Subfamilies and 

Genera of the Soricidae

GEOLOGICAL SURVEY PROFESSIONAL PAPER 565
Subfamilies and 

Genera of the Soricidae

By CHARLES A. REPENNING

GEOLOGICAL SURVEY PROFESSIONAL PAPER 565

Classification, historical zoogeography, 

and temporal correlation of the shrews

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1967
UNITED STATES DEPARTMENT OF THE INTERIOR 

STEW ART L. UDALL, Secretary

GEOLOGICAL SURVEY 

William T. Pecora, Director

Library of Congress catalog-card No. GS 67-175

For sale by the Superintendent of Documents, U.S. Government Printing Office 

Washington, D.C. 20402 - Price 50 cents (paper cover)
CONTENTS

Page

Abstract.___-_--------__-_____________________-____ 1

Introduction.______________________________________ 1

Evaluation of characters...-_________________________ 3

Diagnoses and contents of subfamilies.____________ ____ 7

Subfamily Heterosoricinae Viret and Zapfe, 1951.____ 7

Genus Domnina Cope, 1873____________________ 7

Domnina thompsoni Simpson._-_-_--_.___ 8

Domnina gradata Cope._____--.---.._.._ 8

Domnina greeni Macdonald______________ 9

Domnina n. sp________________________ 9

Genus Paradomnina Hutchison, 1966____________ 10

Genus Trimylus Roger, 1885.__________________ 10

Trimylus compressus (Galbreath)_________ 11

Trimylus aff. neumayrianus (Schlosser)__ 11

Trimylus neumayrianus (Schlosser)______ 12

Trimylus dakotensis Repenning, n. sp_.__ 12

Trimylus roperi Wilson._________________ 13

Trimylus mawbyi Repenning, n. sp________ 13

Trimylus sansaniensis (Lartet)___________ 14

Genus Ingentisorex Hutchison, 1966.____________ 15

Subfamily Crocidurinae Milne-Edwards, 1868-1874. _ _ 15

"Sorex" pusilliformis Doben-Florin____________ 15

"Sorex" dehmiVirei and Zapfe________________ 15

Genus Crocidura Wagler, 1832..________________ 16

Crocidura sp., Butler and Hopwood_______ 17

Corcidura wongi Pei.____________________ 17

Crocidura taungsensis Broom.____________ 17

Crocidura kornfeldi Kormos__-___-_-__-_ 17

Crocidura obtusa Kretzoi________________' 17

Crocidura zorzii Pasa__________________ 17

Genus Diplomesodon Brandt, 1853_-__------_-_- 17

Diplomesodon fossorius Repenning._______ 18

Genus Feroculus Kelaart, 1852________________ 18

Genus Miosorex Kretzoi, 1959._________________ 18

Miosorex grivensis (Depe"ret)_____________ 19

Genus Myosorex Gray, 1838.___________________ 19

Myosorex robinsoni Meester._____________ 20

Genus Paracrocidura Balsac, 1956_____________ 20

Genus Praesorex Thomas, 1913_-_-_-__-___-_-__ 20

Genus Scuti orex Thomas, 1913_ ________________ 21

Genus Solisorex Thomas, 1924__________________ 21

Genus Soricella Doben-Florin, 1964.____________ 21

Genus Suncus Ehrenberg, 1832__-_---____-_____ 21

Suncus sp., Meester_____________________ 22

Genus Surdisorex Thomas, 1906._______________ 22

Genus Sylvisorex Thomas, 1905_________________ 22

Crocidurinae? incertae sedis________________________ 23

"Limnoecus" micromorphus Doben-Florin._______ 23

"Sorex" collongensis Mein______________________ 24

Crocidura?, Mein.____________________________ 24

Subfamily Limnoecinae Repenning, new name___. ____ 24

Genus Angustidens Repenning, n. gen___________ 25

Angustidens vireti (Wilson)_______________ 25

Genus Limnoecus Stirton, 1930.________________ 26

Limnoecus tricuspis Stirton_____________ 26

Limnoecus niobrarensis Macdonald-_ ______ 27

Diagnoses and contents of subfamilies Continued Page

Subfamily Soricinae Fischer von Waldheim, 1817.____ 27

Tribe Soricini Fischer von Waldheim, 1817._______ 29

Genus Crocidosorex Lavocat, 1951______________ 29

Crocidosorex piveteaui Lavocat.___________ 29

Crocidosorex antiquus (Pomel)____________ 29

Genus Antesorex Repenning, n. gen______________ 30

Antesorex compressus (Wilson)_____---_-_- 31

Genus Sorex Linnaeus, 1758.___________________ 31

Genus Drepanosorex Kretzoi, 1941 ___-____-____- 32

Genus Microsorex Baird, 1877_-___-_-_-_-_---_- 33

Genus Alluvisorex Hutchison, 1966___-___-___-_- 33

Genus Petenyia Kormos, 1934__________________ 34

Genus Blarinella Thomas, 1911____--__--__----- 34

Blarinella kormosi (Schlosser)____________ 35

Blarinella quadraticauda (Milne-Edwards). 35

Soricini incertae sedis_______________-_---------- 35

"Sorex" stehlini Doben-Florin.________--------- 35

"Sorex" hibbardi SulimskL_________-_____--_-_- 36

Genus Zelceina Sulimski, l962_-_-_------------- 36

Tribe Blarinini Stirton, 1930___________________ 37

Genus Adeloblarina Repenning, n. gen__ _________ 37

Adeloblarina berklandi Repenning, n. sp 37

Genus Cryptotis Pomel, 1848_ _ _ _ _ _ _ - - - - - 39

Cryptotis adamsi (Hibbard)______________ 39

Cryptotis"? meadensis Hibbard- .__-_____ 40

Cryptotis kansasensis Hibbard-___________ 40

Cryptotis parva (Say)___-_-_-_-__--_----- 41

Cryptotis mexicana (Coues). __________ 41

Genus Paracryptotis Hibbard, 1950 ___________ 41

Genus Shikamainosorex Hasegawa, 1957_________ 42

"Sorex" dehrieli KowalskL_________----_---_--- 42

Genus Blarina Gray, 1838__._..__--------- - 42

Blarina gidleyi Gazin ______-_-_------- 43

Genus Blarinoides Sulimski, 1959.___-_--__---_- 44

Blarinini? incertae sedis.___________------------- 44

"Sorex" kretzoii Sulimski______________-_--_-- 44

Peisorex pohaiensis Kowalski and Li __________ 45

Tribe Neomyini Repenning, new name.____-_----- 45

Genus Neomys Kaup, 1829___--------_--------- 46

Genus Petenyiella Kretzoi, 1956_____-_---------- 46

Genus Episoriculus Eilerman and Morrison-Scott,

1951.___--__.___-------------------- 47

Episoriculus gibberodon (Petenyi) ___-__--- 48

Genus Chodsigoa Kastschenko, 1907__________-_- 48

Chodsigoa bohlini (Young)_________-___-- 50

Genus Beremendia Kormos, 1934_____----------- 50

Genus Nesiotites Bate, 1945.______-----_------- 51

Genus Nectogale Milne-Edwards, l870__-_----__- 51

Genus Soriculus Elyth, 1854._______ ____--_- 52

Genus Chimarrogale Anderson, 1877__---_-_----- 53

Genus Anourosorex Milne-Edwards, 1870. _______ 53

Genus Amblycoptus Kormos, 1926____----------- 54

Genus Hesperosorex Hibbard, 1957__-_---------- 55

Genus Notiosorex Baird, 1877_---------------- 55

Genus Megasorex Hibbard, 1950.________---_--- 56

Neomyini? incertae sedis___________-__--------- 57

Deinsdorfia franconia Heller._______----_------- 57

m
IV CONTENTS

Diagnoses and contents of subfamilies Continued 

Subfamily Allosoricinae Fejfar, 1966.__________

"Sorex" gracilidens Viret and Zapfe._______

Genus Allosorex Fejfar, 1966._____________

Soricidae incertae sedis_______________________

Podihik kura Deraniyagala._______________

Page 

57 

57 

57 

57 

57

Diagnoses and contents of subfamilies Continued 

Names of fossil genera not used in text_________

Ancestry of the Soricidae_______________________

Paleozoogeography and correlation______________

Key to living and fossil genera of the Soricidae____

References.___________________________________

Index_ _______________________________________

Page 

58

58

59

63

66

71

ILLUSTRATIONS

[Figs. 1-41 are drawings of the Soricidae]

FIGURE 1. Taxonomically significant differences.

