatus ornatus Merriam, 1895.

1955. Sorex vagrans parvidens: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):58; 10 December.

1977. S[orex]. m[onticolus]. parvidens: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

2001. S[orex]. o[rnatus]. ornatus (southern clade): Maldonado et al., Molecular Ecology 10:129.
NUMBER 650 • 6 7

2012. Sorex ornatus parvidens: Woodman, J. Mamm. 93(3):837; 28 June. 

First use of current name combination.

Holotype: USNM 56561.

Type Locality: USA: “Spring known as Thurman’s Camp, Bluff 

Lake, altitude 7,500 feet, western side of San Bernardino 

Peak, San Bernardino Mountains, California”; corrected by 

Jackson (1928:124) to “Spring known as Thurmans Camp, 

Bluff Lake, altitude about 7,500 feet, San Bernardino 

Mountains, Calif.” 

Distribution: USA: San Bernardino and San Gabriel Mountains in San Bernardino and Los Angeles Counties, California; elevational distribution: up to 2,286 m (Woodman, 

2012a).

Remarks: Maldonado et al. (2001, 2004) showed genetically 

that shrews of the S. vagrans group from the San Bernardino 

Mountains are part of their southern clade of S. ornatus. 

Woodman (2012a) demonstrated S. obscurus parvidens is 

also morphologically closest to S. ornatus. 

Sorex ornatus relictus Grinnell, 1932

1932. Sorex ornatus relictus Grinnell, Univ. Calif. Publ. Zool. 38(8):389; 

9 June.

Holotype: MVZ 51414.

Type Locality: USA: “excavated slough immediately outside of 

east side levee, Buena Vista Lake, 290 feet altitude, Kern 

County, California.” 

Distribution: USA: formerly found in marshlands associated 

with lakes in the southern San Joaquin Valley in Kings and 

Kern Counties, California; elevational distribution: below 

100 m (Grinnell, 1933).

Remarks: This taxon forms part of the central clade of S. ornatus; hence, it is closely allied with S. ornatus californicus

(Maldonado et al., 2001, 2004).

Sorex ornatus salarius von Bloeker, 1939

1939. Sorex ornatus salarius von Bloeker, Proc. Biol. Soc. Wash. 52:94; 

5 June.

Holotype: MVZ 81548.

Type Locality: USA: “the salt marsh at the mouth of the Salinas 

River, Monterey County, California.” 

Distribution: USA: known only from the coastal region around 

Monterey Bay, Monterey County, California; elevational 

distribution: from near sea level up to 400 m (Hall, 1981; 

Maldonado et al., 2001, 2004).

Remarks: This taxon belongs to Maldonado et al.’s (2001, 

2004) central clade of S. ornatus and is closely allied with 

S. ornatus californicus.

Sorex ornatus salicornicus von Bloeker, 1932

1932. Sorex ornatus salicornicus von Bloeker, Proc. Biol. Soc. Wash. 45:131; 

9 September.

Holotype: MVZ 74679.

Type Locality: USA: “Playa del Rey, Los Angeles County, California”; the original skin tag of the holotype gives the collection locality more accurately as “1 mi. N. of La Playa del 

Rey, Los Angeles Co., California” and provides an elevation 

of “25 ft.” (Chris J. Conroy, MVZ, email communication, 

23 June 2017).

Distribution: USA: known from coastal marshes in Ventura 

and Los Angeles Counties, southwestern California (Grinnell, 1933).

Remarks: This taxon is part of the southern clade of S. ornatus, 

which includes S. ornatus ornatus (Maldonado et al., 2001, 

2004).

Sorex ornatus willetti von Bloeker, 1942

1942. Sorex willetti von Bloeker, Bull. S. Cal. Acad. Sci. 40(3):163; 

31 January.

1967. Sorex o[rnatus]. willetti: von Bloeker, Proceedings of the Symposium 

on the Biology of the California Islands (Santa Barbara: Santa Barbara 

Botanic Garden), p. 247; 18 August.

Holotype: LACM 7400.

Type Locality: USA: “Avalon Canyon, Santa Catalina Island, 

Los Angeles County, California.”

Distribution: USA: known only from Santa Catalina Island, 

California.

Remarks: This taxon is part of the southern clade of S. ornatus, 

and it is closely allied with S. ornatus ornatus (Maldonado 

et al., 2001, 2004).

Sorex pacificus Coues, 1877

Remarks: Subgenus Otisorex. Within the Sorex sonomae

group, S. pacificus is sister to S. sonomae (Demboski and 

Cook, 2001; Hope et al., 2014). Phylogenetic analysis of 

the mitochondrial cytochrome b gene indicated that S. monticola as conceived by Alexander (1996) is not monophyletic but is composed of three genetic clades (Demboski and 

Cook, 2001; Shafer and Stewart, 2007): “southern continental clade” (recognized herein as Sorex monticola [sensu 

stricto]), “northern continental clade” (S. obscurus), and 

“coastal clade” (S. pacificus). Subspecies included in S. pacificus herein are based on my interpretations of Carraway 

(1990) and Alexander (1996) applied to the framework of 

Demboski and Cook (2001), which showed S. bairdi and 

S. pacificus deeply embedded among what had been considered the Pacific coastal subspecies of S. monticola (sensu 

lato) (Alexander, 1996). 

Sorex pacificus bairdi Merriam, 1895

1895. Sorex bairdi Merriam, N. Am. Fauna 10:77; 31 December.

1918. Sorex obscurus bairdi: Jackson, Proc. Biol. Soc. Wash. 31:127; 

29 November.

1955. Sorex vagrans bairdi: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):35; 10 December.
6 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1977. S[orex]. m[onticolus]. bairdii: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July. Incorrect subsequent spelling of Sorex bairdi Merriam, 1895.

1990. Sorex bairdii bairdii: Carraway, Spec. Publ. Mus. Texas Tech Univ. 

32:39; 7 July. Incorrect subsequent spelling of Sorex bairdi Merriam, 

1895. 

1996. Sorex bairdi bairdi: Alexander, Univ. Kansas Mus. Nat. Hist., Misc.

Publ. 88:22; 21 February. 

Holotype: USNM 17414/24318.

Type Locality: USA: “Astoria, Oregon”; Clatsop County.

Distribution: USA: northwestern Oregon from the Columbia 

River south to Lane County; elevational distribution: 70–

490 m (Carraway, 1990).

Sorex pacificus calvertensis Cowan, 1941

1941. Sorex obscurus calvertensis Cowan, Proc. Biol. Soc. Wash. 54:103; 

31 July.

1955. Sorex vagrans calvertensis: Findley, Univ. Kansas Publ. Mus. Nat.

Hist. 9(1):39; 10 December.

1977. S[orex]. m[onticolus]. calvertensis: Hennings and Hoffmann, Occ.

Pap. Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

Holotype: RBCM 1947.

Type Locality: Canada: “Safety Cove, Calvert Island, B.C.”

Distribution: Canada: widely separated Banks Island and Calvert Island, British Columbia (Alexander, 1996).

Sorex pacificus cascadensis Carraway, 1990

1990. Sorex pacificus cascadensis Carraway, Spec. Publ. Mus. Texas Tech.

Univ. 32:55; 7 July.

Holotype: USNM 204479.

Type Locality: USA: “McKenzie Bridge, [Lane County], Oregon” (brackets in original).

Distribution: USA: northeastern Linn County south to Jackson County, Oregon; elevational distribution: 425–1,400 m 

(Carraway, 1990).

Sorex pacificus elassodon Osgood, 1901

1901. Sorex longicauda elassodon Osgood, N. Am. Fauna 21:35; 

26 September.

1905. Sorex obscurus elassodon: Elliot, Field Col. Mus. Publ. 105, Zool. Ser. 

6:450; 6 December.

1955. Sorex vagrans elassodon: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):40; 10 December.

1977. S[orex]. m[onticolus]. elassodon: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

Holotype: USNM 100597.

Type Locality: Canada: “Cumshewa Inlet, Moresby Island, 

Queen Charlotte Islands, British Columbia.” 

Distribution: Canada and USA: individual coastal islands 

from Baranof, Kupreanof, and Mitkof Islands, Alaska, to 

Porcher and Dewdney Islands and part of the Haida Gwaii 

Archipelago (formerly Queen Charlotte Islands), British Columbia (Alexander, 1996).

Sorex pacificus insularis Cowan, 1941

1941. Sorex obscurus insularis Cowan, Proc. Biol. Soc. Wash. 54:103; 31 

July. Not Sorex insularis Okhotina, 1993:64.

1955. Sorex vagrans insularis: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):39; 10 December.

1977. S[orex]. m[onticolus]. insularis: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

Holotype: RBCM 3110.

Type Locality: Canada: “Smythe Island, Bardswell group, 

B. C.”

Distribution: Canada: the western portion of the Bardswell Island Group (including Athlone [formerly Smythe or Smyth], 

Townsend, and Reginald Islands), along the west coast of 

British Columbia (Alexander, 1996).

Sorex pacificus isolatus Jackson, 1922

1922. Sorex obscurus isolatus Jackson, J. Wash. Acad. Sci. 12:263; 4 June.

1955. Sorex vagrans isolatus: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):38; 10 December.

1977. S[orex]. m[onticolus]. isolatus: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

Holotype: USNM 177719.

Type Locality: Canada: “Mouth of Millstone Creek, Nanaimo, 

Vancouver island, British Columbia.” 

Distribution: Canada: Vancouver Island and numerous smaller 

islands (including Denman Island) along the coast of Vancouver Island, British Columbia.

Sorex pacificus longicauda Merriam, 1895

1895. Sorex obscurus longicauda Merriam, N. Am. Fauna 10:74; 

31 December.

1900. S[orex]. longicauda: Merriam, Proc. Wash. Acad. Sci. 2:16; 

14 March.

1902. [Sorex obscurus] longicaudus: Elliot, Field Col. Mus. Publ. 45, Zool.

Ser. 2:372. Unjustified emendation of Sorex obscurus longicauda Merriam, 1895, a noun in apposition.

1955. Sorex vagranslongicauda: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):37; 10 December.

1977. S[orex]. m[onticolus]. longicauda: Hennings and Hoffmann, Occ.

Pap. Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

1996. Sorex monticolus longicaudus: Alexander, Univ. Kansas Mus. Nat.

Hist., Misc. Publ. 88:28; 21 February. Unjustified emendation of Sorex 

obscurus longicauda Merriam, 1895, a noun in apposition. 

Holotype: USNM 74711.

Type Locality: USA: “Wrangel, southeast Alaska.” 

Distribution: Canada and USA: Pacific coastal region and adjacent islands from Taku Inlet, southeast Alaska, to Rivers 

Inlet, British Columbia; includes Pitt, Campania, Princess 

Royal, Swindle, Dufferin, Horsfall, Campbell, Spider, and 
NUMBER 650 • 6 9

the Hunter group of islands; excludes Porcher, the Estevan 

group, the western Bardswell group, and Goose and Hecate 

Islands (Alexander, 1996). 

Sorex pacificus malitiosus Jackson, 1919

1919. Sorex obscurus malitiosus Jackson, Proc. Biol. Soc. Wash. 32:23; 

11 April.

1955. Sorex vagrans malitiosus: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):40; 10 December.

1977. S[orex]. m[onticolus]. malitiosus: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

Holotype: MVZ 8401.

Type Locality: USA: “east side of Warren Island, Alaska.” 

Distribution: USA: Known only from Warren and Coronation 

Islands, along the Pacific Coast of southeastern Alaska (Alexander, 1996).

Sorex pacificus pacificus Coues, 1877

1877. Sorex pacificus Coues, Bull. U.S. Geol. Geog. Surv. Terr. 3(3):650; 

15 May.

1918. Sorex yaquinae Jackson, Proc. Biol. Soc. Wash. 31:127; 29 November. Holotype: USNM 73051, from “Yaquina Bay, Oregon”; Lincoln 

County, USA.

1921. Sorex pacificus pacificus: Jackson, J. Mamm. 2(3):162; 19 August. 

First use of current name combination.

1936. Sorex pacificus yaquinae: Bailey, N. Am. Fauna 55:364; June. 

1955. Sorex vagrans pacificus: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):34; 10 December.

1955. Sorex vagrans yaquinae: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):34; 10 December.

Holotype: USNM 3266.

Type Locality: USA: “Fort Umpqua, Oregon”; mouth of the 

Umpqua River, Douglas County.

Distribution: USA: Pacific coastal region west of the Cascades, 

from Cascade Head in Tillamook County to Coos Bay in 

Coos County, Oregon (Carraway, 1990).

Sorex pacificus permiliensis Jackson, 1918

1918. Sorex obscurus permiliensis Jackson, Proc. Biol. Soc. Wash. 31:128; 

29 November.

1955. Sorex vagrans permiliensis: Findley, Univ. Kansas Publ. Mus. Nat.

Hist. 9(1):36; 10 December.

1977. S[orex]. m[onticolus]. permiliensis: Hennings and Hoffmann, Occ.

Pap. Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

1990. Sorex bairdii permiliensis: Carraway, Spec. Publ. Mus. Texas Tech 

Univ. 32:40; 7 July. 

Holotype: USNM 91048.

Type Locality: USA: “Permilia Lake, west base of Mount Jefferson, Cascade Range, Oregon”; Marion County. 

Distribution: USA: Cascade Range of northern Oregon, from 

the Columbia River south through Lane County; elevational 

distribution: 325–1,400 m (Carraway, 1990).

Sorex pacificus prevostensis Osgood, 1901

1901. Sorex longicauda prevostensis Osgood, N. Am. Fauna 21:35; 

26 September.

1905. Sorex obscurus prevostensis: Elliot, Field Col. Mus. Publ. 105, Zool.

Ser. 6:450; 6 December

1955. Sorex vagrans prevostensis: Findley, Univ. Kansas Publ. Mus. Nat.

Hist. 9(1):41; 10 December.

1977. S[orex]. m[onticolus]. prevostensis: Hennings and Hoffmann, Occ.

Pap. Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

Holotype: USNM 100618.

Type Locality: Canada: “Prevost Island, Queen Charlotte Islands, British Columbia.” 

Distribution: Canada: known only from Kunghit Island (formerly Prevost Island) at the southern tip of the Haida Gwaii 

Archipelago (formerly Queen Charlotte Islands; Alexander, 

1996).

Remarks: Apparently, there were two Prevost Islands in British Columbia. One at the southern tip of the Haida Gwaii 

Archipelago is now called Kunghit Island. A second is located in the channel between the southeastern tip of Vancouver Island and the northwestern corner of Washington. 

Sorex pacificus prevostensis occurs on Kunghit Island.

Sorex pacificus setosus Elliot, 1899

1899. Sorex setosus Elliot, Field Col. Mus. Publ. 32, Zool. Ser. 1(13):274; 

17 May.

1918. Sorex obscurus setosus: Jackson, Proc. Biol. Soc. Wash. 31:127; 

29 November.

1938. Sorex obscurus mixtus Hall, Am. Nat. 72(742):462; 10 September. 

Holotype: MVZ 70376, from “Vanada, Texada Island, Georgia Strait, 

British Colombia,” Canada.

1955. Sorex vagrans setosus: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):36; 10 December.

1955. Sorex vagrans mixtus: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):38; 10 December.

1977. S[orex]. m[onticolus]. setosus: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

1977. S[orex]. m[onticolus]. mixtus: Hennings and Hoffmann, Occ. Pap.

Mus. Nat. Hist. Univ. Kansas 68:4; 15 July.

Holotype: FMNH 6213.

Type Locality: USA: “vicinity of Happy Lake,” Olympic 

Mountains, Clallam County, Washington.

Distribution: Canada and USA: coastal region and western 

slope of the Cascade Mountains from Rivers Inlet in southwestern British Columbia through western Washington and 

northwestern Oregon; elevational distribution from near 

sea level to 1,525 m (Hennings and Hoffmann, 1977; Alexander, 1996).

Sorex palustris Richardson, 1828

Remarks: Subgenus Otisorex; Sorex palustris group. Hope et 

al. (2014) recognized three species-level molecular clades 

among the subspecies formerly united as Sorex palustris:
7 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

eastern S. albibarbis, cordilleran S. navigator, and boreal 

S. palustris (sensu stricto). That study limited S. palustris 

(sensu stricto) to include the subspecies S. palustris hydrobadistes and S. p. palustris. Nagorsen et al. (2017) demonstrated that parapatric S. navigator and S. palustris can 

also be distinguished morphometrically. However, the taxonomic status of many southern and Midwestern populations remains incompletely understood (Hope et al., 2014).

Sorex palustris hydrobadistes Jackson, 1926

1926. Sorex palustris hydrobadistes Jackson, J. Mamm. 7(1):57; 15 February.

Holotype: USNM 229061.

Type Locality: USA: “Withee, Clark County, Wisconsin.” 

Distribution: Extreme northeastern South Dakota, central 

Minnesota, northern Wisconsin, the Upper Peninsula of 

Michigan, and the northern Lower Peninsula of Michigan 

(Hall, 1981; Beneski and Stinson, 1987; Hope et al., 2014); 

elevational distribution: 140–1,100 m.

Sorex palustris palustris Richardson, 1828

1828. Sorex palustris Richardson, Zool. J. 3:517; April.

1838. Amphisorex palustris: Gray, Proc. Zool. Soc. Lond. 5(1837):125; 

14 June.

1847. Crossopus palustris: Reichenbach, Praktische Naturgeschichte Menschen und der Säugethiere für Gebildete aller Stände (Leipzig: Gebhardt 

& Reisland), p. 161 [reference not seen].

1848. [Galemys (Crossopus)] palustris: Pomel, Arch. Sci. Phys. Nat. 9:249. 

1863. Neosorex palustris: Verrill, Proc. Boston Soc. Nat. Hist. 9:167; 

February.

1868. Crossopus fimbripes: Fitzinger, Sitzungsber. Math.-natürwissensch.

Classe Kaiserl. Akad. Wissensch. 57(1):630. Not Sorex fimbripes Bachman, 1837.

1895. Sorex (Neosorex) palustris: Merriam, N. Am. Fauna 10:91; 

31 December.

1901. Neosorex palustrius Elliot, Field Col. Mus. Publ. 45, Zool. Ser. 

2(1):378, fig. 72; 6 March. Incorrect subsequent spelling of Sorex palustris Richardson, 1828.

1915. Neosorex palustris palustris: G. M. Allen, Proc. Biol. Soc. Wash. 

28:17; 12 February.

1926. Sorex palustris palustris: Jackson, J. Mamm. 7(1):57; 15 February. 

First use of current name combination.

Holotype: BM 1842.10.7.1.

Type Locality: Canada: “Marshy places, from Hudson’s Bay to 

the Rocky Mountains.” The holotype was “procured on Capt. 

Franklin’s late Expedition.” John Richardson participated in 

John Franklin’s 1825 Canadian expedition, which traveled 

north down the Mackenzie River, then east (Richardson) or 

west (Franklin) along the coast of the Beaufort Sea. The expedition thereby passed through portions of the Northwest 

Territories, Yukon Territory, Nunavut, and Alaska.

Distribution: Canada and USA: southern Northwest Territory 

and Alberta east of the crest of the Rocky Mountains to 

northern Minnesota and central Ontario; elevational distribution: 15–2,819 m (Hall, 1981; Beneski and Stinson, 

1987; Hope et al., 2014).

Sorex preblei Jackson, 1922

1922. Sorex preblei Jackson, J. Wash. Acad. Sci. 12:263; 4 June.

Holotype: USNM 208032.

Type Locality: USA: “Jordan Valley, altitude, 4,200 feet, Malheur County, Oregon.” 

Distribution: Canada and USA: south central British Columbia, southeastern Washington, central Oregon, and the 

northeastern corner of California east through Montana, 

western Wyoming, and northern Utah; elevational distribution: 343–3,135 m (Cornely et al., 1992; Nagorsen et al., 

2001). 

Remarks: Subgenus Otisorex; clade A2 of Esteva et al. (2010). 

Within the Beringian clade of the Sorex cinereus group, 

S. preblei is a member of a subclade that also contains 

S. lyelli and two undescribed western North American species (Hope et al., 2012). 

Sorex pribilofensis Merriam, 1895

1895. Sorex pribilofensis Merriam, N. Am. Fauna 10:87; 31 December. 

1981. Sorex hydrodromus: Junge and Hoffmann, Occ. Pap. Mus. Nat. Hist.

Univ. Kansas 94:18; 5 November. Not Sorex hydrodromus Dobson, 

1889.

Holotype: USNM 30911.

Type Locality: USA: “St. Paul Island, Pribilof Islands, Bering 

Sea”; Alaska. 

Distribution: Known only from St. Paul Island, Alaska.

Remarks: Subgenus Otisorex; S. cinereus group (Demboski 

and Cook, 2003). Within the Beringian clade of the S. cinereus group, S. pribilofensis is a member of a subclade 

that that is sister to S. haydeni and that also includes 

North American S. jacksoni and S. ugyunak and Siberian 

S. camtschatica Yudin, 1972 and S. portenkoi Stroganov, 

1956 (Hope et al., 2012).

Sorex pribilofensis has sometimes been placed in synonymy with S. hydrodromus Dobson, 1889, which was 

described from a specimen (RAS 2389) obtained from “Unalaska” by I. G. Vosnesensky in 1840–1848 (Hall in Murie, 

1959) and preserved in the Zoological Institute of the Russian Academy of Sciences, St. Petersburg (Baranova et al., 

1981). Sorex hydrodromus was described as a form of water 

shrew with the characteristic “thick comb-like fringe of stiff 

hairs . . . along the outer and inner margin of both manus 

and pes” (Dobson, 1889:373). Later investigators, however, 

variably identified the holotype as (1) “a recognizable kind 

(species or subspecies) of Sorex best placed in the arcticus

group” (Hall in Murie, 1959; see also Jackson, 1928:75), 

(2) synonymous with S. caecutiens (Gromov et al., 1963), 

or (3) “virtually indistinguishable from S. pribilofensis” 
NUMBER 650 • 7 1

(Hoffmann and Peterson, 1967:131–132). The type locality 

of “Unalaska” has been narrowly interpreted to mean Unalaska Island (Hall in Murie, 1959), but no shrews inhabit 

that island (Peterson, 1967). More logically, it refers to the 

old Russian Unalaska District, a region that included the 

Shumagin Islands, the western Alaska Peninsula, Unimak 

Island, the Fox Islands, and the Pribilof Islands (Peterson, 

1967; Rausch and Rausch, 1997). Hall (in Murie, 1959) reported cranial measurements (in mm) from the holotype and 

a second specimen (RAS 2370) of S. hydrodromus collected 

by Vosnesensky from the type locality (condylobasal length: 

16.1, 16.0; cranial breadth: 8.2, 8.8; maxillary breadth: 

4.45, 4.7, respectively). Comparison of these measurements 

with those of S. pribilofensis reported by Hoffmann and 

Peterson (1967; mean and range of measurements for 12 to 

14 individuals (mm): condylobasal length, 15.5, 15.2–15.8; 

cranial breadth, 7.9, 7.8–8.0; maxillary breadth, 4.7, 4.5–

4.9) suggests that S. hydrodromus has slightly larger cranial 

size than S. pribilofensis but a relatively narrower maxillary 

breadth. The holotype of S. hydrodromus is also reported to 

have a substantially longer tail (either 42 or 46 mm) than S.

pribilofensis (mean ± SD for 15 individuals (mm): 34 ± 1; 

van Zyll de Jong, 1982). A better match for S. hydrodromus

may be S. camtschatica Yudin, 1972, which is closer in cranial size (mean ± SD for 34 individuals (mm): condylobasal 

length, 16.2 ± 2; cranial breadth, 8.0 ± 2; breadth across 

M2 – M2

, 3.9 ± 1; van Zyll de Jong, 1982) and is reported 

to possess a “well-developed fringe of stiff hair on the feet” 

(van Zyll de Jong, 1982:1585).

Sorex rohweri Rausch, Feagin, and Rausch, 2007

2007. Sorex rohweri Rausch, Feagin, and Rausch, Mamm. Biol. 72(2):93; 

26 March.

Holotype: UWBM 39812.

Type Locality: USA: “Olympic Peninsula, Washington State, 

near Quilcene, Penny Creek, at lat. 47°51′ N, long. 

122°57′ W.” in Jefferson County.

Distribution: Canada and USA: occurs from the western Cascades to the coast in southwestern British Columbia (south 

of the lower Frasier River) and western Washington and 

west of the Willamette Valley in Oregon; elevational distribution from near sea level to 1,250 m (Rausch et al., 2007; 

Nagorsen and Panter, 2009; Woodman and Fisher, 2016).

Remarks: Subgenus Otisorex; S. rohweri can be considered 

either the most basal branch within the Sorex cinereus

group or the sister taxon to S. longirostris plus the Sorex 

cinereus group (Hope et al., 2012).

Sorex salvini Merriam, 1897

Remarks: Unnamed subgenus; S. salvini group. Esteva et al. 

(2010) showed that S. salvini (as S. veraecrucis locality 14 

and S. saussurei locality 15) is sister to a clade formed by 

S. altoensis (as S. veraecrucis localities 5, 7) and S. saussurei

(as S. saussurei locality 6). Together, these taxa form part of 

Esteva et al.’s (2010) subclade S2 of clade B1.

Matson and Ordóñez-Garza (2017) recognized S. cristobalensis and S. salvini as species, but they exhibit only 

4.0% genetic sequence divergence between them for cytochrome b (Esteva et al., 2010). For this reason, I tentatively 

treat them as subspecies herein, but these taxa should be 

restudied using addition genetic markers. Unfortunately, 

S. salvini oaxacae was not included in Esteva et al.’s (2010) 

study. All three taxa were treated previously as subspecies of S. saussurei and S. veraecrucis. Esteva et al. (2010) 

showed high sequence divergence between S. s. cristobalensis (as S. veraecrucis locality 14) and both (1) S. saussurei

(as S. saussurei locality 6: 10.7% divergence) and (2) S. veraecrucis (as S. veraecrucis locality 11: 16.0%). They also 

demonstrated high sequence divergence between S. s. salvini (as S. saussurei locality 15) and both (1) S. saussurei

(10.1%) and (2) S. veraecrucis (14.6%).

Sorex salvini cristobalensis Jackson, 1925

1925. Sorex saussurei cristobalensis Jackson, Proc. Biol. Soc. Wash. 38:129; 

13 November.

2007. Sorex veraecrucis cristobalensis: Carraway, Monogr. West. N. Am. 

Nat. 3:62; 22 June.

2010. S[orex]. veraecrucis (14): Esteva et al., Zootaxa 2615:55, fig. 1; 17 September. Part; not Sorex saussurei veraecrucis Jackson, 1925.

2012. S[orex]. salvini cristobalensis: Woodman et al., Ann. Carnegie Mus. 

80(3):214; 15 July. First use of current name combination.

2017. Sorex cristobalensis: Matson and Ordóñez-Garza, Zootaxa 4236(3): 

475; 24 February.

Holotype: USNM 75883.

Type Locality: Mexico: “San Cristobal, altitude 8,400 feet, 

state of Chiapas.”

Distribution: Mexico: known from three localities in the highlands of Chiapas; elevational distribution: 1,900–2,560 m 

(Carraway, 2007; Matson and Ordóñez-Garza, 2017).

Sorex salvini oaxacae Jackson, 1925

1925. Sorex saussurei oaxacae Jackson, Proc. Biol. Soc. Wash. 38:128; 

13 November.

2007. Sorex veraecrucis oaxacae: Carraway, Monogr. West. N. Am. Nat. 

3:62; 22 June.

2012. S[orex]. salvini oaxacae: Woodman et al., Ann. Carnegie Mus. 

80(3):214; 15 July. First use of current name combination.

Holotype: USNM 71467.

Type Locality: Mexico: “Mountains near Ozolotepec, altitude 

10,000 feet, state of Oaxaca.” 

Distribution: Mexico: highlands of Oaxaca; elevational distribution: 1,600–3,050 m (Carraway, 2007).

Sorex salvini salvini Merriam, 1897

1897. Sorex salvini Merriam, Proc. Biol. Soc. Wash. 11:229; 15 July.
7 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1897. Sorex godmani Merriam, Proc. Biol. Soc. Wash. 11:229; 15 July. 

Holotype: USNM 77044, from “Volcano Santa Maria, Quezaltenango, 

Guatemala (alt. 9,000 ft. = 2740 meters).”

1904. [Sorex] godmanni Trouessart, Cat. Mamm. Suppl. 1904:135. Incorrect 

subsequent spelling of Sorex godmani Merriam, 1897.

1928. Sorex saussurei godmani: Jackson, N. Am. Fauna 51:158; 24 July.

1928. Sorex saussurei salvini: Jackson, N. Am. Fauna 51:159; 24 July.

2010. S[orex]. saussurei (15): Esteva et al., Zootaxa 2615:55, fig. 1; 17 September. Part; not Sorex saussurei Merriam, 1892.

2012. S[orex]. salvini salvini: Woodman et al., Ann. Carnegie Mus. 80(3):214; 

15 July. First use of current name combination.

Holotype: USNM 77035.

Type Locality: Guatemala: “Calel, Totonicapán, Guatemala 

(alt. 10200 ft. = 3100 meters).”

Distribution: Guatemala: highlands of southwestern and central Guatemala; elevational distribution: 2,040–3,110 m 

(Matson and Ordóñez-Garza, 2017).

Sorex saussurei Merriam, 1892

1892. Sorex saussurei Merriam, Proc. Biol. Soc. Wash. 7:173; 29 September.

1905. Sorex saussurii Elliot, Field Col. Mus. Publ. 105, Zool. Ser. 6:456; 

6 December. Incorrect subsequent spelling of Sorex saussurei Merriam, 

1892.

1912. Sorex saussurei saussurei Miller, Bull. U.S. Natl. Mus. 79:19; 31 December. Part.

Holotype: USNM 33667/45702.

Type Locality: Mexico: “the Sierra de Colima, Jalisco, Mexico 

(altitude 8,000 feet).” 

Distribution: Mexico: highlands in Jalisco, Colima, Michoacán, México, Distrito Federal, Morelos, and Puebla; elevational distribution: 2,370–3,180 m (Carraway, 2007). 

Remarks: Unnamed subgenus; S. saussurei group. Sorex saussurei (as S. saussurei locality 6) is sister to S. altoensis (as 

S. veraecrucis localities 5, 7), and together, they form part of 

Esteva et al.’s (2010) subclade S2 of clade B1.

Sorex sclateri Merriam, 1897

1897. Sorex sclateri Merriam, Proc. Biol. Soc. Wash. 11:228; 15 July.

Holotype: USNM 75872.

Type Locality: Mexico: “Tumbala, Chiapas, Mexico (alt., 

5,000 ft.).” 

Distribution: Mexico: known only from two localities in the 

highlands of Chiapas, 1,525 m elevation (Carraway, 2007; 

Matson and Ordóñez-Garza, 2017).

Remarks: Unnamed subgenus; S. salvini group (Matson and 

Ordóñez-Garza, 2017).