2. Domnina thompsoni.

3. Domnina gradata. ______

4. Domnina n. sp_________

5. Trimylus compressus.

6. Trimylus dakotensis _____

7. Trimylus mawbyi _______

8. Crocidura russu'a. ______

9. Diplomesodon pulchellum10. Miosorex grivensis______

11. Myosorex varius. _______

12. Praesorex goliath. _______

13. Suncus caeruleus.. ______

14. Surdisorex norae. _______

15. Sylvisorex lunaris. ______

16. Angustidens vireti. ______

17. Limnoecus tricuspis.

18. Limnoecus niobrarensis_ _

19. Antesorex compressus....

20. Sorex araneus. _________

21. Microsorex hoyi- ________

Page

9

10

11

12

14

16

17

18

19

20

22

23

23

25

26

27

30

31

32

Page

22. Alluvisorex chasseae- _____________________ 33

23. Petenyia hungarica-.--------------------- 34

24. Blarinella quadraticauda __________________ 35

25. Adeloblarina berklandi-.------------------ 38

26. Cryptotis adamsi,.---------- ------------- 39

27. Cryptotis parva------------------------ _ 1 40

28. Paracryptotis rex...---------------------- 41

29. Blarina brevicauda- ______________________ 43

30. Blarina gidleyi-------------------------- 43

31. Neomys jodiens-------------------------- 46

32. Episoriculus caudatus_____________________ 47

33. Chodsigoa hypsibia,---------------------- 49

34. Chodsigoa salenskii - ____________________ 49

35. Beremendia fissidens______________________ 50

36. Nectogale elegans ----------------------- 51

37. Soriculus nigrescens. _____________________ 52

38. Chimarrogale himalayica,------------------ 53

39. Anourosorex squamipes. __________________ 54

40. Notiosorex crawfordi.--------------------- 56

41. Megasorex gigas__________________________ 56

42. Diagram of a phylogeny of the Soricidae ___ 61
SUBFAMILIES AND GENERA OF THE SORICIDAE

By CHARLES A. KEPENNING

ABSTRACT

The shrews constitute a distinct and uniform but phylogenetically diverse family in which five subfamilies, one subdivided into three tribes, can be recognized by consistent morphologic differences. These differences are traceable back 

through the late and middle Tertiary shrews and point to a 

hypothetical common ancestor. The trends of the morphologic 

change within the subfamilies rather clearly indicate what this 

ancestor should look like. Because the great majority of fossil shrews are represented only by teeth and mandibles, the 

morphologic characters here used to classify most shrews pertain to these parts of the animals.

The extinct subfamily Heterosoricinae is known from the 

Oligocene to the Pliocene and in both the Old and the New 

Worlds. Four genera are recognized in North America; one 

of them is known also from the Old World. At least 12 species are included in these genera. The subfamily is not closely 

related to other shrews, and all shrews in the subfamily lack 

an internal temporal fossa on the mandible, have extreme lingual offset of the lower mandibular articulation, and retain 

at least some pigmentation in the teeth.

The living subfamily Crocidurinae is known from the middle 

Miocene to the Recent in the Old World. In the subfamily are 

11 to 13 fossil species and 10 living genera. The. subfamily 

appears most likely to be of Austro-Tethyan origin (mainly 

Ethiopian Africa and peninsular India) and is not well represented in the known fossil record. It is characterized by an 

internal temporal fossa on the mandible, by little or no separation of the articular facets on the mandibular condyle with 

a labial emargination of the interarticular area, by the retention of a primitive triangular P4, and by no pigmentation of the 

teeth.

The extinct subfamily Limnoecinae is known from the early 

Miocene to the middle Pliocene in North America and closely 

parallels the Old World crocidurines in morphology. Two 

genera and three species are recognized. The subfamily is 

characterized by crocidurine features except that P4 is modifled from a triangular tooth to one in which a central crest 

runs from the apex of the single cusp to the posterior cingulum; at least some pigmentation is present in the teeth.

The living subfamily Soricinae is known from the late Oligocene to the Recent in the Old World and from the early 

Miocene to the Recent in North America. It is characterized 

by an internal temporal fossa on the mandible, by little to great 

separation of the articular facets on the condyle of the mandible with a lingual emargination of the interarticular area, by 

the modification of the primitive triangular cusp of Pt into a 

labial shearing blade with a posterolingual basin, and by retention of tooth pigmentation in most, but not all, genera. 

This subfamily is very large and is divisible into three tribes

largely on the basis of masticatory modifications; this basis of 

separation is supplemented by characters in the teeth. The 

tribe Soricini contains at least 3 extinct genera, about 30 extinct species, and 4 living genera. It is characterized by little modification of its mandibular articulation or teeth and by 

a prominent entoconid crest on lf±. The tribe Blarinini contains at least 4 extinct genera, 10 extinct species, and 2 living 

genera. It is characterized by slight to great modification of 

its mandibular articulation with a broad interarticular area, 

no entoconid crest on M» and by a trapezoidal M2 . The tribe 

Neomyini contains 4 fossil genera, 13 fossil species, and 9 living genera. It is characterized by great modification of its 

mandibular articulation with a narrow interarticular area, an 

entoconid crest on Mt in most forms, and a prominent bifid 

upper incisor in most Old World forms.

The extinct subfamily Allosoricinae is known from the late 

Miocene to the late Pliocene in Europe. It is characterized by 

unusual carnivorelike cheek teeth that are unequaled in specialization by any other shrew. Two genera (one unnamed) 

are included in this subfamily.

INTRODUCTION

The living shrews are generally placed in two subfamilies with the exception of the African armored 

shrew Scutisorex, which some workers (for example, 

Alien, 1917") have placed in a separate subfamily. The 

Soricinae are holarctic in distribution, and the Crocidurinae are entirely of Old World distribution and 

largely tropical. As stated by Ellerman and Morrison-Scott (1951, p. 41), this division into two subfamilies "seems to be based chiefly on the Soricinae 

having teeth pigmented at the tips, and the Crocidurinae having teeth entirely white." These authors, 

Friant (1947), and several others have pointed out 

that, although the Soricinae normally have pigmented 

teeth, pigmentation in the subfamily is variable 

very light in several genera and entirely missing in 

a few genera. Because of this variability, several 

zoologists, including Ellerman and Morrison-Scott 

(1951) and Heim de Balsac and LaMotte (1956, p. 

141-142), have not recognized subfamilies of the 

Soricidae; however, others, including Friant, have 

recognized them.

In considering fossil shrews, there has been a 

similar tendency to abandon the subfamilies of living

1
SUBFAMILIES AND GENERA OF THE SORICIDAE

shrews (Hibbard, 1950, p. 129; and Viret and Zapfe, 

1951, p. 414). With the study of fossil shrews, however, greater attention has been directed toward additional criteria to recognize suprageneric taxa within the Soricidae. Although he preferred to consider 

his groups as of less than subfamily rank, Stirton 

(1930, p. 223-225) reviewed and grouped many of 

the living shrews to evaluate a Miocene fossil. He 

selected five divisions of the family, which he called 

groups, on the basis of eight features (10 features 

for the Anourasorex group). Those features that 

Stirton used to distinguish his groups include tooth 

pigmentation, shape of the internal temporal fossa 

of the mandible (fossa pterygoidea, intertemporal 

fossa, or posterointernal ramal fossa of authors), 

degree of separation of the condyles of the mandible 

(or articular facets on the condyle), height of tooth 

cusps, separation of the tips of the protoconid and 

the metaconid, position of the foramen ovale, form 

of the talonid of Mz, and nature of the entoconid and 

basin of the talonid of J/i and M2 .

Earlier Schlosser (1924, p. 5) suggested that the 

absence of secondary cuspules on the cutting edge of 

the lower incisor and the shape of the ascending 

ramus of his species Crocidura, korm^osi were distinctive of the genus.. Still earlier Miller (1912, p. 30, 

86) pointed out that the Crocidurinae could be separated from the Soricinae on the basis of the structure 

of P4 , Lavocat (1951, p. 23-24), following Miller's 

criterion, seems to have been one of the first to place 

much significance on the morphology of P4 in fossil 

shrews. He regarded his Crocidosvrex piveteaui as 

intermediate between Crocidura and Sorex largely on 

the basis of the form of P±. Wilson (1960, p. 35, 36, 

38) also placed emphasis on the form of the F4 in 

Sorex. The slightly more anterior average position 

of the mental foramen in the Crocidurinae has also 

been mentioned by several authors as of possible value 

in distinguishing this subfamily from the Soricinae.