Sorex sonomae Jackson, 1921

Remarks: Subgenus Otisorex; S. sonomae group. Molecular 

phylogenetic analyses indicate that S. sonomae is sister to 

S. pacificus (Demboski and Cook, 2001; Esteva et al., 2010; 

Hope et al., 2014). 

Sorex sonomae sonomae Jackson, 1921

1921. Sorex pacificus sonomae Jackson, J. Mamm. 2(3):162; 19 August.

1955. Sorex vagrans sonomae: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

9(1):32; 10 December.

1990. Sorex sonomae sonomae: Carraway, 1990, Spec. Publ. Mus. Texas 

Tech Univ. 32:29; 7 July. First use of current name combination.

Holotype: MVZ 19658.

Type Locality: USA: “Sonoma County side of Gualala River, 

Gualala, California.”

Distribution: USA: Pacific Coast from Lincoln and Benton 

Counties, Oregon, south to Marin County, California; elevational distribution: from near sea level to 1,700 m (Carraway, 1990).

Sorex sonomae tenelliodus Carraway, 1990

1990. Sorex sonomae tenelliodus Carraway, 1990, Spec. Publ. Mus. Texas 

Tech Univ. 32:33; 7 July.

Holotype: USNM 565663.

Type Locality: USA: “4 mi. S, 101⁄2 mi. E Blue River, East Fork 

Creek, T17S, R6E, sec. 7, Lane County, Oregon.”

Distribution: USA: Lincoln County, Oregon, south to northern 

Siskiyou County, California; elevational distribution: 160–

1,315 m (Carraway, 1990).

Sorex stizodon Merriam, 1895

1895. Sorex stizodon Merriam, N. Am. Fauna 10:98; 31 December.

Holotype: USNM 75885.

Type Locality: Mexico: “San Cristobal, Chiapas.” 

Distribution: Mexico: known from only two localities near San 

Cristóbal de las Casas in the highlands of northern Chiapas, 

2,740 m elevation (Carraway, 2007; Matson and OrdóñezGarza, 2017).

Remarks: Unnamed subgenus; S. salvini group (Matson and 

Ordóñez-Garza, 2017).

Sorex tenellus Merriam, 1895

1895. Sorex tenellus Merriam, N. Am. Fauna 10:81; 31 December.

1902. Sorex tenellus myops Merriam, Proc. Biol. Soc. Wash. 15:76; 22 

March. Holotype: USNM 29559/41634, from “White Mountains, California.” On the basis of E. W. Nelson’s field catalog, Howell (1923:266) 

corrected the type locality to “Cottonwood Creek (about 9550 feet) on 

east slope of the White Mountains, Inyo County, California.” 

1912. Sorex tenellus tenellus: Miller, Bull. U.S. Natl. Mus. 79:18; 

31 December.

Holotype: USNM 25083/32495.

Type Locality: USA: “summit of Alabama Hills near Lone 

Pine, Owens Valley, Calif[ornia].” On the basis of information in the field catalog of the collector (E. W. Nelson), 

Howell (1923:266) corrected the type locality to “Lone 

Pine Creek (about 4500 feet), Owens Valley, Inyo County, 

California.” 
NUMBER 650 • 7 3

Distribution: USA: highlands in western California (from 

Mount Lassen to the Inyo Mountains) to eastern Nevada 

(from the Ruby Mountains south to the Spring Mountains) 

and western Utah; elevational distribution: 1,370–4,270 m 

(Hoffmann and Owen, 1980; Rickart et al., 2004, 2011, 

2017; Shohfi et al., 2006).

Remarks: Subgenus Otisorex; S. oreopolus group, which is 

part of Esteva et al.’s (2010) clade A3. 

Sorex trowbridgii Baird, 1857

FIGURE 25

Remarks: Unnamed subgenus (George, 1988:456); S. trowbridgii group. This species forms part of Esteva et al.’s 

(2010) clade B1, in which it appears to be sister to their 

subclade S2 (S. salvini and S. saussurei groups). In contrast, 

Maldonado et al. (2015) showed S. trowbridgii to be sister 

to the S. merriami group.

Sorex trowbridgii destructioni Scheffer and Dalquest, 1942

1942. Sorex trowbridgii destructioni Scheffer and Dalquest, J. Mamm. 

23(3):334; 14 August.

Holotype: USNM 271634.

Type Locality: USA: “Destruction Island, Jefferson County, 

Washington.” 

Distribution: USA: known only from Destruction Island, 

Washington (Hall, 1981; George, 1989).

Sorex trowbridgii humboldtensis Jackson, 1922

1922. Sorex trowbridgii humboldtensis Jackson, J. Wash. Acad. Sci. 12(11): 

264; 4 June.

Holotype: USNM 97271.

Type Locality: USA: “Carson’s Camp, Mad River, Humboldt 

Bay, Humboldt County, California.” 

Distribution: USA: coastal region of southern Humboldt and 

Mendocino Counties, California; elevational distribution: 

from near sea level to 700 m (Grinnell, 1933; Hall, 1981; 

George, 1989).

Sorex trowbridgii mariposae Grinnell, 1913

1913. Sorex montereyensis mariposae Grinnell, Univ. Calif. Publ. Zool. 

10(9):189; 20 March.

1923. Sorex trowbridgii mariposae: Grinnell, Univ. Calif. Publ. Zool. 

21(10):314; 27 January. First use of current name combination.

Holotype: MVZ 12979.

Type Locality: USA: “Yosemite Valley at 4,000 feet altitude, Mariposa County, California.” Miller and Kellogg 

(1955:19) restricted the type locality to “near the old 

Sentinel Hotel”; which is approximately 37°44′30′′N, 

119°35′30′′W.

Distribution: USA: from south central Oregon south along 

the higher inner Coast Ranges to southern Mendocino 

County, California, and from western Siskiyou and Modoc 

Counties, California, south through the Warner Mountains and northern Sierra Nevada of eastern California 

and the south central corner of Nevada to Tulare County; 

elevational distribution: 385–2,290 m (Grinnell, 1933; 

Hall, 1981; George, 1989).

Sorex trowbridgii montereyensis Merriam, 1895

1895. Sorex montereyensis Merriam, N. Am. Fauna 10:79; 31 December.

1922. Sorex t[rowbridgii]. montereyensis: Jackson, J. Wash. Acad. Sci. 

12:264; 4 June. First use of current name combination.

Holotype: USNM 32000/44810.

Type Locality: USA: “Monterey, Calif[ornia].”; Monterey 

County. 

Distribution: USA: coastal region of west central California, 

from southern Mendocino County south to southern Santa 

Barbara County; elevational distribution: from near sea 

level to 610 m (Grinnell, 1933; Hall, 1981; George, 1989).

Sorex trowbridgii trowbridgii Baird, 1857

FIGURE 25

1857. Sorex trowbridgii Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):13, pl. 26.

1884. Sorex trowbridgei: True, U.S. Natl. Mus. Circ. 29:22; 29 November. 

Incorrect subsequent spelling of Sorex trowbridgii Baird, 1857.

1922. Sorex trowbridgii trowbridgii: Jackson, J. Wash. Acad. Sci. 12:264; 

4 June. First use of current name combination.

Lectotype: USNM 813/3088, designated by A. J. Poole and 

Schantz (1942:196).

Type Locality: USA: “Astoria, Oregon.” Miller and Rehn 

(1901:239) added that the type locality is at the “mouth of 

Columbia River, Clatsop County.”

Distribution: Canada and USA: southwestern British Columbia 

through western Washington and western Oregon to eastern 

Humboldt and western Trinity Counties, California; elevational distribution: from near sea level to 1,985 m (Grinnell, 

1933; Hall, 1981; George, 1989).

Sorex tundrensis Merriam, 1900

Distribution: Holarctic: in Eurasia, the species ranges from the 

Pechora River in European Russia east through Siberia to 

Chukotka and south to the Altai Mountains, Mongolia and 

northeastern China; in North America, it occurs through 

much of Alaska, the northwestern corner of British Columbia, northern Yukon Territory, and northern Northwest Territories (Hutterer, 2005; Naughton, 2012).

Remarks: Subgenus Sorex; Sorex araneus group (Fumagalli et 

al., 1999). A single subspecies occurs in North America.

Sorex tundrensis tundrensis Merriam, 1900

1900. Sorex tundrensis Merriam, Proc. Wash. Acad. Sci. 2:16; 14 March.

1956. Sorex arcticus tundrensis: Bee and Hall, Univ. Kansas Mus. Nat. Hist.,

Misc. Publ. 8:13; 10 March.

1983. S[orex]. t[undrensis]. tundrensis: Okhotina, Zool. Zh. 62(3):410; 

March. First use of current name combination.
7 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Holotype: USNM 99286.

Type Locality: USA: “St. Michaels, Alaska.”

Distribution: Canada and USA: northern, western, and central 

Alaska, the northwesternmost corner of British Columbia, 

northern Yukon Territory, and northern Northwest Territories (Naughton, 2012).

Sorex ugyunak Anderson and Rand, 1945

1945. Sorex cinereus ugyunak Anderson and Rand, Can. Field-Nat. 59(2):62; 

16 October.

1983. Sorex ugyunak: van Zyll de Jong, Handbook of Canadian Mammals, 

pp. 5, 76. First use of current name combination.

Holotype: AMNH 31365.

Type Locality: Canada: “Tuktuk (Tuktuyaktok), northeast side 

of Mackenzie River delta, south of Toker Point, Mackenzie 

District, Northwest Territories, Canada.” 

Distribution: Canada and USA: tundra areas of northern 

Alaska, northern Yukon Territory, northern Northwest Territories, and Nunavut (Naughton, 2012).

Remarks: Subgenus Otisorex; Beringian clade of the Sorex cinereus group (Hope et al., 2012). Hope et al. (2012) found 

low resolution among high-latitude tundra/taiga species in 

the Beringian clade of the Sorex cinereus group (North American S. jacksoni Hall and Gilmore, 1932; S. pribilofensis Merriam, 1895; and S. ugyunak and Siberian S. camtschatica

Yudin, 1972; S. leucogaster Kuroda, 1933; and S. portenkoi

Stroganov, 1956) that they interpreted as evidence of recent 

speciation. Their analysis indicated S. ugyunak is sister to a 

clade comprising S. camtschatica, S. jacksoni, and S. portenkoi. Despite the relative lack of evidence of paraphyly or 

polyphyly in their cytochrome b tree, they urged caution in 

revising the taxonomic status of these taxa.

Sorex vagrans Baird, 1857

FIGURES 23, 25

Remarks: Subgenus Otisorex; Sorex vagrans group. Molecular 

analyses of S. ornatus and S. vagrans by Maldonado et al. 

(2001, 2004) demonstrated the existence of three genetic 

clades (northern, central, and southern) along the western 

coast of North America. Their northern clade, which corresponds to S. vagrans, includes what were previously called 

S. ornatus sinuosus as well as some northern populations of 

S. ornatus californicus. 

Sorex vagrans halicoetes Grinnell, 1913

1913. Sorex halicoetes Grinnell, Univ. Calif. Publ. Zool. 10(9):183; 

20 March.

1928. Sorex vagrans halicoetes Jackson, N. Am. Fauna 51:108; 24 July. First 

use of current name combination.

Holotype: MVZ 3638.

Type Locality: USA: “salt marsh near Palo Alto, Santa Clara 

County, California.” 

Distribution: USA: salt marshes bordering the southern arm of 

San Francisco Bay (San Mateo, Santa Clara, and Alameda 

Counties); known only from near sea level (Grinnell, 1933; 

Gillihan and Foresman, 2004).

Remarks: Specimens originally identified as S. vagrans halicoetes from San Gregorio, along the Pacific shore of San 

Mateo County (Jackson, 1928; Grinnell, 1933), were subsequently referred to the newer subspecies S. v. paludivagus

(von Bloeker, 1939).

Sorex vagrans paludivagus von Bloeker, 1939

1939. Sorex vagrans paludivagus von Bloeker, Proc. Biol. Soc. Wash. 52:93; 

5 June.

Holotype: LACM 5053.

Type Locality: USA: “the salt marsh at mouth of Elkhorn 

Slough, Moss Landing, Monterey County, California.” 

Distribution: USA: Pacific coastal salt marshes of west central 

California, potentially from Rockaway Beach, San Mateo 

County, south at least to Seaside Lagoon, Monterey County; 

known only from near sea level (von Bloeker, 1939; Gillihan 

and Foresman, 2004).

Sorex vagrans sinuosus Grinnell, 1913

1913. Sorex sinuosus Grinnell, Univ. Calif. Publ. Zool. 10(9):187; 20 March.

1981. Sorex o[rnatus]. sinuosus, Junge and Hoffmann, Occ. Pap. Mus. Nat.

Hist. Univ. Kansas 94:34; 5 November.

Holotype: MVZ 16470.

Type Locality: USA: “Grizzly Island, near Suisun, Solano 

County, California.” 

Distribution: USA: salt marshes of Grizzly Island; known only 

from near sea level.

Remarks: Previously considered a subspecies of Sorex ornatus, Maldonado et al. (2001, 2004) demonstrated that this 

taxon is more closely related genetically to Sorex vagrans 

vagrans, with which it forms a clade that is the sister group 

to S. ornatus. 

Sorex vagrans vagrans Baird, 1857

FIGURES 23, 25

1857. Sorex vagrans Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):15, pl. 26.

1857. Sorex suckleyi Baird, Rep. Expl. Surv. Railr. Pacif. 8(1):18, pl. 27. 

Lectotype: USNM 1677, designated by Lyon and Osgood (1909:249) 

from “Steilacoom, W[estern]. T[erritories].,” Pierce County, Washington, USA. 

1891. Sorex dobsoni Merriam, N. Am. Fauna 5:33; 30 July. Holotype: 

USNM 24274/31678, from “Saw Tooth or Alturas Lake, Saw Tooth 

Mountains, Idaho . . . Altitude about 2,200 meters (7,200 feet)”; Blaine 

County, USA.

1895. Sorex vagrans dobsoni: Merriam, N. Am. Fauna 10:68; 31 December.

1895. Sorex amoenus Merriam, N. Am. Fauna 10:69; 31 December. 

Holotype: USNM 29784/41863, from “Mammoth Pass, head of 

Owens River, east slope Sierra Nevada, California (altitude, about 
NUMBER 650 • 7 5

10,000 feet),” USA. On the basis of the field catalog of the collector (E. W. Nelson), Howell (1923:266) corrected the type locality 

to “somewhere in the immediate vicinity of Mammoth (8000 feet), 

Mono County, California.”

1895. Sorex nevadensis Merriam, N. Am. Fauna 10:71; 31 December. Holotype: USNM 24891/32302, from “Reese River, Nevada,” USA. Miller 

and Kellogg (1955:20) further restricted the type locality to “about 

6,000 feet, at line between Lander and Nye Counties.”

1899. Sorex shastensis Merriam, N. Am. Fauna 16:87; 28 October. Holotype: USNM 95450, from “Wagon Camp, Mount Shasta (alt. 5,700 

ft. in the lower part of the Canadian zone)”; Siskiyou County, California, USA. 

1912. Sorex vagrans vagrans: Miller, Bull. U.S. Natl. Mus. 79:14; 31 December. First use of current name combination.

1912. Sorex californicus californicus: Miller, Bull. U.S. Natl. Mus. 79:18; 

31 December. Part; not S. ornatus californicus Merriam, 1895.

1922. Sorex trigonirostris Jackson, J. Wash. Acad. Sci. 12:264; 4 June. Holotype: USNM 203608, from “Ashland, altitude 1,975 feet, Jackson 

County, Oregon,” USA.