Therefore, it seems that there are a large number 

of dental and mandibular features that may be useful in separating the Crocidurinae from the Soricinae 

besides tooth pigmentation, which has been considered 

to be a fallible criterion. Cranial and postcranial 

skeletal features, as well as criteria based upon the 

soft parts, may also be useful, but with very few exceptions these are not known in the fossil record. A 

review of the above features, of the dental formulae, 

and of the morphology of P4 and J/1 in most fossil 

species and in most living genera leads to the conclusion that subfamilies are recognizable in both living 

and fossil shrews. Five subfamilies are clearly recognizable in the fossil record: the Crocidurinae and the

Soricinae, representing the surviving subfamilies; and 

the Heterosoricinae, the Allosoricinae, and the Limnoecinae (new name), representing extinct groups of 

subfamily rank. The living subfamilies are of approximately equal size; there are 21 soricine species 

and 21 crocidurine species in the Palearctic and Indian 

regions (Ellerman and Morrison-Scott, 1951), no 

soricines occur in Africa, and there are no crocidurines in North America.

Specimens of nearly all North American fossil 

shrews older than Pleistocene were examined during 

this review as were specimens representing all but 

two x living genera of the world; however, specimens 

of only six Old World fossil species were examined. 

In this review the description of Old World fossil 

forms is largely from published accounts. Such a 

review has obvious disadvantages as the criteria here 

utilized are not always discussed or clearly figured 

in published descriptions. For this reason little attempt to revise the nomenclature of the Old World 

fossil shrews is made, although they are placed in the 

subfamilies which seem most appropriate as based 

upon published descriptions. The specimens examined 

are listed in the discussion of the taxa they represent.

Subfamily, tribe (where applied), genus, and many 

extinct species diagnoses are given in terms of the 

criteria here used. For many shrews, particularly 

for the living genera, other criteria are available in 

the literature to enlarge the diagnoses but these criteria are not mentioned here because they cannot be 

applied uniformly to both living and fossil shrews. 

This does not imply that other criteria are not valid 

or useful.

The illustrations are somewhat stylized but were 

done carefully with a camera lucida attached to a 

microscope. There is an advantage in having all 

forms illustrated by the same technique and by one 

person. The illustrations are not drawn to a uniform 

scale because size has appeared to be of no more than 

specific significance. When possible, P^ MI, and Ms 

are shown in both occlusal and buccal view; P 4 and 

M 1 in occlusal view. M 2 almost always parallels 

the development of M t and hence is not shown in the 

illustrations, even of types. In the few shrews where 

M 2 shows noticeable difference from M i, the difference 

is noted in the text. The mandible is drawn in lingual 

view, and the condyles of the mandible are shown in 

posterior view.

Embryological studies to determine the dental 

formulae of the soricids in terms of incisor, canine, 

premolar, and molar teeth (Xrnbach-Christie-Linde,

i Feroculus and Solisorex from Ceylou. Photographs of British 

Museum (Natural History) specimens of these genera were examined.
EVALUATION OF CHARACTERS

1912; Kindahl, 1960) are not in entire agreement and 

the formulae seem to vary with the genus under study. 

Although this work is certainly important in evaluation 

of the phylogeny of the Soricidae, it has not been considered in this review. Dental formulae given in this 

report are arbitrarily based upon the frequent use of 

the term "antemolar" in published descriptions of 

shrews. The term applies to those teeth between the 

soricid incisor and the molars. The formula used is: 

Total=soricid incisor-antemolars including P 4-molars/ 

soricid incisor-antemolars including P4-molars

Acknowledgments. The following persons have 

loaned or donated specimens used in this review and 

are here sincerely thanked: Kazimierz Kowalski, 

Polish Academy of Science, Krakow; Oldrich Fejfar, 

Central Geological Survey of Czechoslovakia, Prague; 

R. W. Hayman, British Museum (Natural History), 

London; David H. Johnson and C. Lewis Gazin, U.S. 

National Museum, Washington; Barbara Lawrence, 

Harvard University, Cambridge; Malcolm C. McKenna and Sydney Anderson, American Museum of 

Natural History, New York; Claude W. Hibbard, 

University of Michigan, Ann Arbor; Rainer Zangerl, 

Philip Hershkovitz, and Joseph C. Moore, Chicago 

Natural History Museum, Chicago; William A. 

Clemens, University of Kansas, Lawrence; Peter 

Robinson, University of Colorado Museum, Boulder; 

Richard H. Tedford, University of California, Riverside; J. Reid Macdonald and Charles A. McLaughlin, 

Los Angeles County Museum, Los Angeles; Seth B. 

Benson and Ruben A. Stirton, University of California, Berkeley; and J. Arnold Shotwell, University 

of Oregon, Eugene.

S. B. Benson, C. W. Hibbard, J. H. Hutchison, 

Miklos Kretzoi, D. E. Savage, R. A. Stirton, Heinz 

Tobien, and F. C. Whitmore, Jr., have reviewed the 

manuscript and have offered many helpful suggestions. The author has profited from extended discussions of the crocidurine shrews by P. M. Butler, 

University of London.

EVALUATION OF CHARACTERS

Of the features observed in the dental and mandibular morphology during this review, the structure of 

the P4 is most useful in the recognition of subfamilies. 

(See fig. 1.) The structure of the mandibular condyle 

is nearly as useful, and pigmentation of the teeth 

seems significant but is subject to exceptions. The 

absence of the internal temporal fossa is a subfamily 

character of the Heterosoricinae. The other morphologic characters, when considered singly, appear to be 

of taxonomic value only after the identity of the 

subfamily has been established. In combination, most

characters may be mutually supporting of subfamily 

assignment, and some are of subfamily importance 

when developed to an extreme degree. Most appear 

to represent evolutionary trends in several shrew 

lineages that are parallel in direction but not equal in 

rate of change.

During this review it has been found that in the 

characters here examined the shrews are remarkably 

stable in comparison with other mammals. Unless 

otherwise noted, the reliability of these characters 

does not seem to be reduced by individual variability. 

Atavism is rather rare although it does occur in living 

and presumably in fossil forms.

Structure of P4

Domnina gradata Cope (Patterson and McGrew, 

1937) is, in several respects, including the structure 

of the P4 , ideal as the prototype of all soricids. In 

other respects it is clearly a heterosoricine shrew. In 

occlusal view the P4 of D. gradata has a simple equilaterally triangular outline and a single large triangular cusp (figs. 1, 3). There is a posterior basal 

cingulum that extends nearly to the anterior corner 

of the tooth on both the lingual and the labial sides. 

Although the cingulum is prominent at the posterolabial corner, it does not overhang the root and cover 

the alveolus as in most modern shrews. The posterior 

surface of the crown slopes uniformly to the cingulum 

and is slightly concave between the posterolateral 

corners of the crown. The posterolateral corners of 

the triangular cusp are sharp so that slight ridges 

run from the apex of the cusp to the cingulum at the 

posterolabial and posterolingual corners of the tooth. 

The -P4 is the largest tooth of the antemolar series with 

the exception of the procumbent incisor and possibly 

the first antemolar, which is unknown. The antemolar 

alveoli of some species of Trimyhis are similar to those 

of Domnina gradata, but P4 is not the largest postincisor antemolar in Trimylus.

Domnina is here placed in the subfamily Heterosoricinae on the basis of features other than the crown 

of P4 . The P4 in the genera Trvmylus and Ingentisorex appear to be similar to that of Domnma except 

that details of the cingulum and cusp morphology 

have been lost in the development of a rounded, bulbous tooth. The P4 of Paradomnina is like that of 

Domnina.

The basic triangular pattern that is retained in the 

Crocidurinae differs only slightly from that in Domnma,. In the Soricinae, however, the crown is converted into an oblique blade by emphasis of a crest 

connecting the anterior cusp with the cingulum at the 

posterolabial corner of the tooth and by suppression
SUBFAMILIES AND GENERA OF THE SORICIDAE

Lingual

Anterior

Posterolingual 

basfn

Posterior

Heterosoricinae

and 

Crocidurinae

Limnoecinae Soricinae

OCCLUSAL PATTERNS OF LEFT P.

Labial 

emargination Lingual 

emargination

Heterosoricinae Limnoecinae

and 

Crocidurinae

Subfamilies of the Soricidae

Soricinae

Broad

interarticular 

area

Narrow

interarticular'

area

Soricini Blarinini Neomyini

Tribes of the Soricinae 

POSTERIOR PATTERNS OF LEFT MANDIBULAR CONDYLES

FIGURE 1. Taxonomically significant differences in structure of P4 

and mandibular condyles.

of the anteroposterior lingual crest and development 

of a basin in its place. In most soricines the labial 

blade combines with the posterior cingulum to form a 

continuous L-shaped crest which encloses the posterolingual basin on both labial and posterior sides. The 

labial crest of the triangular P4 is also accentuated in 

the Limnoecinae, but it has shifted lingually nearly to 

a median position. The lingual crest of the triangle 

is almost completely suppressed but no posterolingual 

basin has formed because of the medial position of the 

labial crest.

These basic patterns of the P4 are diagrammatically 

shown in figure 1. With few exceptions these patterns

are clearly recognizable in all shrews considered in 

this review. The exceptions are all within the group 

of heavy-toothed and stout-jawed soricines where 

other characters, such as widely separated mandibular 

condyles, leave no doubt of subfamily affinity.

No soricid has ever developed a molariform P4 , and 

for this reason the late Eocene Scrtwrnimia gracttis 

Stehlin (1940), which has a well-developed trigonid 

and talonid in its P±, cannot be considered ancestral 

to any shrew; as recognized by Stehlin (1940, p. 306), 

Saturnima, appears to be an erinaceoid similar to 

Leptacodon.
EVALUATION OF CHARACTERS

Pigmentation

Pigmentation in soricid teeth seems to be remarkably stable insofar as fossilization is concerned and 

is usually preserved even in Oligocene specimens. It 

may be obscured, however, by coloration of the entire 

tooth during fossilization or altered to such a degree 

that it is not visible in normal light. In some genera, 

when its presence is not detectable in ordinary light, 

it can be seen under ultraviolet light as described by 

Patterson and McGrew (1937, p. 247). Hibbard 

(1950, p. 128) noted that reaction of pigmentation to 

ultraviolet light is not consistent in either Recent or 

fossil specimens that he examined.

In many shrews, pigmentation is present only at 

the tips of the cusps and along the crests of the lophs 

so that heavy wear removes all pigmented parts of 

the tooth. Although pigmentation is usually of a 

very conspicuous color, the teeth of some shrews, like 

Nectogale, are pigmented over much of their surface, 

but the pigmentation is of such a light value and 

slight saturation that it may be overlooked. Apparent 

lack of pigmentation in a fossil shrew, therefore, must 

be evaluated cautiously, particularly if the teeth have 

been darkened by fossilization.

Nevertheless, pigmentation is an important character in the recognition of the subfamilies of the 

Soricidae. The teeth of Domnina gradata are extensively pigmented (the pigment is a pale yellowish 

orange), indicating that pigmentation was acquired 

early in the history of the Soricidae (Patterson and 

McGrew, 1937, p. 247) or was inherited from the 

soricoid ancestor of the family. In Trwvylus, pigmentation is present but restricted; it is certainly 

present on the lower incisor, on PI, and on the upper 

incisor. In the Crocidurinae and in the Allosoricinae, 

tooth pigmentation is lost. In the Limnoecinae it 

seems to be present in most specimens, but Macdonald 

(1947, p. 125) and also Hibbard (1957, p. 329) stated 

that Limnoecus niobrarensis Macdonald shows no pigmentation. James (1963, p. 36) pointed out, however, 

that in Macdonald's type the teeth are worn to a level 

beneath the base of the pigmented area on other specimens of the genus. In the Soricinae, pigmentation is 

characteristic but is very light in some genera and 

absent in a few; some soricines have been classified as 

crocidurines because of the lack of tooth pigmentation.

Internal temporal fossa of the mandible

This fossa (also called fossa pterygoidea, intertemporal fossa, and posterointernal ramal fossa by other 

authors) is not pocketed in the Heterosoricinae (fig. 

2). It is deeply pocketed in all other subfamilies 

(figs. 8-41), and in these groups its form seems to be

of no more than generic importance as far as diagnosis 

of most shrews is concerned.

The soricid internal temporal fossa is a consequence 

of the loss of the zygomatic arch and much of the 

masseter muscle. The internal part of the temporal 

muscle assumes the function of the masseter in this 

situation, and its insertion on the medial side of the 

coronoid process is enlarged and is shifted ventrally, 

to a level approximating that of the masseter in other 

mammals, for a more masseterlike force and leverage. 

Loss of the zygomatic arch and development of the 

internal temporal fossa also are associated with loss 

of the masseteric fossa on the lateral side of the 

mandible, for the same reason. With the exception 

of heterosoricines, no shrew has a masseteric fossa; 

also, the heterosoricines do not have a soricine internal 

temporal fossa and have retained a reduced zygomatic 

arch.

Patterson and McGrew (1937, p. 256-257), apparently in a lapse of memory while thinking of the 

internal temporal fossa, stated that a deep masseteric 

fossa "extending downward much nearer the ventral 

margin of the jaw" (than in the bat Mvniopterw) 

is a soricid character. This and several other features 

led them to place the genus Mystipterus Hall in the 

Soricidae rather than in the Chiroptera. The deep 

masseteric fossa of Mystipterus clearly indicates, however, that the animal has a normal mammalian masseter system because the fossa cannot form without 

the masseteric muscle.

Hall's illustration (1930, pi. 38b) shows that the 

masseteric fossa of Mystipterus is well developed, and 

his opinion that the genus is a bat seems much more 

reasonable than other opinions which label it a shrew. 

The lack of reduction in the size of the talonid of MS 

(the only tooth known) and the full development of 

the V-shaped pattern on the talonid and on the 

trigonid of M3 suggest, however, that Mystipterus may 

be a talpid. It resembles Mydecodon martini Wilson 

(1960, fig. 32) and is somewhat less like the living 

Neurotrichus.

Mandlfoular condyle

Domnina (D. thompsoni Simpson, see McDowell, 

1958, p. 181-182, fig. 30E) has virtually one broad 

mandibular condyle which is constricted and offset 

in its center foreshadowing the doubled articular 

facets of later soricids. Separation, to form two 

articulations, has chiefly taken place in the early 

Miocene shrews although in some the articular facets 

may still be joined. This situation is little changed 

in the later limnoecines and in the modern crocidurines. Some of the living soricines and the extinct

240-305 O-67 2
6 SUBFAMILIES AND GENERA OF THE SORICIDAE

allosoricines have greatly accentuated the separation 

of the facets.

On many mandibular condyles it is difficult to 

clearly distinguish the extent of the articular facets. 

Compare, for example, the variations in interpretation 

of the facets on Limnoecus niobrarensis made by Hibbard (1957, fig. 3A), by James (1963, fig. 9c), again 

by James (1963, fig. 11), and in this report (fig. 18). 

All were made by careful examination of the same 

specimen.

There are three basic patterns to be seen in the 

mandibular condyles of the shrews (fig. 1). In nearly 

all shrews these are recognizable regardless of the 

degree of separation of the articular facets, regardless 

of the vagueness of the limits of the articular surfaces, and regardless of the antiquity of the shrew 

considered insofar as the present record is known.

In the subfamily Heterosoricinae the lower articular 

facet of the mandibular condyle is not much offset 

ventrally from the upper facet (more so in Trimytus 

than in Donwdna) but is placed lingually to an extreme degree so that virtually no part of the lower 

facet lies below the upper facet. This situation is 

not found in any other subfamily.

In the subfamilies Crocidurinae and Limnoecinae, 

the lower facet of the mandibular condyle is beneath 

the upper, and the two facets are continuous along 

the lingual side of the posterior face of the condyle. 