1922. Sorex ornatus californicus: Jackson, J. Wash. Acad. Sci. 12:264; 

4 June. Part; not Sorex ornatus californicus Merriam, 1895.

1928. Sorex vagrans nevadensis: Jackson, N. Am. Fauna 28:107; 

31 December.

1928. Sorex vagrans amoenus: Jackson, N. Am. Fauna 28:109; 31 December.

1936. Sorex ornatus trigonirostris: Bailey, N. Am. Fauna 55:366; 29 August.

Lectotype: USNM 1675 (Figure 25), designated by Lyon and 

Osgood (1909:251).

Type Locality: USA: “Shoalwater Bay, W[estern]. T[erritories].” 

[Pacific County, Washington]. 

Distribution: Canada and USA: southern British Columbia 

south through Washington and Oregon to San Francisco 

Bay in western California and through the Sierra Nevada in 

eastern California; western Montana to western Wyoming, 

northern Utah, and central Nevada; elevational distribution: 55–3,050 m (Findley, 1955; Hennings and Hoffmann, 

1977; Gillihan and Foresman, 2004).

Sorex vagrans vancouverensis Merriam, 1895

1895. Sorex vancouverensis Merriam, N. Am. Fauna 10:70; 31 December. 

1928. Sorex vagrans vancouverensis: Jackson, N. Am. Fauna 28:106; 31 December. First use of current name combination.

Holotype: USNM 71913.

Type Locality: Canada: “Goldstream, Vancouver Island, British Columbia.” 

Distribution: Canada: Vancouver, Bowen, Saltspring, Saturna, 

and South Pender Islands, southwestern British Columbia 

(Findley, 1955; Hall, 1981).

Sorex ventralis Merriam, 1895

1895. Sorex obscurus ventralis Merriam, N. Am. Fauna 10:75; 31 December.

1903. Sorex ventralis: Elliot, Field Col. Mus. Publ. 71, Zool. Ser. 3 (8):148; 

20 March. First use of current name combination.

1955. Sorex oreopolus ventralis: Findley, Univ. Kansas Publ. Mus. Nat. Hist. 

7(14):617; 10 June.

Holotype: USNM 68342.

Type Locality: Mexico: “Cerro San Felipe, Oaxaca, Mexico 

(altitude, 10,000 feet).”

Distribution: Mexico: highlands of México, Distrito Federal, 

Tlaxcala, Puebla, and Oaxaca; elevational distribution: 

1,990–3,900 m (Carraway, 2007).

Remarks: Subspecies Otisorex; S. oreopolus group. Sorex ventralis forms part of Esteva et al.’s (2010) clade A3, within 

which it is sister to S. oreopolus.

Sorex veraecrucis Jackson, 1925

1925. Sorex saussurei veraecrucis Jackson, Proc. Biol. Soc. Wash. 38:128; 

13 November.

2007. Sorex veraecrucis veraecrucis: Carraway, Monogr. West. N. Am. Nat. 

3:57; 22 June.

2012. S[orex]. salvini veraecrucis: Woodman et al., Ann. Carnegie Mus. 

80(3):214; 15 July.

Holotype: USNM 55106.

Type Locality: Mexico: “Xico, altitude 6,000 feet, state of Vera 

Cruz.” 

Distribution: Mexico: Veracruz, Puebla, and Oaxaca; elevational distribution: 1,300–2,860 m (Carraway, 2007).

Remarks: Subspecies Otiosorex; S. veraecrucis group. Sorex 

veraecrucis (as Sorex veraecrucis locality 11) is sister to 

S. ixtlanensis (Esteva et al., 2010). Together, they form Esteva et al.’s (2010) subclade S1 within clade A3.

Sorex veraepacis Alston, 1877

FIGURE 22

1843. Corsira temlyas J. E. Gray, Proc. Zool. Soc. London, 1843(11):79; 

10 January. Nomen nudum, “from Coban, in Central America.”

1877. Sorex verae-pacis Alston, Proc. Zool. Soc. London, 1877(3):445; 

1 October.

1877. C[orsira]. teculyas Alston, Proc. Zool. Soc. London, 1877(3):445; 

1 October. Nomen nudum.

1925. Sorex veraepacis veraepacis: Jackson, Proc. Biol. Soc. Wash. 38:130; 

13 November.

Lectotype: BM 43.6.13.8

Type Locality: Guatemala: “Coban (Vera Paz), Guatemala”; 

Alta Verapaz Department. Matson and Ordóñez-Garza 

(2017) noted that all specimens of Sorex veraepacis (sensu 

stricto) collected after the species was first described are 

from the Sierra de los Cuchumatanes, Guatemala. On the 

basis of this fact, they suggested that the holotype was most 

likely also collected there and transported to Cobán, which 

was an early center for trade in natural history specimens.

Distribution: Guatemala: Sierra de los Cuchumatanes of Alta 

Verapaz and Huehuetenango Departments (Matson and 

McCarthy, 2005). 
7 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Remarks: Unnamed subgenus; S. veraepacis group. Esteva et al.’s 

(2010) clade B2 consists of Sorex veraepacis (as S. veraepacis

locality 16), S. macrodon, and S. mutabilis (as S. veraepacis locality 10). Sorex chiapensis, S. ibarrai, and S. mutabilis

were formerly included in S. veraepacis as a subspecies.

Acknowledgments

I am sincerely grateful to my friend and colleague Alfred L. 

Gardner for suggesting this project several years ago and then 

gently nagging me into actually doing it. He also closely read several previous versions of this work and graciously provided extensive thoughts, insights, and suggestions that greatly improved 

the manuscript. I am indebted to Robert M. Timm and an anonymous reviewer, who read the entire manuscript and provided 

relevant comments, corrections, and additions. Any remaining 

problems are entirely my own fault. 

The following curators, collections managers, and data wranglers provided valuable information about specimens, notes, catalogs, and data under their care and occasionally chased wild geese 

for me: Ted Daeschler (ANSP); Roberto Portela Miguez (BM); 

Rhian Rowson (Bristol Museum and Art Gallery, Bristol, UK); 

Maureen Flannery (California Academy of Sciences, San Francisco); Adam Ferguson and Larry Heaney (FMNH); Jim Dines 

(LACM); Cécile Callou and Violaine Nicolas (MNHN); Chris J. 

Conroy (MVZ); Joseph Bopp (New York State Museum, Albany); Andrew J. Hebda (Nova Scotia Museum, Halifax); Craig 

Ludwig, Ingrid Rochon, Suzanne Peurach, and Darrin Lunde 

(USNM); Eric J. Sargis and Krystof Zyskowski (Yale Peabody 

Museum, New Haven, Connecticut); and Anneke van Heteren 

(Zoologische Staatssammlung München, Munich, Germany). Gil 

Taylor (Smithsonian Institution Libraries, Washington, D.C.) and 

the staff of the Raymond H. Fogler Library, University of Maine, 

Orono, arranged for an interlibrary loan of a rare microfilm 

copy of the Eastern Star and Washington Advertiser newspaper. 

Library assistance was also provided by Joan Stahl, University 

Libraries, Catholic University of America, Washington, D.C. and, 

as always, Leslie K. Overstreet and Alexandra K. Newman, Joseph F. Cullman 3rd Natural History Library, Smithsonian Institution, Washington, D.C. Support for this work came from the 

Biological Survey Unit (BSU), a field station of the U.S. Geological 

Survey’s Patuxent Wildlife Research Center that is on the verge 

of being disbanded by its parent agency. This may be one of the 

last publications to issue from this historical unit, which has been 

working collaboratively with the staff of the Smithsonian Institution and other government agencies to provide competent systematic collection services to the biological community and publish 

solid scientific research since 1885. Any use of trade, product, or 

firm names is for descriptive purposes only and does not imply 

endorsement by the U.S. government.
Relationships among species within the Nesophontidae, Solenodontidae, and Notiosoricini remain unresolved, as do relationships among the tribes of the Talpidae. Relationships of the three species of Scapanus follow He et al. (2016). 

Within the Soricidae, relationships within the Blarinini are based on Brant and Ortí 

(2002), Reilly et al. (2005), Guevara and Cervantes (2014), He et al. (2015), and Baird et 

al. (2017); those within the Soricini follow George (1988), Demboski and Cook (2001, 

2003), Maldonado et al. (2001, 2004), Shafer and Stewart (2007), Esteva et al. (2010), 

and Hope et al. (2012, 2014).

Order Eulipotyphla

Family Nesophontidae 

 Genus Nesophontes

 N. edithae

 N. hypomicrus

 N. major

 N. micrus

 N. paramicrus

 N. zamicrus

Family Solenodontidae 

 Genus Solenodon

 S. arrendondoi

 S. cubanus

 S. marcanoi

 S. paradoxus

Family Talpidae 

 Tribe Condylurini 

 Genus Condylura

 C. cristata

 Tribe Neurotrichini 

 Genus Neurotrichus

 N. gibbsii

 Tribe Scalopini 

 Genus Parascalops

 P. breweri

 Genus Scalopus

 S. aquaticus

Appendix A: Phylogenetic Listing of Species 

of New World Eulipotyphla
7 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

 Genus Scapanus

 S. anthonyi

 S. latimanus

 S. orarius

 S. townsendii

Family Soricidae 

 Subfamily Soricinae

 Tribe Blarinini 

 Genus Blarina

 B. brevicauda

 B. hylophaga

 B. carolinensis

 B. peninsulae

 B. shermani

 Genus Cryptotis

 C. mexicanus group 

 C. nelsoni

 C. magnus

 C. mexicanus

 C. obscurus

 C. phillipsii

 C. nigrescens group 

 C. brachyonyx

 C. colombianus

 C. hondurensis

 C. merus

 C. nigrescens

 C. lacandonensis

 C. mayensis

 C. merriami

 C. parvus group 

 C. berlandieri

 C. orophilus

 C. parvus

 C. pueblensis

 C. soricinus

 C. tropicalis

 C. goldmani group 

 C. alticola

 C. goldmani

 C. peregrinus

 C. goodwini group 

 C. gracilis

 C. cavatorculus

 C. celaque

 C. griseoventris

 C. mccarthyi

 C. lacertosus

 C. magnimanus

 C. mam

 C. goodwini

 C. oreoryctes

 C. thomasi group 

 C. aroensis

 C. dinirensis

 C. endersi

 C. equatoris

 C. evaristoi

 C. medellinius

 C. meridensis

 C. monteverdensis

 C. montivagus

 C. niausa

 C. osgoodi

 C. perijensis

 C. peruviensis

 C. squamipes

 C. tamensis

 C. thomasi

 C. venezuelensis

 Tribe Notiosoricini 

 Genus Megasorex

 M. gigas

 Genus Notiosorex

 N. cockrumi

 N. crawfordi

 N. evotis

 N. villai

 Tribe Soricini 

 Genus Sorex

 Subgenus Sorex

 S. araneus group 

 S. arcticus

 S. maritimensis

 S. tundrensis

 S. minutissimus group

 S. minutissimus

 Unnamed subgenus 

 Clade B1

 S. trowbridgii group

 S. trowbridgii

 S. merriami group 

 S. arizonae

 S. merriami

 S. saussurei group 

 S. altoensis

 S. mediopua

 S. saussurei

 S. salvini group 

 S. mccarthyi

 S. salvini

 S. sclateri

 S. stizodon

 Clade B2

 S. veraepacis group 

 S. chiapensis

 S. ibarrai

 S. macrodon
NUMBER 650 • 7 9

 S. madrensis

 S. mutabilis

 S. veraepacis

 Subgenus Otisorex

 S. dispar group 

 S. dispar

 Clade A3 

 S. oreopolus group 

 S. fumeus

 S. tenellus

 S. nanus

 S. oreopolus

 S. orizabae

 S. ventralis

 S. veraecrucis group (subclade S1)

 S. ixtlanensis

 S. veraecrucis

 Clade A2 

 S. cinereus group, basal lineages

 S. rohweri

 S. longirostris

 S. cinereus group, southern clade

 S. cinereus

 S. fontinalis

 S. lyelli

 S. emarginatus

 S. milleri

 S. cinereus group, Beringian clade

 S. preblei

 S. haydeni

 S. pribilofensis

 S. ugyunak

 S. portenkoi [Siberian]

 S. jacksoni

 S. camtschatica [Siberian]

 S. leucogaster [Siberian]

 Clade A1 

 S. hoyi group 

 S. hoyi

 S. vagrans group 

 S. ornatus

 S. vagrans

 S. palustris group 

 S. albibarbis

 S. palustris

 S. bendirii

 S. navigator

 S. sonomae group 

 S. pacificus

 S. sonomae

 S. monticola group 

 S. monticola

 S. obscurus

Appendix B: Literature Abbreviations

List of abbreviations used in synonymies for titles of journals, books, and other printed media. 

Abhand. Physik. Klasse Abhandlungen der Physikalischen Klasse der KöniglichPreussischen Akademie der Wissenschaften aus den Jahren 

1804–1811 (Illiger, 1815) 

Acct. Exped. Pittsb. Rocky Mtns. Account of an Expedition from Pittsburgh to the Rocky 

Mountains (James, 1822)

Am. J. Sci. American Journal of Science and Arts

Am. Midl. Nat. American Midland Naturalist

Am. Month. Mag. American Monthly Magazine and Critical Review

Am. Mus. Novit. American Museum Novitates

Am. Nat. American Naturalist

Animal Kingdom The Animal Kingdom (Kerr, 1792).

Ann. Carnegie Mus. Annals of the Carnegie Museum

Ann. Franç. Anat. Phys. Annales Françaises et Étrangères d’Anatomie et de 

Physiologie, Appliquées a la Mèdecine et a l’Histoire 

Naturelle

Ann. Mag. Nat. Hist. Annals and Magazine of Natural History

Ann. Rep. Provancher Soc. Nat. Hist. Annual Report of the Provancher Society of Natural History 

of Canada (Rapport Annuel de la Société Provancher 

d’Histoire Naturelle du Canada)

Arch. Sci. Phys. Nat. Archives des Sciences Physiques et Naturelles (Geneve)

Ark. Zool. Arkiv för Zoologi

Atlantic J. Atlantic Journal and Friend of Knowledge

Boston J. Nat. Hist. Boston Journal of Natural History

Boston J. Philos. Arts Boston Journal of Philosophy and the Arts

Bull. Am. Mus. Nat. Hist. Bulletin of the American Museum of Natural History

Bull. Geol. Nat. Hist. Surv. Minn. Bulletin of the Geological and Natural History Survey of 

Minnesota

Bull. Mus. Comp. Zool. Bulletin of the Museum of Comparative Anatomy

Bull. S. Cal. Acad. Sci. Bulletin of the Southern California Academy of Sciences

Bull. U.S. Geol. Geog. Surv. Terr. Bulletin of the United States Geological and Geographical 

Survey of the Territories

Bull. U.S. Natl. Mus. Bulletin of the United States National Museum
8 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Byull. Moscov. Ovshch. Ispyt. Pryr. Otd. Biol. Byulleten Moscovskogo Ovshchestva Ispytateley Pryrodi 

Otdel Biologycheskyy [Бюллетень Московского Общества 

Испытателей Природы, Отдел Биологический; Bulletin of 

the Moscow Society of Naturalists, Biological Series]

Cambr. Nat. Hist. The Cambridge Natural History, Vol. 10: Mammalia

Can. Field-Nat. Canadian Field-Naturalist

Cat. Mamm. Asia. Soc. Mus. Catalogue of the Mammals in the Asiatic Society Museum

Cat. Mamm. Mus. Natl. Hist. Nat. Catalogue des mammifères du Museum National d’Histoire 

Naturelle (Geoffroy Saint-Hilaire, 1803)

Cat. Mamm. Suppl. Catalogus Mammalium tam Viventium quam Fossilium.