If separated, an elevated and usually rounder bridge 

on the lingual side of the posterior face of the condyle, 

the interarticular list of Sulimski (1959, p. 128-129), 

connects the two facets, and there is an excavation 

of the interarticular area on the labial side of the 

posterior face of the condyle. This condylar structure 

is invariable in these two subfamilies but in a few 

shrews, as in the giant species of Suncus, the articulating surface connecting the upper and lower facets 

is so broad that it is impossible to decide whether 

the interarticular excavation is on the lingual or labial 

side of the posterior face of the condyle.

In the subfamily Soricinae the lower facet is below 

the upper in posterior view, and the two facets are 

continuous along the labial side of the condyle. If 

separated, as they are in all known forms younger 

than middle Miocene, the two articular facets are 

connected by an interarticular list that is on the labial 

side of the posterior face of the condyle, and the 

lingual side of the interarticular area is excavated. 

This pattern is also invariable, but the position of the 

articular surfaces is otherwise greatly variable in the 

subfamily, largely in degree of anterior placement of 

the lower facet. This variation in articulation is not 

known in other subfamilies, the most extreme exceptions being in the extinct crocidurine Diplomesodon 

fossorvus Repenning (1965) and on the extinct allosoricine AUosorex stenodus Fejfar (1966).

Height of cusps

There is a general trend throughout the soricids 

to increase the crown height in the evolution of the 

separate lineages, except in the Heterosoricinae; however, relative cusp height is difficult to judge. In this 

review it has been found easier, in most shrews, to 

estimate the stage of evolution by observing the relative height above the labial cingulum at which the 

bottom of the reentrant valley between the protoconid 

and the hypoconid opens on the labial side of M±. 

In Domnina and the many earlier Miocene shrews, 

this reentrant opens at or very close to the level of 

the cingulum. In many of the more advanced forms, 

there is an appreciable distance between the reentrant 

and the cingulum. In many living shrews, however, 

this is not so and the value of this character is quite 

variable with different lineages. Although partly 

related to cusp height, this feature may also be related 

to a tendency of the metalophid-protolophid junction 

to shift slightly labially in geologically later genera.

Heel of M3

Reduction of the talonid of the M3 in Croddura to 

a single trenchant cusp has frequently been cited as 

a character distinctive of the crocidurines. It is prevalent in this subfamily but is also found in some 

soricines, like Cryptotis, It also occurs in the Limnoecinae but not in the Heterosoricinae. The heel of 

Ms is virtually lost in a few shrews, and the entire 

Mz is missing in five genera.

Position of the mental foramen

The mental foramen lies beneath the middle of Mi or 

the anterior root of M2 in the Heterosoricinae; beneath 

P4 to beneath the anterior root of Mj in the Crocidurinae; beneath or slightly ahead or behind the anterior 

root of MI in the Limnoecinae; and in the Soricinae 

beneath the middle of M± in most Miocene to Recent 

species examined but beneath the F4 in Croddosarew and 

in Antesorex, new genus, of the late Oligocene to early 

Miocene. In all lineages it tends to have shifted 

posteriorly in geologically younger forms. The position 

of the foramen seems fairly constant within the subfamilies but the overlap in position between subfamilies 

is great. In addition there have been a very few 

individuals of living shrews noted during this review 

that have either doubled mental foramina or have the 

mental foramen significantly anterior to its position in 

other individuals from the same local population.
DIAGNOSES AND CONTENTS OF SUBFAMILIES

In the figures, the mental foramen is usually shown 

with the labial view of the lower teeth. These teeth are 

generally separated for clearer illustration, and a line is 

shown between the foramen and the tooth (either P4 or 

MI) with which it is properly oriented.

Closeness of the protoconld to tlie metaconid on Ma

Stirton (1930, p. 244) cited the closeness of these two 

cusps in his "Crocidura group" as being distinctive. 

Although it seems that it might reflect the height of cusp 

to some degree, it does appear to be characteristic of the 

living crocidurines and particularly of the extinct 

limnoecines. I have had considerable difficulty in making the comparison with even moderately worn teeth 

and consider it virtually impossible to judge the closeness of these cusps in figures of worn teeth. As a consequence, the character has not been considered to be of 

much value in this review except where the two cusps are 

obviously close. In many shrews where it seems of 

importance, an anterior view of MI is figured.

Structure of P* and M1

There appears to be no subfamily distinctiveness in 

these teeth other than for the Heterosoricinae. This 

subfamily is alone in the absence of an emargination of 

the posterior basal outline of the tooth in all forms and 

in the consequent failure to develop a posterior projection of the hypoconal flange. This structure is otherwise characteristic of most, but certainly not all, genera 

of the Soricidae.

Miscellaneous features

Secondary cuspules on the cutting edge of the lower 

incisor seem to have no significance in subfamily characterizations or in evolutionary patterns. The dental 

formulae show a broad trend toward reduction in number of antemolars, but the trend cannot be correlated 

with subfamily distinctions. Overlap of the alveolus 

and part of the ramus by the labial cingulum of P4 is 

most extreme in some soricine genera but is present to 

some degree in both limnoecine and crocidurine shrews. 

A prominent entoconid on MI is characteristic of most 

shrews, but its variation in prominence and in position 

seems to be a tribal or generic character. A depression 

on the labial surface of the horizontal ramus in the 

region of the mental foramen is also a generic character 

in the Heterosoricinae and in some Soricinae. A bifid 

or fissident upper incisor is conspicuous in the Heterosoricinae and in the tribe Neomyini; and is slightly 

developed in many incisors in the tribe Soricini. It is 

known in only one genus of the Crocidurinae. This 

feature is formed by a smaller cusp branching from the 

medial side of the principal (anterior) hooklike cusp of 

the falciform first upper soricid incisor. The occlusal

outline of M2, and the position of the zygomatic process 

of the maxillary relative to M2, the nature of development of the superior pterygoid fossa of the mandible, 

the extent of the external temporal fossa of the mandible, and a few other features are characters of tribal 

significance in the Soricinae.

DIAGNOSES AND CONTENTS OF SUBFAMILIES 

Subfamily HETEROSORICINAE Viret and Zapfe, 1951

P± has triangular cusp, little or no posterior sulcus on 

cusp, and root transversely broadened or even doubled 

in most species; MI low cusped reentrant between 

protonid and hypoconid opens at level of cingulum; Ms 

has double-cusped or crescentic talonid (this tooth is 

absent in Ingentisorex from the late Miocene of Oregon) ; P4 and M1 have no emargination of posterior 

basal outline and no posterior projection of hypoconal 

flange; in Trimylus, M2 is trapezoidal and the zygomatic 

process of maxillary originates opposite it, but in the 

less specialized Domnina, M2 is rectangular and the 

zygomatic process originates behind this tooth; tooth 

pigment present at least to some degree; masseteric fossa 

well developed; internal temporal fossa lacking or very 

shallow and never pocketed; ventral condyle of mandible offset lingually much farther from plane of mandible 

at lower sigmoid notch than in any other subfamily of 

shrews and at approximately right angles to vertical 

axis of horizontal ramus; mental foramen below middle 

of MI to below anterior root of M2 and usually associated 

with depressed area on labial face of mandible or with 

groove running anterodorsad from foramen to region 

of P4 , or both; a small foramen of unknown function 

(Mawby, 1960, p. 955) on lingual face of mandible near 

inferior margin below MI or M2 ; medium to very large 

size; early Oligocene to late Miocene, North America; 

middle Oligocene to early Pliocene, Europe. "Sorex" 

kinkelini Zinndorf (1901) appears to be from middle 

Oligocene (Eupelian) deposits. The specimen is a 

lower incisor that clearly represents Trimylus, but the 

species is not definable. As noted by Zinndorf (p. 134), 

it is closest to Trimylus neumayria/rms.

Genus DOMNINA Cope, 1873 

Genotype: Domnina gradata Cope, 18Y3 

P4 triangular in occlusal view with sharp-cornered 

triangular cusp; MI has very prominent entoconid distinctly separated from hypolophid and united to metaconid by high ridge (entoconid crest); teeth not bulbous ; cingulum strong but not inflated; mental foramen 

below middle of MI and associated only with minor 

depression on labial face of mandible. Dental formula 

1-5-3/1-5-3 or 1-5-3/1-4-3.
8 SUBFAMILIES AND GENERA OF THE SORICIDAE

Domnina is the oldest and most primitive shrew here 

recognized. In form of P4 , height of crown of M^ 

relatively slight reduction of the heel of M$, little separation of the mandibular condyles, lack of a deep and 

pocketed internal temporal fossa, lack of a posterior 

emargination on P4 and the upper molars, pigmentation 

of the teeth, and other features related to the structure 

of the skull (see McDowell, 1958, p. 182-184, and Patterson and McGrew, 1937, p. 255), Domnina could easily 

be considered ancestral to most other shrews. Domnina 

is precocious, however, in the posterior position of the 

mental foramen and in the extreme lingual placement of 

that part of the mandibular condyle homologous to the 

lower condyle of other shrews. It is atypical, in comparison with genera of the other subfamilies, in its transversely broadened or double-rooted premolars. These 

last features clearly indicate that Domnina is a specialized shrew belonging in the Heterosoricinae and not 

a generalized ancestor to all shrews.

Domnina thompsoni Simpson, 1941

Figure 2

A species of Domnina with lower antemolars not 

crowded anteroposteriorly; five (?) antemolars in addition to soricid incisor; roots of antemolars not transversely broadened; somewhat smaller size than D. 

gradata. Early Oligocene (Chadronian) of Montana.

This is the only known specimen of Domnina on which 

the mandibular condyles are well preserved. They are 

drawn apart but not separated (see McDowell, 1958, 

fig. SOD, E). It is interesting to note, even in this 

early stage of evolution of the soricid doubled condyle, 

that the lower anticular facet is well displaced lingually 

in the heterosoricine fashion and to a far greater extent

than in any other subfamily. The presumed reduction 

in number of antemolars in D. gradata, their conspicuous crowding between the molars and the insicor, and 

the transversely broadened or doubled roots of these 

antemolars are marked differences between these two 

species.

Specimen examined. The type, American Museum of 

Natural History 32647.

Domnina gradata Cope, 1873 

Figure 3

A species of Domnina with dental formula 1-5-3/ 

1-4-3; lower antemolars crowded together, four in number, with transversely broadened or incipiently doubled 

roots; somewhat larger size than D. thompsoni; heavily 

pigmented teeth nearly all of MI above cingulum is 

pigmented. Middle Oligocene (Orellan) of Colorado 

and Nebraska and Oligocene of Wyoming.

This is a very well-known shrew (see Patterson and 

McGrew, 1937). The first postincisor antemolar of the 

lower dentition is not known. The similarity in alveolar 

pattern between this species and Trimylus from the late 

Miocene of Oregon suggests that the P4 of D. gradata 

may not be larger than the first postincisor antemolar. 

The second and third lower antemolars are of a form 

similar to, but much smaller than, P4 .

The P4 is typically soricid but differs in certain 

characters. The protocone, typically small in most 

shrews, is either not present or is represented by a small 

bladelike elevation on the hypoconal cingulum. The 

paracone is more prominent and of a more conical form 

than in most shrews. The parastyle is minute and 

hardly more than a cingular irregularity. Finally, 

there is no emargination of the posterior basal outline

FIGURE 2. Domnina thompsoni Simpson, type, AMNH 32647. Left mandible. Medial view from Simpson (1941); posterior view from McDowell 

(1958).
DIAGNOSES AND CONTENTS OF SUBFAMILIES 9

although the basin between the parametaconal blade 

and the hypoconal cingulum is very deep. The P4 is 

known only on USNM 12841.

The M1 is typically soricid except that it lacks the 

posterior emargination and the protruding hypoconal 

flange which results from this emargination.

Consistent with its slightly younger geologic age, the 

mental foramen is slightly posterior of the position in 

D. thompsoni and is a little behind the midpoint of MI 

rather than a little behind the anterior root of J/i.

On the basis of USNM 12841 from Wyoming, 

McDowell (1958, fig. SOB, C) concluded that Domnina 

gradata lacked the zygomatic arch as do most shrews. 

Further preparation of the damaged skull of this specimen leads me to feel not only that the lack of an arch 

is not demonstrable on this specimen but that the size 

and shape of the broken end of the maxillary root of the 

zygoma suggests that a relatively large and horizontally 

broad arch or zygomatic process was present. This 

conclusion is almost mandatory when one considers the 

very large, deep, and extensive masseteric fossa on the 

mandible. It should be noted, also, that McDowell's 

illustration (fig. 30C) shows a slight posterior emargination on P4 and M1 . This emargination is not present 

on the specimen, but its absence could not be established 

until some matrix and a thick coating of glue were 

removed from the teeth.

Specimens examined. Chicago Natural History Museum P25750, P15320, P25754, and P25749; U.S. National Museum 12841.

Domnina greeni Macdonald, 1963

According to Macdonald (1963, p. 168), the ridge 

(the entoconid crest) connecting the entoconid to the

metaconid is lower than in D. gradata, and the labial 

cingulum is not present below the protoconid (although 

his fig. 5A seems to show that it is). His measurements 

indicate a shrew the size of D. gradata. Early Miocene 

(Arikareean) of South Dakota. 

This specimen was not seen.

Domnina n. sp.

Figure 4

Material. U.S. National Museum 23095, isolated left 

P± and MI with posterior half of the talonid missing. 

These two teeth were found in situ within 3 inches of 

each other and have the same degree of wear; it is 

assumed that they belong to the same individual. Collected by C. A. Repenning in 1959.

Locality and age. U.S. Geological Survey vertebrate 

loc. M1033, eastern Caliente Range, San Luis Obispo 

County, Calif. Early Miocene. Found in association 

with ArcJiaeolagus and ParaMppus in beds which contain, in the same area, mammals believed to belong in 

the Arikareean North American Provincial Age 

(Repenning and Vedder, 1961).

Diagnosis. A small species of Domnina with P4 that 

has transversely broadened root; size one-fourth smaller 

than D. gradata; pigment reduced to only tips of cusps.

Description. USNM 23095 is a shrew differing from 

Domnina gradata, in the parts represented, only by its 

size and the amount of pigmentation of its teeth. P4 is 

a triangular tooth with a single triangular cusp and a 

posterior cingulum which extends forward on both sides 

nearly to the anterior apex of the tooth. The cingulum 

does not overhang the root and would not have hidden 

either the root or the alveolus. The root of this speci-

.Entoconid

Entoconid crest

Occlusal-lateral view to show 

antemolar alveoli

1-5-3

1-4-3

FIGURE 3. Domnina gradata Cope, USNM 12841. Left mandibles; P 4 and 

M1 . M 3 is from CNHM P15320, and lower antemolar alveoli are from 

CNHM UC 1552.
10 SUBFAMILIES AND GENERA OF THE SORICIDAE

men was lost, but on the basis of the area of attachment 

P4 was either double rooted or at least transversely 

widened and anteroposteriorly compressed to about half 

of the width. Only a faint trace of yellowish pigment 

can be seen near the cusp apex. Maximum width of P4 

is 0.8 mm, and the anteroposterior length is about 

1.0 mm. There is a cavity formed by caries on the 

cingulum near the posterolingual corner of the tooth. 

The posterior half of the talonid is missing from the 

MI, and the tooth is well worn. The cingulum is strong 

from the paraconid to its termination at the fracture 

across the talonid. The entoconid is prominent and is 

connected to the metaconid by a crest that appears to 

be relatively as high as that in D. gradata. The metalophid joins the protolophid slightly labial of the midline 

of the tooth. The reentrant between the protoconid 

and the hypoconid opens on the labial side of the tooth 

at the cingulum and actually blends with the cingulum 

as in D. gradata. A faint trace of yellowish pigment is 

detectable on the labial face of the paralophid. The 

MI is about 1.5 mm in length (including an estimation of 

the missing part of the talonid) and about 1.0 mm wide 

across the protolophid. D. n. sp. (USNM 23095) is 

about as much smaller than D. thompsoni as D. thomp 

soni is smaller than D. gradata,:

D. gradata* 

Mi D. n. sp. D. thompsoni* (mean]

Length...________________ 1. 5± 1.8 2.3

Width_____-_-_____---___- 1.0 1.2 1.5 

*Data from Simpson (1941, p. 2).

The closer approach in size of D. n. sp. to D. thompsoni would suggest a closer relationship. The anteroposteriorly compressed root of the P4 of D. n. sp., however, and the relative age of the three species indicates 

a closer relationship between D. n. sp. and D. gradata. 

Except for its alveolus, P4 is not known in D. thompsoni.