Quinquennale Supplementum (Trouessart, 1904)

Cat. Marsup. Monotrem. Catalogue of the Marsupialia and Monotremata in the 

Collection of the British Museum (Natural History)

Checkl. Mamm. Suppl. A Check-List of Mammals of the North American Continent: 

Supplement (Elliot, 1917)

Contr. Mamm. N. Mtn. Reg. A Contribution to the Mammalogy of the North Mountain 

Region (Green, 1930)

Contr. Mus. Paleo. Univ. Mich. Contributions from the Museum of Paleontology, University 

of Michigan

Cyclop. Anat. Phys. Cyclopaedia of Anatomy and Physiology

Das Thierreich Das Thierreich eingetheilt nach dem Bau der Thiere als 

Grundlage ihrer Naturgeschichte und der vergleichenden 

Anatomie von den Herrn Ritter von Cuvier. Erster Band,

Säugethiere und Vögel (Schinz, 1821)

Dict. Class. d’Hist. Nat. Dictionnaire classique d’histoire naturelle (Geoffroy SaintHilaire, 1827a)

Die Säugthiere Suppl. Die Säugthiere in Abbildungen nach der Natur mit 

Beschreibungen von Dr. Johann Christian Danierl von 

Schreber, Supplementband (Wagner, 1855)

Diver. Distr. Diversity and Distributions

East. Star Eastern Star and Washington Advertiser (a weekly newspaper 

from Machias, Maine)

Essai Hist. Nat. Taupe Essai sur l’histoire naturelle de la taupe (De la Faille, 1769)

Fauna Am. Fauna Americana (Harlan, 1825)

Fauna Bor.-Am. Fauna Boreali-Americana (Richardson, 1829)

Field Col. Mus. Publ., Zool. Ser. Field Columbian Museum Publication, Zoological Series

Field. Zool., n.s. Fieldiana: Zoology, new series

Gen. Mamm. Insect. Galeop. Genera Mammalium. Insectivora. Galeopithecia (Cabrera, 

1925)

Gen. Mamm. Monot. Marsup. Genera Mammalium. Monotrema, Marsupialia (Cabrera, 

1919)

Gen. Zool. General Zoology, or, Systematic Natural History (Shaw, 1800)

Geog. Gesch. Mensch. Vierf. Thiere Geographische Geschichte des Menschen, und der allgemein 

verbreiteten vierfüßigen Thiere (Zimmermann, 1780)

Guthrie’s Geography A New Geographical, Historical, And Commercial 

Grammar; and Present State of the Several Kingdoms of 

the World (Guthrie, 1815)

Hist. Am. Fauna History of the American Fauna (Holder et al., 1877)
NUMBER 650 • 8 3

J. Acad. Nat. Sci. Phila. Journal of the Academy of Natural Sciences of Philadelphia

J. Elisha Mitchell Sci. Soc. Journal of the Elisha Mitchell Scientific Society

J. Mamm. Journal of Mammalogy

J. Phys. Chim. Hist. Nat. Arts Journal de Physique, de Chimie, d’Histoire Naturelle et des 

Arts

J. Wash. Acad. Sci. Journal of the Washington Academy of Science

Kung. Svenska Vet. Hand. Kungliga Svenska Vetenskapsakademiens Handlingar 

List Mamm. Brit. Mus. A List of the Specimens of Mammalia in the Collection of the 

British Museum (Gray, 1843)

Mag. Zool. Anat. Comp. Pal. Magasin de Zoologie, d’Anatomie Comparée et de 

Palaeontologie

Mamm. Biol. Mammalian Biology

Mamm. Res. Mammal Research

Mammalogie Mammalogie, ou description des espèces de mamifères, Part 1 

(Desmarest, 1820)

Mast. Neotr. Mastozoología Neotropical 

Mém. Acad. Imp. Sci. St. Petersbourg, ser. 6, Sci. Math. Phys. et Nat. Mémoires de l’Académie Impériale des Sciences de St.-

Pétersbourg. 6e série, Sciences, Mathématiques, Physiques 

et Naturelles

Mem. Am. Mus. Nat. Hist. Memoirs of the American Museum of Natural History

Mém. Mus. Hist. Nat. Mémoires du Muséum d’Histoire Naturelle (Paris)

Mem. Soc. Cienc. Nat. La Salle Memoria, La Sociedad de Ciencias Naturales La Salle

Mém. Soc. Mus. d’Hist. Nat. Strasbourg Mémoires de la Société du Muséum d’Histoire Naturelle de 

Strasbourg 

Misc. Publ. Mus. Zool., Univ. Mich. Miscellaneous Publications. Museum of Zoology, University 

of Michigan

Monatsb. K. Preuss. Akad. Wiss. Berlin Monatsberichte der Königlichen Preussische Akademie des 

Wissenschaften zu Berlin

Monogr. Insectiv. A Monograph of the Insectivora (Dobson, 1890)

Monogr. West. N. Am. Nat. Monographs of the Western North American Naturalist

MSW2 Mammal Species of the World, 2nd ed., ed. D. E. Wilson and 

D. M. Reeder, Washington, D.C.: Smithsonian Institution 

Press, 1993

MSW3 Mammal Species of the World, 3rd ed. (Wilson and Reeder, 

2005) 

Mus. Nat. Hist. The Museum of Natural History (Richardson et al., 1877)

N. Am. Fauna North American Fauna

Nasek. Mlekop. Sibiri Nasekomoyadnye Mlekopitayushchie Sibiri [Насекомоядные 

млекопитающие Сибири; Insectivorous Mammals of 

Siberia] (Yudin, 1989)

Natl. Mus. Canada Bull. National Museum of Canada Bulletin

Natürl. Syst. Natürliches System der Amphibien, mit vorangehender 

Classification der Säugthiere und Vögel. Ein beitrag zur 

vergleichenden Zoologie (Wagler, 1830) 

Northwest. Nat. Northwestern Naturalist

Nouv. tableau mamm. Nouveau tableau du règne animal. Première classe.

Mammifères (Lesson, 1842)

Occ. Pap. Mus. Nat. Hist. Univ. Kansas Occasional Papers of the Museum of Natural History,

University of Kansas
8 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Occ. Pap. Mus. Texas Tech Univ. Occasional Papers of the Museum of Texas Tech University

Occ. Pap. Mus. Zool. Louisiana State Univ. Occasional Papers of the Museum of Zoology, Louisiana 

State University

Occ. Pap. Mus. Zool. Univ. Mich. Occasional Papers of the Museum of Zoology, University of 

Michigan

Précis découv. Précis des décourvertes et travaux somiologiques ou 

zoologiques et botaniques (Rafinesque, 1814)

Proc. Acad. Nat. Sci. Phila. Proceedings of the Academy of Natural Sciences of 

Philadelphia

Proc. Biol. Soc. Wash. Proceedings of the Biological Society of Washington

Proc. Boston Soc. Nat. Hist. Proceedings of the Boston Society of Natural History

Proc. Calif. Acad. Sci. Proceedings of the California Academy of Sciences

Proc. Indiana Acad. Sci. Proceedings of the Indiana Academy of Science

Proc. New Engl. Zool. Club Proceedings of the New England Zoological Club

Proc. Penna. Acad. Sci. Proceedings of the Pennsylvania Academy of Science

Proc. Trans. Nova Scot. Inst. Nat. Sci. Proceedings and Transactions of the Nova Scotian Institute 

of Natural Science

Proc. U.S. Natl. Mus. Proceedings of the United States National Museum

Proc. Wash. Acad. Sci. Proceedings of the Washington Academy of Sciences

Proc. Zool. Soc. Lond. Proceedings of the Zoological Society of London

Prodr. Syst. Mamm. Avium Prodromus Systematis Mammalium et Avium (Illiger, 1811)

Recherches des Mamm. Recherches pour servir a l’histoire naturelle des mammifères 

Regnum animale Regnum animale in classes IX (Brisson, 1756, 1762)

Rep. Expl. Surv. Railr. Pacif. Reports of Explorations and Surveys, to Ascertain the Most 

Practicable and Economical Route for a Railroad from the 

Mississippi River to the Pacific Ocean. Volume 8: General 

Report upon the Zoology of the Several Pacific Railroad 

Routes. Part 1: Mammals. Washington, D.C.: A. O. P. 

Nicholson

Rep. W. Va. Board Agric. Report of the West Virginia State Board of Agriculture for 

the Quarter Ending December 30, 1910. Forestry

Rev. Franc. Mamm. Revue Française de Mammalogie

Rev. Mus. Argentino Revista del Museo Argentino de Ciéncias Nautales 

“Bernardino Rivadavia” e Instituto Nacional de 

Investigación de las Ciéncias Naturales

Rev. Peruana Biol. Revista Peruana de Biología

Säug. Mitteil. Säugetierkundliche Mitteilungen

Sci. Publ. Cleveland Mus. Nat. Hist. Scientific Publications of the Cleveland Museum of Natural 

History

Sitzungsber. Math.-natürwissensch. Classe Kaiserl. Akad. Wissensch. Sitzungsberichte der Mathematisch-natürwissenschaftlichen 

Classe der Kaiserlichen Akademie der Wissenschaften

Smithson. Misc. Coll. Smithsonian Miscellaneous Collections

Southwest. Nat. Southwestern Naturalist

Spec. Mamm. Brit. Mus. A List of the Specimens of Mammalia in the Collection of the 

British Museum

Spec. Publ. Carnegie Mus. Special Publication of the Carnegie Museum of Natural 

History

Spec. Publ. Mus. Texas Tech Univ. Special Publication of the Museum, Texas Tech University

Spec. Zool. Geog. Specimen Zoologiae Geographicae, Quadrupedum Dimicilia 

et Migrationes Sistens (Zimmermann, 1777)

Synop. Insectiv. Mamm. Synopsis of Insectivorous Mammals (Gill, 1875)
NUMBER 650 • 8 5

Syst. Biol. Systematic Biology

Systema Naturae, 10th ed. Systema Naturae, 10th edition of Linnaeus (1758)

Systema Naturae, 13th ed. Systema Naturae, 13th edition of Gmelin (1788)

Syst. Reg. Anim. Systema Regni Animalis per Classes, Ordines, Genera,

Species, Varietates: cvm Synonymia et Historia 

Animalivm: Classis I. Mammalia (Erxleben, 1777)

Syst. Zool. Systematic Zoology

Thesaurus Locupletissimi Rerum Naturalium Thesauri Accurata 

Descriptio (Seba, 1734)

Thierreich Das Thierreich eingetheilt nach dem Bau der Thiere als 

Grundlage ihrer Naturgeschichte und der vergleichenden 

Anatomie

Trans. Kansas Acad. Sci. Transactions of the Kansas Academy of Science

Trans. Linn. Soc. New York Transactions of the Linnean Society of New York

Univ. Calif. Publ. Zool. University of California Publications in Zoology

Univ. Kansas Mus. Nat. Hist., Misc. Publ. University of Kansas, Museum of Natural History,

Miscellaneous Publications

Univ. Kansas Publ. Mus. Nat. Hist. University of Kansas Publications, Museum of Natural 

History 

Univ. Kansas Sci. Bull. University of Kansas Science Bulletin

U.S. Natl. Mus. Circ. United States National Museum, Circular (True, 1884)

Virginia Mus. Nat. Hist. Mem. Virginia Museum of Natural History Memoir

Voll. Natursyst. Suppl. Vollständigen Natursystems Supplements und Register Band

(Müller, 1776, 1789)

Zeit. Geol. Wiss. Zeitschrift für Geologische Wissenschaften

Zoognosia Zoognosia Tabulis Synopticis Illustrata (Fischer, 1814)

Zool. J. Zoological Journal

Zool. J. Linn. Soc. Zoological Journal of the Linnean Society

Zool. New York Zoology of New-York, or the New-York Fauna. Part 1:

Mammalia (De Kay, 1842)

Zool. Record Zoological Record

Zool. Zh. Zoologicheskii Zhurnal [Зоологический Журнал; Zoological 

Journal]

Abbreviations used in the text for institutions with systematic collections in which 

primary types (holotypes, lectotypes, neotypes) of Eulipotyphlans were deposited. Numbers in brackets are the numbers of primary types of Eulipotyphla 

currently known to be held by the institution. 

AMNH Department of Mammalogy, Division of Vertebrate Zoology, American Museum of Natural History, New York, New York, USA (17)

AMNH VP Department of Vertebrate Paleontology, Division of Paleontology, 

American Museum of Natural History, New York, New York, USA (2)

ANSP Mammalogy Collection, Academy of Natural Sciences of Philadelphia (now Academy of Natural Sciences of Drexel University), 

Philadelphia, Pennsylvania, USA (7)

ANSPVP Paleontology Collection, Academy of Natural Sciences of Philadelphia 

(now Academy of Natural Sciences of Drexel University), Philadelphia, Pennsylvania, USA (1)

BM The Natural History Museum [formerly the British Museum (Natural History)], London, UK (11)

CM Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, 

USA (5)

CMN Canadian Museum of Nature, Ottawa, Canada (5)

CMNH Cleveland Museum of Natural History, Cleveland, Ohio, USA (4)

CMVP Division of Vertebrate Paleontology, Carnegie Museum of Natural 

History, Pittsburgh, Pennsylvania, USA (1)

CNMA Colección de Mamíferos, Instituto de Biología, Universidad Nacional 

Autónoma de México, Mexico City, Mexico (1)

CUMV Cornell University Museum of Vertebrates, Ithaca, New York, USA (1)

CVULA Colección de Vertebrados de la Universidad de Los Andes, Mérida, 

Venezuela (1)

EBRG Museo Estación Biológica de Rancho Grande, El Limón, Carabobo, 

Venezuela (2)

FLMNH Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA (1)

FMNH Field Museum of Natural History, Chicago, Illinois, USA (7)

GEC Grupo de Exploraciones Científicas, Havana, Cuba (2)

KU Collection of Mammals, Natural History Museum and Biodiversity 

Institute, University of Kansas, Lawrence, Kansas, USA (11)

Appendix C: Institutional Abbreviations 

for Systematic Collections
8 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

KUVP Collection of Vertebrate Paleontology, 

Natural History Museum and Biodiversity 

Institute, University of Kansas, Lawrence, 

Kansas, USA (1)

LACM Natural History Museum of Los Angeles 

County, Los Angeles, California, USA (2)

LSU Louisiana State University Museum of Zoology, Baton Rouge, Louisiana, USA (1)

MB Museum für Naturkunde an der HumboldtUniversität zu Berlin, Paläontologisches 

Museum, Berlin, Germany [Ma. 1967 = 

Mammalia, 1967 gesammelt] (1)

MBLUZ Museo de Biología, Universidad del Zulia, 

Maracaibo, Venezuela (1)

MCZ Museum of Comparative Zoology, Harvard 

University, Cambridge, Massachusetts, 

USA (6)

MCZ BANGS Collection of E. A. and Outram Bangs, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (8)

MEPN Museo de Historia Natural “Gustavo 

Orces V.,” Escuela Politécnica Nacional, 

Quito, Ecuador (1)

MNHN Museum National d’Histoire Naturelle, 

Paris, France (0)

MNHNC Museo Nacional de Historia Natural de 

Cuba, Havana, Cuba (1)

MSB Museum of Southwestern Biology, University 

of New Mexico, Albuquerque, New Mexico, 

USA (1)

MUSA Museo de Historia Natural de la Universidad 

Nacional de San Agustín, Arequipa, Peru (1)

MUSM Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Lima, 

Peru (1)

MVZ Museum of Vertebrate Zoology, University of 

California, Berkeley, California, USA (24)

MZFC Museo de Zoología “Alfonso L. Herrera,” 

Universidad Nacional Autónoma de México, 

Mexico City, Mexico (1)

MZSL Museum of the Zoological Society of London, whose collections were transferred to 

the British Museum (Natural History), now 

the Natural History Museum, London, UK

OMNH Sam Noble Oklahoma Museum of Natural 

History, University of Oklahoma, Norman, 

Oklahoma, USA (1)

PM Philadelphia Museum, Philadelphia, Pennsylvania, USA, an 18th and early 19th century 

museum; nearly all of the collection has been 

lost or destroyed (Woodman, 2009)

PSMNH Puget Sound Puget Sound Museum of 

Natural History, University of Puget Sound, 

Tacoma, Washington, USA (1)

RAS Zoological Museum, Zoological Institute, 

Russian Academy of Sciences, Saint Petersburg, Russia (1) 

RBCM Royal British Columbia Museum, Victoria, 

British Columbia, Canada (2)

SMNH Sternberg Museum of Natural History, Fort 

Hays State University, Hays, Kansas, USA (1)

TCWC Texas Wildlife Cooperative Collection, Texas 

A&M University, College Station, Texas, 

USA (3)

TTU Museum of Texas Tech University, Lubbock, 

Texas, USA (1)

UAM University of Alaska Museum of the North, 

Fairbanks, Alaska, USA (1)

UF Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA (1)

UIMNH Museum of Natural History, University of 

Illinois, Urbana, Illinois, USA [defunct; collections transferred to MSB]

UMMZ University of Michigan Museum of Zoology, 

Ann Arbor, Michigan, USA (5)

USNM Division of Mammals, Department of Vertebrate Zoology, National Museum of Natural 

History, Smithsonian Institution, Washington, D.C., USA (formerly the United States 

National Museum) (147)

USNM PAL Department of Paleobiology, National 

Museum of Natural History, Smithsonian 

Institution, Washington, D.C., USA (formerly 

the United States National Museum) (3)

UWBM University of Washington Burke Museum, 

Seattle, Washington, USA (1)

UWYMV University of Wyoming Museum of Vertebrates, Laramie, Wyoming, USA (1)

ZMB Humboldt-Universität zu Berlin, Museum für 

Naturkunde (formerly Zoologisches Museum 

Berlin), Berlin, Germany (2)
There are approximately 327 known type localities for New World eulipotyphlans distributed among 17 countries. 

Canada (30)

No locality (2)

British Columbia (8)

Manitoba (3)

Newfoundland and Labrador (2)

Northwest Territories (2)

Nova Scotia (7)

Ontario (3)

Quebec (3)

Canada/Alaska (2)

Colombia (7)

Antioquia (2)

Cauca (1)

Cesar (1)

Cundinamarca (1)

Distrito Capital (2)

Costa Rica (6)

Cartago (3)

Limón (1)

Puntarenas (1)

San José (1)

Cuba (7)

Dominican Republic (2)

Ecuador (4)

Azuay (1)

Bolívar (1)

Napo (1)

Pichincha (1)

Appendix D: Distribution of Type Localities
9 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Guatemala (12)

Alta Verapaz (4)

El Progreso (1)

Huehuetenango (3)

Quetzaltenango (3)

Totonicapán (1)

Haiti (3)

Hispaniola (Haiti/Dominican Republic) (1)

Honduras (6)

Comayagua (1)

Cortés (1)

Francisco Morazán (1)

Lempira (2)

Santa Barbara (1)

Mexico (51)

Baja California (4)

Chiapas (6)

Coahuila (2)

Distrito Federal (1)

Durango (1)

Guanajuato (1)

Guerrero (2)

Hidalgo (1)

Jalisco (7)

México (1)

Nayarit (1)

Oaxaca (12)

Puebla (2)

Sinaloa (1)

Tamaulipas (4)

Veracruz (4)

Yucatan (1)

Nicaragua (1)

Jinotega (1)

Panama (4)

Bocas del Toro (1)

Chiriquí (2)

Darién (1)

Peru (2)

Cajamarca (2)

Puerto Rico (1)

Russia (1)

Krasnoyarskii Krai (1)

USA (182)

Unknown locality (5)

Alabama (1)

Alaska (14)

Arizona (4)

Arkansas (2)

California (35)

Colorado (1)

Connecticut (1)

Florida (9)

Idaho (4)

Indiana (2)

Kansas (3)

Kentucky (2)

Louisiana (1)

Maine (1)

Maryland (2)

Massachusetts (3)

Michigan (1)

Minnesota (1)

Mississippi (1)

Missouri/Illinois (1)

Montana (2)

Nebraska (5)

Nevada (1)

New Hampshire (1)

New Jersey (1)

New Mexico (2)

New York (10)

North Carolina (3)

North Dakota (1)

Ohio (2)

Oklahoma (2)

Oregon (14)

Oregon/Washington (1)

Pennsylvania (5)

South Carolina (3)

Tennessee (1)

Texas (5)

Utah (1)

Vermont (3)

Virginia (4)

Washington (16)

West Virginia (1)

Wisconsin (3)

Wyoming (1)

Venezuela (5)

Aragua (1)

Merida (1)

Táchira (1)

Trujillo (1)

Yaracuy (1)
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acadica, Sorex, 51