Genus PARADOMNINA Hutchison, 1966 

Genotype: Paradomnina r&lictw Hutchison, 1966 

A heterosoricine shrew resembling Domnina but lacking a prominent entoconid crest joining the entoconid 

to the metaconid on MI and J/2 ; postentoconid valley 

lacking on M\ and M? so that hypolophid unites with 

entoconid; five postincisor antemolars including P4 in

the mandible; antemolar roots compressed anteroposteriorly as in most heterosoricines; cheek teeth 

elongate and not bulbous, mandible elongate in contrast 

to Trimylus. Late Miocene (Barstovian) of Oregon. 

Only one species, the genotype, is known.

Specimens examined. The type and referred specimens listed by Hutchison (1966, p. 5).

Genus TRIMYLTTS Roger, 1885 2

Genotype: Trimylus schlosseri Roger, 1885 = 

Sorex sansaniensis Lartet, 1851

P4 triangular in occlusal view (lacking other details 

because of bulbous form), smaller than first antemolar 

behind incisor (but first antemolar known only in late 

Miocene and early Pliocene forms); MI very low 

crowned, swollen, with inflated cingulum, these characters increasing in prominence in geologically younger 

forms; entoconid present on M\ in earlier forms but 

more detached from metaconid than in Domnina and 

closely attached to hypolophid, decreasing in importance and becoming a distal part of hypolophid in 

geologically youngest forms; talonid of M3 with crescentic loph and not distinctly bicuspid as in Domni/na; 

mental foramen below middle of MI to middle of M2 

(more posterior than in any other shrew) and placed 

in a depression on labial face of mandible which leads 

anterodorsally to premolar region; horizontal ramus 

massive and increasingly so in geologically younger 

forms; mandibular condyles widely separated in heterosoricine pattern (lower condyle displaced more lingually than ventrally); pigmentation reduced and in 

most forms only on incisors and anterior antemolars. 

Dental formula 1-5-3/1-4^3 or 1-^3/1-3-3.

There are at least three species of Trimylus in Europe 

and at least four in North America. The relationship 

between these is obscure but suggests two lineages. One 

is confined to North America, and the other to Europe. 

If two lineages are represented, they are not readily 

recognizable because of parallel, but not identical, evolutionary trends on each continent. The presence of two

2 In a recent publication, Doben-Florin (1964, p. 15-16) traced the 

history of the names Heterosorex Gaillard (1915) and Trimylus Roger 

(1885) and unquestionably established the priority of Trimylus. See 

synonomy under Trimylus sansaniensis.

Anterior Lingual 

FIGURE 4. Domnina n. sp., USNM 23095. Right P ^ and MI.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 11

lineages is suggested by the position of the conspicuous 

mental foramen which is below the trigonid of M2 in all 

European forms for which illustrations have been seen. 

In North American forms the foramen is below the 

talonid of M± except in Trimylus roperi in which it is 

below the trigonid of M^ as in the European forms. 

Progressive changes throughout the genus include increase in massiveness of the mandible, increase in 

bulbousness of the molars, increase in fusion of the 

entoconid to the posterior loph of the talonid of Mi and 

M2 with ultimate loss of this cusp as a distinct feature 

in geologically youngest forms, and decrease in the space 

for the lower antemolars between the incisor and M^ 

which results in reduction in size of the posterior antemolars and (in European forms) loss of an antemolar 

in both upper and lower jaws. The suggested relationships cannot be further evaluated without study of 

the European specimens.

Trimylus compressus (Galbreath), 1953 

Figure 5

Dommna compressa Galbreath, 1953, Kansas Univ. Paleont.

Contr., no. 13, Vertebrata, art. 4, p. 46-49, fig. 14. 

Heterosorex compressus (Galbreath). Mawby, 1960, Jour.

Paleontology, v. 34, no. 5, p. 954-956, fig. 2.

A little specialized species of Trimylus with molars 

distinctly more bulbous and cingula more inflated than 

in Domnina but less so than in other North American 

species of Trimylus; entoconid prominent on M^ connected to posterior lophid (or hypolophid) of talonid 

and separated from metaconid; lower antemolars not 

greatly crowded between incisor and MI although more 

so than in D. gradata; mental foramen below talonid 

of MI. Middle Oligocene (Orellan) of Colorado and 

middle or late Oligocene of Nebraska.

Only two specimens are known, both right mandibles. 

The type, University of Kansas 8154, has been described 

by Galbreath (1953, p. 46-49) and by Mawby (1960, p. 

954-955, text fig. 2). The referred specimen is Chicago

Natural History Museum UM400 and was collected 

north of Pine Kidge, Sioux County, Nebr. Neither 

specimen has the antemolars, but the type shows the 

alveoli of the posterior two and part of a third (Mawby, 

p. 955) from which an estimate of the crowding between 

the incisor and MI can be made. The Nebraska specimen shows a very light orange pigmentation on the 

upper half of the paraconid blade of M^ and, thus, 

indicates greater tooth pigmentation than is present in 

younger species. T. compressus has been compared to 

T. aff. neumayrianus (described by Viret and Zapfe, 

1951, p. 425-426) by Mawby (1960, p. 955). Perhaps 

the most significant comparison that can be made 

between the two is the slight crowding between the 

incisor and MI relative to later species on both 

continents.

Specimen examined. Chicago Natural History Museum UM400.

Trimylus aff. neumayrianus (Schlosser), 1887

Heterosorex aff. neumayrianus (Schlosser). Viret and Zapfe, 

1951, Eclogae geol. Helvetiae, v. 44, no. 2, p. 425-426, 

fig. 12.

Viret and Zapfe described a mandible of Trimylus 

from upper Oligocene (upper Stampian) rocks of 

Kickenbach, Switzerland. Their illustration shows 

three antemolar alveoli that are not at all crowded 

between the incisor and Mlf The posteriormost alveolus 

is long anteroposteriorly and, as suggested by Wilson 

(1960, p. 31), may have held the roots of two antemolars, 

which would bring the antemolar count to four. 

Domnina gradata and all North American species of 

Trimylus in which the premolar count is reasonably 

well established have four antemolars behind the incisor. 

Insofar as I am aware, however, this is the only 

Trimylus of Europe that may have had four antemolars. 

Wilson's suggestion that there were roots of two antemolars in the last alveolus of T. aff. neumayrianus is 

supported by the strong anteroposterior compression of

FIGURE 5. Trimylus compressus (Galbreath), CNHM UM400. Right mandible

and dentition.
12 SUBFAMILIES AND GENERA OF THE SORICIDAE

the roots of all heterosoricines except D. thompsoni 

that the elongate alveolus should have held just one root 

seems contradictory to a very well established character 

of the subfamily.

Trimylus aff. neumayrianus is also characterized by a 

very posterior mental foramen, a characteristic of all 

European species of the genus but not of any American 

species except T. roperi. In size T. aff. neumayrianus is 

comparable to T. compressus but it has a stouter horizontal ramus. Viret and Zapfe's illustration seems to 

show low teeth on the cutting edge of the lower incisor, 

a characteristic of earlier European species.

This specimen was not seen, and the only illustration 

and description seen are those of Viret and Zapfe.

Trimylus neumayrianus (Schlosser), 1887

Sorex neumayrianus Schlosser, 1887, Beitr. Palaontologie Geologie Osterreich-Ungarns, v. 6, p. 122, pi. 2, figs. 46-50.

Heterosorex neumayrianus (Schlosser). Gaillard, 1915, Soc. 

linne'ene Lyon Annales, v. 62, p. 83.

Heterosorex neumayrianus (Schlosser). Viret and Zapfe, 

1951, Eclogae geol. Helvetiae, v. 44, no. 2, p. 425.

Viret and Zapfe have compared their Stampian 

Trimylus with Schlosser's description of his Aquitanian 

species as well as possible from the published record. 

The lower antemolars, as well as their alveoli, according 

to Schlosser (1887, p. 122), are unknown. To judge 

from his figure 60 of plate 2, however, the proximal 

termination of enamel on the labial side of the lower 

incisor seems to be very nearly below the anterior edge 

of MI, a fact indicating greater anteroposterior compression of the antemolars than in Trimylus aff. 

neumayrianus from Rickenbach. The incisor to MI 

relationship seems to be very close to that in the 

Arikareean specimen from South Dakota, and there is 

less anteroposterior compression of the antemolars than 

in the late Arikareean Trimylus roperi from Colorado. 