acadicus, Neosorex, 47

Sorex, 47, 51, 52, 54

aenea, Talpa, 27

aeneus, Scalops, 27

aequatoris, Blarina, 36

aereus, Scalops, 20

Scalopus, 20, 21

alascensis, Sorex, 64, 65

alaskanus, Sorex, 64, 

albibarbis, Neosorex, 47, 48

Sorex, 47, 64, 70, 79

albiventer, Atophyrax, 49

Neosorex, 49

Sorex, 49, 

alleni, Scalopus, 21

alnorum, Microsorex, 58, 59

Sorex, 58, 59

aloga, Blarina, 28

alpinus, Scapanus, 26

Sorex, 47

alticola, Blarina, 35

Cryptotis, 35, 78

altoensis, Sorex, 48, 71, 72, 78

americana, Monodelphis, 6, 

Talpa, v, 10, 19

amoenus, Sorex, 74, 75

Amphiosorex, 47

Amphisorex, vi, 47, 51, 55, 56, 70

Amphi-Sorex, 46

anastasae, Scalops, 22

Scalopus, 22

anastasiae, Scalops, 22

angusta, Blarina, 32, 

angusticeps, Blarina, 32

Anotus, 28

anthonyi, Scapanus, 25, 78

Antillogale, 12

aquatica, Talpa, 22

aquaticus, Scalops, 20, 22, 24, 25

Scalopus, v, 15, 20–25, 27, 77

Scalpos, 22

Sorex, v, 15, 16, 20, 22, 23, 27

araneus, Mus, 6

Sorex, 5, 6, 46, 47, 48, 61, 73, 78

arcticus, Sorex, v, vi, 8, 47–52, 55, 57, 61, 

70, 73, 78

argentatus, Scalops, 24, 25

arizonae, Sorex, 49, 78

aroensis, Cryptotis, 35, 78

arredondoi, Solenodon, 12

Astromycter, 14–16

Astromyctes, 15

Astromydes, 15, 16

Atophyrax, 47, 49

Atopogale, 12

australis, Scalopus, 22

avia, Cryptotis, 44

avius, Cryptotis, 44

bachmani, Crocidura, 60

Musaraneus, 60

bairdi, Sorex, 67, 68

bairdii, Sorex, 68, 69

bassi, Scalopus, 22

belli, Sorex, 48

bendirei, Atophyrax, 49

Sorex, 49

bendiri, Atophyrax, 49

Sorex, 49

bendirii, Atophyrax, 47, 49

Neosorex, 49

Sorex, 49, 64, 79

berlandieri, Blarina, v, 18, 35

Cryptotis, v, 18, 35, 36, 78

Blaria, 28, 40

Blarina, iii, v, vi, 5, 9, 18, 28–45, 78

blitchi, Sorex, 56, 57

brachyonyx, Cryptotis, 36, 78

Brachysorex, 28, 34, 42

brasiliensis, Musaraneus, 6

Sorex, 6

Taxonomic Index
102 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

brevicauda, Blarina, v, vi, 

28–34, 44, 78

brevicaudata, Didelphys, 9

Monodelphis, 9

brevicaudatus, Brachysorex, 28

Peramys, 9

Sorex, v, 28, 30

brevicaudus, Sorex, v, 28–30

breweri, Parascalops, v, 19–21, 77

Scalops, v, 15, 19–21

Scapanus, 20

Scaphanus, 20

Talpa, 20

brooksi, Sorex, 64

buchariensis, Sorex, 47

caecutiens, Sorex, 70

californicus, Scalops, 25, 26

Scapanus, 25, 26

Sorex, 66, 67, 74, 75

calvertensis, Sorex, 68

campi, Scapanus, 26

canadensis, Scalops, 22, 27

Talpa, v, 15

carolinensis, Blarina, vi, 28, 

32–34, 78

Sorex, vi, 28, 32–34

caryi, Scalopus, 22

cascadensis, Sorex, 68

caudata, Talpa, 19, 20

caudatus, Episoriculus, 63

Sorex, 63

caurinus, Scapanus, 26

cavatorculus, Cryptotis, 36, 

37, 78

celaque, Cryptotis, 36, 37, 78

celatus, Cryptotis, 43

cerulescens, Sorex, 6

chiapensis, Sorex, 49, 61, 63, 

76, 78

christata, Condylura, 16

churchi, Blarina, 32

cinereus, Sorex, v, vi, 8, 18, 33, 

35, 41, 42, 46, 47, 51–58, 

60–62, 70, 71, 74, 79 

cockrumi, Notiosorex, 45, 46, 78

Codylura, 6, 14

colombiana, Cryptotis, 36

colombianus, Cryptotis, 36, 78

compacta, Blarina, 28

concolor, Sorex, 32

Condylura, iii, v, 6, 14–17, 

19–21, 28, 77

Condylurus, 14

Condylus, 15, 16

Condytura, 14, 16

constrictus, Sorex, 7

cooperi, Crocidura, 51

Sorex, v, vi, 8, 51, 53

Corsira, 28, 32, 44, 46, 51, 75

costaricensis, Blarina, 28

crawfordi, Notiosorex, 45, 

46, 78

Sorex, 45, 46

Criptotis, 35

cristata, Condylura, v, 6, 15–17, 

20, 21, 77

Condylus, 16

Condytura, 16

Talpa, 15

Talpasorex, 15

cristatus, Astromydes, 16

Rhinaster, 16

Scalops, 15

Scalopus, 15

Sorex, v, 14–16, 21, 22

cristobalensis, Sorex, 71

Crocidura, 7, 9, 35, 47, 51, 60

crockrumi, Notiosorex, 46

Croscopus, 7

Crossopus, 7, 47, 70

Cryptotis, iii, v, vi, 1, 5, 6, 9, 

18, 28, 29, 33, 34–45, 

48, 78

Cryptotus, 35

Cryptotys, 35

cryptus, Scalopus, 24

Ctenomys, 9

cubana, Atopogale, 12

cubanus, Solenodon, v, 12, 13, 77

cumberlandensis, Blarina, 30

cupreata, Talpa, 22

daphaenodon, Sorex, 47

davisi, Scalopus, 22

dekayi, Sorex, vi, 32, 33

delmarvensis, Blarina, 30

destructionensis, Neurotrichus, 17

destructioni, Sorex, 73

dichrurus, Sorex, 7

dicrurus, Sorex, 7

dilatus, Scapanus, 26

dinirensis, Cryptotis, 36, 78

Dipus, 7

dispar, Sorex, 47, 56, 57, 79

dobsoni, Sorex, 74

durangae, Sorex, 63

edithae, Nesophontes, 11, 77

eionis, Sorex, 60

elassodon, Sorex, 68

elasson, Cryptotis, 42, 43

emarginatus, Sorex, 57, 62, 79

endersi, Cryptotis, 36, 78

equatoris, Blarina, 36

Cryptotis, 36, 40, 41, 43, 78

etruscus, Sorex, 49

Suncus, 49

europaea, Talpa, 19, 22

europea, Talpa, 19, 22

Eurosorex, 47

euryrhynchis, Cryptotis, 35

evaristoi, Cryptotis, 37, 78

evotis, Notiosorex, 46, 78

Sorex, 46

exilipes, Blarina, v, 18, 42

exilis, Sorex, 7

eximius, Blarina, 42

Microsorex, 58

Sorex, 58

fimbriata, Sorex, 7

fimbripes, Crocidura, 7

Croscopus, 7

Crossopus, 7, 70

Neosorex, 7

Sorex, v, 7, 8, 47, 48, 70

fisheri, Sorex, 60

fissipes, Codylura, 6

Condylura, 6

Talpa, 6

Talpasorex, 6

flava, Talpa, 15, 22

flavescens, Talpa, 22

floridana, Blarina, vi, 33, 41, 42

Cryptotis, 41

floridanus, Cryptotis, vi, 33, 35, 

41, 42

fodiens, Neomys, 46, 56

Sorex, 56

fonitnalis, Sorex, 57

fontinalis, Sorex, 57, 79

forsteri, Amphisorex, 51

Corsira, 51

Sorex, v, vi, 8, 51, 52 57

fossilis, Blarina, 29

fossor, Blarina, 37

Cryptotis, 37

fosteri, Sorex, 51

frankstounensis, Sorex, 52

frontalis, Cryptotis, 37

fumens, Sorex, 58

fumeus, Sorex, 8, 57, 58, 79

fusca, Talpa, 22

Galemys, 14, 28, 34, 44, 70

gaspensis, Sorex, 56, 57

gibbsi, Neurotrichus, 17

Neürotrichus, 17

gibbsii, Neurotrichus, v, 17–19, 77

Neürotrichus, 17

Nëurotrichus, 17

Urotrichus, v, 17, 18

gibsii, Urotrichus, 17

gigas, Megasorex, 45, 78

Notiosorex, 45

glacialis, Sorex, 64, 65

gloveralleni, Sorex, 47, 48

godmani, Sorex, 72

goldmani, Blarina, 37, 38

Cryptotis, 35, 37, 38, 43, 78

goodwini, Cryptotis, 35–39, 

41, 78

gracilis, Cryptotis, 37, 78

grinnelli, Scapanus, 27

griseoventris, Cryptotis, 37, 

38, 78

guerrerensis, Cryptotis, 37

haitiensis, Solenodon, 13, 14

halicoetes, Sorex, 74

harlani, Brachysorex, 28, 42

Cryptotis, 42, 43

Galemys, 28

Sorex, vi, 28, 42

haydeni, Sorex, vi, 50, 58, 70, 79

hermanni, Sorex, 46

Hesperoscalops, 20

hollisteri, Sorex, 55

Homalura, 47

Homalurus, 47

hondurensis, Cryptotis, 38, 78

hooperi, Blarina, 32

howelli, Scalopus, 24

hoyi, Microsorex, 58, 59

Sorex, v, vi, 18, 47, 50, 58, 

59, 79

hudsonius, Dipus, 7

Zapus, 7

hulophaga, Blarina, 34

humboldtensis, Sorex, 73

hyacinthinus, Neurotrichus, 17

Neürotrichus, 17

hydrobadistes, Sorex, 70

hydrodromus, Sorex, 70, 71

Hydrogale, 47

hylophaga, Blarina, 28–30, 

34, 78 

hypomicrus, Nesophontes, 11, 

12, 77

ibarrai, Sorex, 60, 63, 76, 78

idahoensis, Sorex, 51

inflatus, Scalopus, 24

insularis, Scapanus, 25

Sorex, 68

intermedius, Neurotrichus, 17

Scalops, 20

Scalopus, 20

intervectus, Microsorex, 58

isolatus, Sorex, 68

ixtlanensis, Sorex, 60, 75, 79

jacksoni, Cryptotis, 37

Sorex, 60, 70, 74, 79

jerryrchoatei, Blarina, 30

jucensis, Sorex, 66

juncensis, Sorex, 66

kirtlandi, Blarina, 32

knoxjonesi, Blarina, 30

labradorensis, Sorex, 47, 48

lacandonensis, Cryptotis, 38, 78

lacertosus, Cryptotis, 37, 38, 78

lagunae, Sorex, 66

laricorum, Sorex, vi, 49, 50

lasertosus, Cryptotis, 38

latimanus, Scalops, 25, 26

Scapanus, 25–27, 78

leseueri, Amphisorex, 55

leseurii, Amphisorex, 55

lesneurii, Sorex, 55

lesueri, Amphisorex, 55

Sorex, 55

lesuerii, Amphisorex, 55

lesueuri, Amphisorex, 55

Sorex, 55

lesueurii, Amphisorex, vi, 55, 56

Sorex, 54, 55

leucogenys, Sorex, 62
NUMBER 650 • 103

longicauda, Sorex, 68, 69

Talpa, 19

longicaudata, Condylura, 19

Talpa, 15, 19, 20 

Talpasorex, 19

longicaudatus, Rhinaster, 20

longicaudus, Sorex, 68

longiquus, Sorex, 65

longirostris, Nesophontes, 11

Otisorex, 60

Sorex, vi, 33, 55, 60, 61, 

71, 79 

lyelli, Sorex, 61, 70, 79

macer, Cryptotis, 35

machetes, Blarina, 37, 43

Cryptotis, 37, 43

machrina, Talpa, 24

machrinoides, Scalopus, 24

machrinus, Scalops, 20, 24

Scalopus, 24

macra, Cryptotis, 35

macrodon, Sorex, 61, 76, 78

macroura, Condylura, v, 15, 16, 

Rhinaster, 16

macrourus, Sorex, 56

macrurus, Episoriculus, 56

Rhinaster, 16

Sorex, 56

Soriculus, 56

madrea, Cryptotis, 41

madrensis, Sorex, 61, 79

magna, Blarina, 38

Cryptotis, 38

magnimana, Cryptotis, 38

magnimanus, Cryptotis, 37, 

38, 78

magnus, Cryptotis, 38, 78

major, Nesophontes, 11, 77

Neurotrichus, 17

malitiosus, Sorex, 69

mam, Cryptotis, 37, 38, 78

manitobensis, Blarina, 29

marcanoi, Antillogale, 12

Solenodon, 12, 77

mariposae, Sorex, 73

maritimensis, Sorex, 61, 78

mayensis, Blarina, 38

Cryptotis, 38, 39, 78

mccarthyi, Cryptotis, 37, 39, 78

Sorex, 61, 78

medellinia, Cryptotis, 39

medellinius, Cryptotis, 39, 78

medellinus, Cryptotis, 39

mediopua, Sorex, 62, 78

Megasorex, iii, 45, 78

melanogenys, Sorex, 63

melanotis, Sorex, v, 9

mera, Cryptotis, 39

meridensis, Blarina, 36, 39

Cryptotis, 39, 78

merriami, Cryptotis, 39, 78

Sorex, 47, 49, 55, 62, 73, 78

merus, Cryptotis, 39, 78 

metallescens, Scalops, 27

mexicana, Blaria, 40

Blarina, 37–39, 43

Cryptotis, 37, 40, 41, 43

Soriciscus, 39 

mexicanus, Cryptotis, 37–41, 

43, 78

Sorex, 9, 10

Microsorex, 47, 58, 59

micrura, Blarina, vi, 44, 45

Cryptotis, 38, 44

micrurus, Galemys, 28, 32, 44

Sorex, 38, 44

micrus, Nesophontes, 11, 77

milleri, Sorex, 54, 57, 62, 79

minima, Blarina, 34

minor, Neurotrichus, 19

minusculus, Scapanus, 26

minutissimus, Sorex, 47, 62, 78

mirabilis, Sorex, 47

miscix, Sorex, 55

mixtus, Sorex, 69

monoensis, Scapanus, 26

monsvairani, Soricidus, 47

montanus, Microsorex, 59

Scalopus, 24

Sorex, 59

Suncus, 59

montereyensis, Sorex, 73

monteverdensis, Cryptotis, 40, 78

monticola, Sorex, 47, 62–67, 79

monticolus, Sorex, 63, 68

montivaga, Blarina, 40

Cryptotis, 40

montivagus, Cryptotis, 36, 40, 

43, 78

Mus, 6, 11, 12

Musaraneus, 6, 34, 35, 46, 60

musculus, Mus, 11, 12

mutabilis, Sorex, 60, 63, 76, 79

myops, Sorex, 72

nanus, Scalopus, 24

Sorex, 23, 79

navigator, Neosorex, vi, 47, 

53, 64

Sorex, vi, 47, 53, 64, 70, 79

nayaritensis, Cryptotis, 35

nebrascensis, Scalopus, 24

nelsoni, Blarina, 40

Cryptotis, 40, 78

neomexicanus, Sorex, 63

Neomys, 46, 47, 56

Neosorex, vi, 7, 47–49, 53, 

64, 70

Neourotrichus, 17

Nesophontes, iii, 10–12, 77

Neurotrichus, iii, v, 14, 

17–19, 77

Neürotrichus, 17

Nëurotrichus, 17

nevadensis, Sorex, 75

niausa, Cryptotis, 40, 78

niger, Sorex, v, 31, 32

Talpa, v, 22, 23

nigra, Condylura, 16, 17

nigrescens, Blarina, 39, 40

Cryptotis, 36, 38–41, 78 

Soriciscus, 40 

nigriculus, Sorex, 55

Notiosorex, iii, 35, 43, 45, 46, 78

oaxacae, Sorex, 71

obscura, Blarina, 41

Cryptotis, 41

obscuroides, Sorex, 65

obscurus, Cryptotis, 41, 78

Sorex, 62–69, 75, 79

occultus, Scapanus, 26, 27

Ognevia, 47

ohionensis, Sorex, 52

olivaceus, Blarina, 41

olympicus, Scalops, 27

Scapanus, 27

orarius, Scapanus, 25, 27, 78

oreinus, Sorex, 66

oreopolus, Sorex, 47, 57, 63, 

65, 73, 75, 79

oreoryctes, Cryptotis, 37, 41, 78

orinus, Sorex, 66

orizabae, Sorex, 65, 79

ornatus, Sorex, 65–67, 74, 75, 79

orophila, Blarina, 41 

Cryptotis, 41

Soriciscus, 41

orophilus, Cryptotis, 41, 78

osgoodi, Blarina, 41

Cryptotis, 41, 78

Otisorex, vi, 46, 47, 49, 51, 54, 

56–58, 60–67, 69–75, 79 

Oxyrhin, 46

ozarkensis, Blarina, 32

pachyurus, Sorex, vi, 49, 50

pacificus, Sorex, 62, 64, 67–69, 

72, 79

pallida, Blarina, 32

palmeri, Atophyrax, 49

Neosorex, 49

Sorex, 49

paludivagus, Sorex, 74

palustris, Amphisorex, 70

Crossopus, 70

Galemys, 70

Neosorex, 47, 64, 70

Sorex, 7, 8, 47, 49, 64, 69, 

70, 79

paradoxus, Solenodon, v, 3, 

12–14, 77

paramicrus, Nesophontes, 11, 

12, 77

Parascalops, iii, v, 14, 19–21, 77

parva, Blarina, 35

Condylura, 17

Cryptotis, 35, 41–44

parvidens, Sorex, 66, 67

parvus, Cryptotis, v, vi, 18, 28, 

29, 33, 35, 36, 41–44, 

48, 78

Scalops, 24, 25

Scalopus, 24, 25

Scapanus, 27

Sorex, v, 18, 28, 29, 35, 42, 48

peninsulae, Blarina, 34, 78

pennantii, Talpa, 24

pensylvancius, Talpa, 22

Talpasorex, 22

pennsylvanica, Scalops, 20, 22

pensylvanica, Talpasorex, 22

Perascalops, 19

peregrina, Blarina, 43

Cryptotis, 40, 43

peregrinus, Cryptotis, 40, 43, 78

pergracilis, Blarina, 35

Cryptotis, 35, 43

perijensis, Cryptotis, 43, 78

permiliensis, Sorex, 69, 

personatus, Sorex, 8, 51, 55, 56, 

58, 60

peruviensis, Cryptotis, 40, 43, 78

phillipsii, Cryptotis, 35, 43, 78

Notiosorex, 35, 43

Xenosorex, 43

platyrhinus, Otisorex, vi, 46, 

51, 54, 57

Sorex, v, 18, 51, 57

platyrrhinchus, Sorex, 51

platyurus, Talposorex, 28

plumbea, Blarina, 34

poeyanus, Solenodon, 12

porteri, Scalopus, 25

prarinatus, Astromycter, 16

prasinata, Condylura, 14, 16

prasinatus, Astromycter, 16

preblei, Sorex, 61, 70, 79

prevostensis, Sorex, 69

pribilofensis, Sorex, 60, 70, 71, 

74, 79

pueblensis, Cryptotis, 43, 78

pulcher, Scalopus, 21

punctulatus, Sorex, 47, 48

purpurascens, Talpa, 15, 22

pyrhonota, Cryptotis, 5

pyrrhonota, Blarina, 5, 6, 9, 10

radiata, Talpa, 15

radiatus, Sorex, 15

Rattus, 11, 12

rattus, Rattus, 11, 12

relictus, Sorex, 66, 67

reposta, Talpa, 20

rexroadi, Hesperoscalops, 20

Rhinaster, 14

richardsoni, Sorex, v, 8, 48, 

51, 57

rohweri, Sorex, 71, 79

rubra, Talpa, v, 10, 

rufa, Chrysochloris, 10

Talpa, 10

salarius, Sorex, 66, 67

salicornicus, Sorex, 66, 67

salvini, Sorex, 47, 48, 61, 

71–73, 75, 78

saussurei, Sorex, 47, 48, 61–63, 

71–73, 75, 78
104 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Scalops, v, 4, 15, 19–22, 24–27

Scalopus, iii, v, 14, 15, 20–25, 

27, 28, 77

Scalpos, 20, 22

Scapanus, iii, 14, 20, 25–27, 

77, 78

Scapasius, 25

Scaphanus, 20, 25

schefferi, Scapanus, 27

sclateri, Sorex, 72, 78

sericatus, Scapanus, 26

sericea, Talpa, 24

setosus, Sorex, 69

shastensis, Sorex, 75

shermani, Blarina, 34, 78

shumaginensis, Sorex, 65, 

similis, Asoriculus, 65 

Sorex, 65

simplicidens, Blarina, 32

sinuosus, Sorex, 66, 74

Solenodon, iii, v, 3, 12–14, 77

sonomae, Sorex, 47, 67, 72, 79

soperi, Sorex, 65, 

Sorax, 47

Sore, 47

Sorex, iii, v, vi, 2, 4–10, 14–16, 

18, 20–23, 27–35, 38, 39, 

41, 42, 44–76, 78, 79

Soricidus, 47

soricina, Blarina, 44

Cryptotis, 44

soricinus, Cryptotis, 44, 78

Soriciscus, 35, 39–41

Soriculus, 56, 

Sorieiscus, 35

sphagnicola, Sorex, 48

sphagnicolus, Sorex, 48

squamipes, Blarina, 44

Cryptotis, 44, 78

stizodon, Sorex, 72, 78

streatori, Sorex, 56

Strogonovia, 47

submicrus, Nesophontes, 11

suckleyi, Sorex, vi, 50, 74

Suncus, 49, 59

superstes, Nesophontes, 11

surinamensis, Crocidura, 9

Cryptotis, 5, 9

Sorex, 5, 6, 9, 10

taeniata, Talpa, 27

Talpa, v, 4, 6, 10, 14, 15, 19, 

20, 22–24, 27

Talpasorex, 6, 14, 20, 22, 28, 

talpoides, Blarina, v, vi, 28, 

31–33, 44

Corsira, 32

Sorex, v, vi, 28, 31–33, 44

Talposorex, 28

Talva, 20, 22

tamensis, Cryptotis, 44, 78

teculyas, Corsira, 75

telmalestes, Blarina, 32

temlyas, Corsira, 75

tenelliodus, Sorex, 72

tenellus, Sorex, 61, 63, 72, 79

tersus, Cryptotis, 41

texanus, Scalops, 20

Scalopus, 25

texensis, Scalops, 25

Scalopus, 25

thomasi, Blarina, 39, 43, 44

Cryptotis, 35–37, 39–41, 

43–45, 78

thompsoni, Microsorex, 58, 59

Sorex, vi, 50, 58, 59

thomsoni, Sorex, 59

townsendi, Scalops, 26, 27

Scapanus, 26

townsendii, Scalops, 25, 26, 27

Scapanus, 26, 27, 78

Talpa, 27

towsendii, Scapanus, 27

trigonirostris, Sorex, 75

tropicalis, Blarina, vi, 38, 44, 45 

Corsira, 44

Cryptotis, vi, 38, 44, 45, 78

trowbridgei, Sorex, 73

trowbridgii, Sorex, vi, 47, 53, 

73, 78

truei, Scapanus, 26

truii, Scapanus, 26

tundrensis, Sorex, 73, 78

turneri, Sorex, 47, 48

ugyunak, Sorex, 60, 70, 74, 79

umbrosus, Sorex, 57, 58, 

Urotrichus, v, 14, 17, 18

vagrans, Sorex, vi, 47, 50, 53, 

63–69, 72, 74, 75, 79

vancouverensis, Sorex, 75

venezuelensis, Cryptotis, 44, 78

ventralis, Sorex, 65, 75, 79

veraecrucis, Sorex, 47, 48, 60, 

71, 72, 75, 79

veraepacis, Sorex, vi, 45, 47, 

49, 51, 60, 61, 63, 75, 76, 

78, 79

verae-pacis, Sorex, 75

villai, Notiosorex, 46, 78

virginiana, Talpa, 22

virginianus, Scalopus, 20, 22

Talpa, v, 22, 23

virginea, Talva, 22

Talpa, 22

vulgaris, Sorex, 46, 48

wagneri, Sorex, 60

washingtoni, Microsorex, 58

willetti, Sorex, 66, 67

winnemana, Microsorex, 59

Sorex, 59

woodi, Solenodon, 12, 13, 14

Xenosorex, 35, 43

yakimensis, Scapanus, 27

yaquinae, Sorex, 69

yukonicus, Sorex, 62

zamicrus, Nesophontes, 11, 

12, 77

Zapus, 7

zeteki, Cryptotis, 40, 41
Albuja Viteri, Luís, 40

Alexander, Lois F., 63, 65, 68

Allen, Glover Morrill, 11, 44, 47, 70

Allen, Joel Asaph, 25, 28, 39, 40, 41, 44, 

58, 65

Alston, Edward Richard, vi, 44, 45, 49, 63, 75

Altobello, Giuseppe Carlo Emilio, 47

Anderson, Rudolph Martin, 29, 32, 60, 64, 

65, 74

Anthony, Harold Elmer, iii, 10, 11, 40,56 

Arredondo, Oscar, 11

Audubon, John James, 24

Ayres, William Orville, 25

Bachman, John, v, vi, 7, 8, 15, 19, 20, 21, 24, 

25, 26, 27, 28, 32, 33, 34, 35, 41, 42, 

47, 48, 51, 53, 57, 60, 61, 70

Bailey, Vernon Orlando, 20, 25, 63, 69, 75

Baird, Spencer Fullerton, v, vi, 17, 18, 19, 26, 

27, 28, 32, 35, 42, 46, 47, 49, 50, 51, 

53, 55, 57, 58, 59, 60, 64, 67, 73, 74, 

Baker, Robert James, 45, 46

Baker, Rollin Harold, 21, 24,

Bangs, Outram, 17, 20, 22, 26, 28, 32, 55

Barbour, Thomas, 12

Batchelder, Charles Foster, 56

Beddard, Frank Evers, 19, 25

Bee, James W., 73

Bell, Thomas, 14, 16

Benedict, Russell A., 34

Benson, Seth B., 62

Blainville, Henri-Marie Ducrotay de, 9, 14, 22

Blair, W. Frank, 34

Blanford, William Thomas, 56

Blyth, Edward, 15, 16

Boddaert, Pieter, 19

Bole, B. Patterson, Jr., 32, 42, 52, 55

Bond, Richard M., 62

Borisenko, A. V., 59

Brandt, Johann Friedrich von, iii, v, 3, 12, 

13, 14

Brisson, Mathurin Jacques, 6, 34, 46

Brooks, Fred E., 20, 22

Brown, Barnum, 32

Brown, Larry N., 59

Brown, Wennona A., 34

Bryant, Walter E., 17

Burgner, Robert L., 19

Burt, William H., 47

Butler, Amos W., 55

Cabrera, Ángel, 5, 9, 12, 36, 39, 40, 41, 44

Carraway, Leslie N., 37, 46, 48, 60, 62, 68, 

69, 71, 72, 75

Cassin, John, 27

Ceballos, Gerardo, 37

Chapman, Frank Michler, 22

Choate, Jerry Ronald, 30, 34, 35, 37, 38, 39, 

40, 41, 43, 44

Cook, Joseph A., 47

Cope, Edward Drinker, 32, 47, 48

Cory, Charles B., 22

Coues, Elliott, iii, 20, 24, 35, 37, 38, 45, 46, 

47, 48, 58, 67, 69

Cowan, Ian McTaggart, 68

Cox, Philip, 58

Cuvier, Georges [Jean Léopold Nicolas Frédéric, Baron Cuvier], 8, 20

Dalquest, Walter Woelber, 19, 27, 73

Davis, Joseph Anthony, 60

Davis, William Bennoni, 24, 34, 35

De Kay, James Ellsworth, v, vi, 18, 32, 46, 

51, 54, 57, 60

De la Faille, Clément, v, 15

Demboski, John R., 47

Desmarest, Anselme Gaëtan, 10, 15, 19, 22, 

27

Díaz, Alba, 39

Diersing, Victor E., 49, 58, 59, 

Dobson, George Edward, 48, 49, 51, 57, 62, 

70, 74

Authority Index
106 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Dokuchaev, Nikolai E., 62

Dolgov, V. A., 47

DoNascimiento, Carlos, 36

Durant, Pedro, 35

Duvernoy, Georges Louis, v, 

vi, 28, 30, 34, 42, 46, 47, 

55, 56

Elliot, Daniel Giraud, 17, 20, 

22, 24, 25, 26, 34, 35, 49, 

51, 55, 58, 64, 65, 66, 68, 

69, 70, 72, 75

Erxleben, Johann Christian 

Polycarp, 6, 9, 10, 15, 19, 

20, 22

Esteva, Martha E., 48, 63, 71, 72

Feagin, Jean E., 71

Findley, James S., 57, 63, 64, 

65, 66, 67, 68, 69, 72, 75

Fischer, Joanne Baptista, 22

Fischer, K., 11

Fischer von Waldheim, Gotthelf, 

iii, 1, 14, 15, 28, 46

Fitzinger, Leopold Joseph, 7, 9, 

47, 51, 60, 70

Forbes, William Alexander, 17

Gapper, Anthony, v, vi, 28, 31, 

32, 33, 44

Gaumer, George F., 38

Genoways, Hugh H., 30, 34, 

35, 37

Geoffroy Saint-Hilaire, Étienne, 

iii, v, 6, 7, 15, 20, 22, 28

Geoffroy Saint-Hilaire, Isidore, 

8, 51, 60

George, Sarah B., 34

Geptner, Vladimir Georgievich, 

47, 

Gill, Theodore, iii, 1, 12, 14, 19

Gilmore, Raymond Maurice, 

60, 74

Gilpin, John Bernard, 16, 47, 

51, 59

Gmelin, Johann Friedrich, 5, 6, 

7, 9, 22

Goldman, Edward Alphonso, 

39, 66

Goodwin, George Gilbert, 37, 

40, 41, 43, 56

Gray, John Edward, iii, 15, 28, 

32, 35, 40, 44, 46, 51, 

70, 75

Green, Morris Miller, 7, 47, 55

Grinnell, Joseph, 26, 67, 73, 74

Guevara, Lázaro, 38

Güldenstaedt, Johann Anton, 8

Günther, Albert Carl Ludwig 

Gotthilf, iii, 17

Hall, Eugene Raymond, 39, 47, 

60, 63, 65, 69, 73, 74

Hamilton, William J., Jr., 34

Handley, Charles O., Jr., 39, 

41, 44

Harlan, Richard, v, 15, 16, 19, 

20, 22, 27

Harper, Frances, 41, 42

Harris, Thaddeus William, 14, 

15, 16

Hasegawa, Masami, 10

Hennings, Darwen, 63, 64, 65, 

66, 68, 69

Hensel, Reinhold F., 65

Hermann, Johann, 7, 46

Herrick, Clarence Luther, 20, 

25, 55

Hibbard, Claude William, iii, 

20, 29, 45

Hodgson, Brian Houghton, 

56, 63

Hoffmann, Robert Shaw, 63, 

64, 65, 66, 68, 69, 70, 74

Hoffmeister, Donald Frederick, 49

Holder, Joseph Bassett, 7

Hollister, Ned, 47, 55, 57

Hooper, Emmet Thurman, 47

Hope, Andrew G., 47, 62, 64

Horsfield, Thomas, 63

Howell, Arthur Holmes, 22

Husson, A. M., 5

Hutterer, Rainer, iii, 17, 34, 43

Illiger, Carl, iii, 6, 14, 15, 20, 28

Jackson, Hartley Harrad 

Thompson, vi, 8, 21, 22, 

24, 25, 26, 27, 35, 37, 38, 

43, 47, 48, 49, 50, 52, 55, 

56, 57, 58, 59, 60, 61, 62, 

63, 65, 66, 67, 68, 69, 70, 

71, 72, 73, 74, 75

Jentinck, Fredericus Anna, 5, 9

Johnson, David Horn, 47

Johnson, Murray L., 27

Jones, Cheri A., 32

Jones, J. Knox, Jr., 37, 46

Junge, Jane Ann, 66, 70, 74

Kaup, Jakob J., 34, 46, 47

Kelaart, Edward Frederick, 59

Kellogg, [Arthur] Remington, 

11, 12, 58, 62, 73, 75

Kelson, Keith R., 39, 47, 65

Kerr, Robert, v, vi, 8, 9, 10, 18, 

39, 41, 46, 47, 48, 51, 52, 

53, 54, 55, 56, 57

Kretzoi, Miklós, iii, 28

Kruskop, S. B., 59

Kuroda, Nagamichi, 60, 74

Küster, H. S., 49

Layne, James N., 34

Le Conte, John, 20, 22, 24, 27

Lehmann, Joannes Georgius 

Christianus, 56

León Paniagua, Livia, 38

Lesson, Réné Primeverre, 14, 

20, 22, 28

Linares, Omar J., 39

Linnaeus, Carl, iii, v, 6, 7, 14, 

15, 16, 19, 20, 21, 22, 23, 

27, 28, 46, 47, 48, 

Linsley, James Harvey, 51

Long, Charles A., 59

Lönnberg, Einar, 36

Lord, John Keast, 17

Lowery, George H., 34

Lyon, Marcus Ward, Jr., 25, 35, 

42, 49, 58, 59, 74, 75

Maldonado, Jesús E., 66

Marcgrave, Georg, 6, 

Matson, John Orville, 38, 49, 

60, 61, 71

McAliley, Lee Rex, 45, 46

McCarthy, Timothy John, 60

Medina, César E., 37

Meigen, Johann Wilhelm, 47

Merriam, Clinton Hart, vi, 17, 

26, 32, 33, 34, 35, 37, 38, 

39, 40, 41, 42, 43, 44, 45, 

46, 47, 48, 49, 51, 55, 56, 

60, 61, 62, 63, 64, 65, 66, 

67, 68, 70, 71, 72, 73, 

74, 75

Miller, Gerrit Smith, Jr., 11, 12, 

17, 20, 21, 22, 26, 35, 37, 

38, 40, 41, 42, 43, 44, 46, 

49, 55, 57, 58, 62, 63, 64, 

65, 66, 72, 73, 75

Milne-Edwards, Alphonse, 35

Molinari, Jesús, 35

Moncrief, Nancy D., 30

Moreno Cárdenas, Pablo 

Alejandro, 40

Morgan, Gary S., 12

Moulthrop, Philip N., 32, 42, 

52, 55

Müller, Philipp Ludwig Statius, 

6, 20, 22

Murie, Adolph [Winstrom], 38

Nathusius, Hermann, 46, 48

Nelson, Edward William, 66

Oberholser, Harry Church, 22

Ognev, Sergei Ivanovitch, 47

Okada, H., 10

Okhotina, M. V., 48, 68, 73

O’Neill, Mark B., 45, 46

Ord, George, v, 31, 32

Ordóñez Garza, Nicte, 38, 49, 

60, 61, 71

Osgood, Wilfred Hudson, 25, 

26, 35, 42, 49, 58, 59, 62, 

65, 68, 69, 74, 75

Ottenwalder, José A., 12, 13, 14

Pacheco Torres, Victor R., 40, 43

Palmer, Fletcher G., 20, 25, 26, 27

Paradiso, John L., 17

Pari, Alexander, 37

Patterson, Bryan, 12

Pavlinov, I. Y., 59, 62

Péfaur, Jaime E., 35

Pennant, Thomas, 46, 56

Peters, Wilhelm Karl Hartwig, v, 

12, 13, 26

Peterson, Richard S., 52

Pino, Kateryn, 37

Pomel, Auguste, iii, 1, 20, 25, 

27, 28, 32, 34, 35, 44, 47, 

60, 70

Poole, Arthur J., 73

Poole, Earl L., 57

Preble, Edward Alexander, 58, 59

Quiroga-Carmona, Marcial, 35, 

36, 43, 44

Rafinesque, Constantine Samuel, v, 6, 7, 9, 16, 22, 24

Rajevsky, V. V., 59

Ramírez Pulido, José, 48

Rand, Austin Loomer, 60, 65, 74

Rausch, Robert L., 71

Rausch, Virginia R., 71

Reed, Katherine Milmine, 3

Rehn, James A. G., 17, 20, 55

Reichenbach, A. B., 55

Reumer, Jelle W. F., iii, 45

Rhoads, Samuel N., 24, 47

Rhymer, Judith M., 56

Richardson, John, v, vi, 7, 8, 27, 

48, 51, 52, 57, 69, 70

Rye, E. C., 17

Saban, Roger, 35

Sánchez Cordero, Víctor, 38

Savi, Paolo, 49

Say, Thomas, v, vi, 18, 28, 29, 

30, 35, 41, 42, 48

Schaldach, William. J., 35, 43

Schantz, Viola S., 73

Scheffer, Victor Blanchard, 27, 73

Schinz, Heinrich Rudolf, 6, 14, 

15, 19, 20, 22, 28, 47

Schmidly, David James, 24, 34

Schulze, Erwin, 47

Schwartz, Albert, 25, 56

Sealander, John A., 34

Seba, Albertus, v, 6, 10, 22, 23

Setzer, Henry W., 36, 40

Shaw, George, 10, 15, 22

Skalon, Vasily Nikolaevich, 59

Smith, Ronald W., 16, 32, 52, 61

Speer, Kelly A., 47

Stephens, Frank, 25, 26, 49, 64

Stewart, Donald T., 61

Stone, Witmer, 41

Storer, Tracey Irwin, 26

Stroganov, Sergei Ul’janovich, 

47, 60, 70, 74
NUMBER 650 • 107

Sundevall, Carl Jakob, 48, 55

Swarth, Harry Schelwald, 26

Talbot, Sandra L., 47

Tate, George Henry Hamilton, 

39, 41, 44

Thomas, [Michael Rogers] 

Oldfield, 9, 36, 39, 40, 41, 

44, 47

Timm, Robert Mark, 36, 38, 

39, 40, 43, 46

Tomes, Robert Fisher, 38, 44

Trouessart, Édouard-Louis, 34, 

44, 49, 72

True, Frederick William, iii, 19, 

20, 22, 24, 25, 26, 27, 42, 

49, 73

Turvey, Samuel T., 14

Valqui, Michael H., 40, 43

Van Hyning, Thompson, 15, 16

van Zyll de Jong, C. G., 74

Verrill, Addison Emery, 7, 64, 70

Viret, Jean, 3

Vivar, Elena, 36, 40, 41, 43

von Bloeker, John Christian, 

67, 74

Waddell, Peter J., 1, 3, 10

Wagler, Johann Georg, 14, 35, 47

Wagner, Johann Andreas, 16, 

20, 28, 51

Webster, William David, 29, 30

Woodman, Neal, 36, 38, 39, 40, 

41, 43, 44, 49, 51, 67, 71, 

72, 75

Yahontov, E. L., 59

Yates, Terry Lamon, 17, 22, 

24, 27

Yudin, Boris Stepanovich, 47, 

60, 70, 71, 74

Zapfe, Helmuth, 3

Zeballos, Horacio, 36, 37

Zimmermann, Eberhard August 

Wilhelm von, 6, 7, 15, 19, 

20, 22, 62
SUMMARY OF REQUIREMENTS FOR SMITHSONIAN CONTRIBUTIONS SERIES

For comprehensive guidelines and specifications, visit https://scholarlypress.si.edu. 

ABSTRACTS must not exceed 300 words. 

TEXT must be prepared in a recent version of Microsoft Word; use a Times font in 12 point for regular text; be doublespaced; and have 1′′ margins.

REQUIRED ELEMENTS are title page, abstract, table of contents, main text, and references.

FIGURES must be numbered sequentially (1, 2, 3, etc.) in the order called out; have components lettered consistently (in 

size, font, and style) and described in captions; include a scale bar or scale description, if appropriate; include any legends 

in or on figures rather than in captions. Figures must be original and must be submitted as individual TIF or EPS files.

FIGURE FILES must meet all required specifications in the Digital Art Preparation Guide. Color images should be 

requested only if required. 

TAXONOMIC KEYS in natural history manuscripts should use the aligned-couplet form for zoology. If cross-referencing 

is required between key and text, do not include page references within the key, but number the keyed-out taxa, using the 

same numbers with their corresponding heads in the text.

SYNONYMY IN ZOOLOGY must use the short form (taxon, author, year:page), with full reference at the end of the 

manuscript under “References.”

REFERENCES should be in alphabetical order, and in chronological order for same-author entries. Each reference should 

be cited at least once in main text. Complete bibliographic information must be included in all citations. Examples of the 

most common types of citations can be found at SISP’s website under Resources/Guidelines & Forms.