The incisor bears two prominent cusps along its cutting

The masseteric fossa (Schlosser, 1887, plate 2, fig. 50) 

seems to be larger, deeper, and more extensive than in 

any other shrew, including the Arikareean and older 

species from North America, but the fossa is not preserved in the earlier European specimens. The mental 

foramen is below the trigonid of M2 as in all Old World 

forms. Early Miocene (Burdigalian) from Germany 

(Weisenau and Wintershof-West) and possibly middle 

dligooene (Rupelian) as "/Sotrex" kinkelini Zindorf, 

1901. No specimen seen.

Trimylus dakotensis Repenning, n. sp. 

Figure 6

Heterosorex. Macdonald, 1951, Soc. Vertebrate Paleontology

[Guide Book], 5th Field Conf., p. 72. 

Heterosorex. Mawby, 1960, Jour. Paleontology, v. 34, no. 5,

p. 951, 952, 955, 956.

Holotype. University of California Museum of 

Paleontology 37270; a nearly complete left mandible 

lacking the angular process and the antemolars. Collected by R. A. Stirton, 1934.

Type locality and age. University of California Museum of Paleontology loc. V 3417, Flint Hill fauna, 

Bennett County, S. Dak. Early Miocene (Arikareean).

Diagnosis. A small species of Trimylus with mental 

foramen below talonid of M\\ MI has entoconid flush 

against but not confluent with hypolophid, labial 

cingulum heavy and inflated but absent anterior to 

protoconid; M3 has inflated cingulum and tooth unreduced in contrast to late Miocene species; all cheek 

teeth moderately bulbous and low crowned.

Description. T. dakotensis is comparable in size to 

T. aff. neumayrianus and T. compressus; it is smaller 

than T. maiobyi, n. sp., and slightly smaller than the 

smallest specimens of T. roperi described by Wilson 

(1960, p. 32). Although the antemolars and that part 

of the mandible containing their alveoli are destroyed, 

the distance from the incisor to MI would indicate conFIGURE 6. Trimylus dakotensis Repenning, n. sp. Left mandible and dentition.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 13

siderably greater anteroposterior compression of the 

antemolars than is evident in either T. aff. neumayrianus or T. compressus. T, dakotensis differs from T. 

compressus in greater inflation of the labial cingula and 

in greater bulbousness of the cheek teeth. These same 

characters are further accentuated in the late Miocene 

species to be described in the following pages. T. dakotensis differs only slightly, however, from the nearly 

contemporaneous T. roperi. The chief differences 

appear to be the somewhat smaller size, the absence of a 

labial cingulum anterior of the protoconid of M\, 

slightly less massive mandible below the molars, and, 

perhaps most importantly, the anterior position of the 

mental foramen in T. dakotensis. Measurements of the 

type specimen have been given by Mawby (1960, table 

1, column e).

The coronoid and articular processes of the mandible 

of T. dakotensis are very well preserved. In typical 

heterosoricine fashion the lower articular facet is offset 

lingually so that no part of it lies beneath the upper 

facet. Although similar in pattern to Domnina thompsoni, the articular facets are separated and this pattern 

matches all known species of Trimylus for which the 

articular structure is known.

The type of Trimylus dakotensis appears to be the 

first specimen of this genus found in North America 

(there is no record of the date of discovery of CNHM 

UM400). Certainly it was the first Trimylus to be 

recognized from North America and was described in 

manuscript many years ago by Joseph T. Gregory. 

Unfortunately, the manuscript was never published and 

thanks are here given to Dr. Gregory for permission to 

describe the specimen in this report.

Trimylus roperi (Wilson), 1960

Heterosorex roperi Wilson, 1960, Kansas Univ. Paleont. Contr., 

Vertebrata, art. 7, p. 27-33, figs. 14-22.

A large species of Trimylus with dental formula 

1-5-3/1-4-3; four antemolars closely appressed between 

the lower incisor and MI ; entoconid of MI distinct in 

lingual view but completely fused to hypolophid in 

occlusal view; MI lower crowned than Oligocene and 

early Miocene forms and higher crowned than late 

Miocene forms from North America; mental foramen 

below or slightly behind trigonid of M2 in North 

America a character unique to this form; P4 and M1 

very similar to Domnina. gradata but more massive; 

size large, comparable to T. sansaniensis and T. mawbyi, 

n. sp. (see measurements in Mawby, 1960, and Wilson, 

1960). Late early Miocene (late Arikareean) of 

Colorado.

This is the earliest Trimylus in which the upper teeth 

are known (Wilson, 1963). They are also known from

the late Miocene T. sansaniensis and T. mawbyi, n. sp. 

All are strikingly like Domnina, and all have bifid upper 

incisors similar to the living genus Neomys.

These specimens were not seen but are well illustrated 

and described by Wilson.

Trimylus mawbyi Repenning, n. sp.

Figure 7

Heterosorex sp. Mawby, 1960, Jour. Paleontology, v. 34, no. 5, 

p. 950-956, text fig. 1.

Type. University of Oregon 10486, right mandible 

with incisor, antemolars 1 and 4, M^z, and part of the 

ascending ramus. Collected by F. B. Van Houten in 

about 1955.

Type locality and age. Five miles southwest of the 

south end of Guano Lake and about 24 miles southwest 

of Beatty Butte, Lake County, Oreg. The exact locality 

is not known to me; however, there are many outcrops 

in the general area that could represent the locality, and 

all are of the same strata. The referred material comes 

from these strata at localities from within 10 miles to 

about 40 miles from the type locality.

Late Miocene (Barstovian). The associated fauna is 

virtually identical to that from the Mascall Formation.

Referred material. U.S. National Museum 23096, a 

left mandible with incisor, alveoli for four antemolars, 

and MI and M2 , U.S. National Museum 23097, a right 

M1 ; both from U.S. Geological Survey loc. M1041, 

Beatty Butte fauna. Also five lower incisors, three 

upper incisors, and a mandible fragment with M\ from 

the following localities: Beatty Butte (loc. M1041), 

Harney County; Guano Eanch (loc. M1042), Lake 

County, 15 miles southwest of Beatty Butte; and Snyder 

Creek (loc. M1043), Lake County, which is 37 miles 

west-northwest of Beatty Butte. All referred specimens were associated with Barstovian mammals.

Diagnosis. Of all known species of Trimylus, T. 

mawbyi has the greatest reduction of Ms relative to MI, 

the greatest bulbousness of MI, the greatest inflation of 

the cingulum of M!, the lowest cusp height of MI, and 

the greatest massiveness of the mandible relative to the 

molar size. The metalophid joins the protolophid of 

M\ far more labially than in any other species. Also 

conspicuous is the fact that the stout molars are farther 

to the rear, relative to the anterior edge of the ascending 

ramus, than in any other heterosoricine.

Description. In addition to the above characters, 

T. mawbyi has the mental foramen below the talonid of 

MI, a position typical of most American members of this 

subfamily. The entoconid of MI is a blunt cuspid 

thoroughly merged with the hypolophid. The upper 

incisor is bifid. The M1 is quite comparable to T. roperi 

only more stout and bulbous. The antemolar alveoli of

240-305 O-67 3
14 SUBFAMILIES AND GENERA OF THE SORICIDAE

UO 10486

1-4-3

Antero-occlusal view to show antemolar 

alveoli USNM 23096

FIGURE 7. Trimylus mawbyi Repenning, n. sp., type UO 10486 and referred specimens. Type: right mandible and dentition. USNM 23096: left mandible 

and dentition. USNM 23097: right Ml.

the lower jaw, USNM 23096, show clearly that four 

transversely widened or doubled roots were present 

much as in Domnina gradata only more closely compressed between the incisor and MI. The alveoli of the 

first and last antemolars are nearly the same size and 

much larger than the intervening two. The type specimen, however, shows that the crown of P4 was much 

smaller than that of the first antemolar. These alveoli 

are nearly identical to those shown by Wilson for T. 

roperi (1960, fig. 20), which he feels (p. 31) are indicative of four antemolars behind the incisor. Measurements of the type have been given by Mawby (1960, 

table 1, column d).

The general mandibular and dental structure of T. 

mawbyi immediately gives the impression of a nutcracker type of action rather than the usual soricid 

condition which could be compared to the weak but 

quick action of chopsticks. Perhaps the heterosoricine 

mandibular characters in general reflect a molluscan, 

crustacean, or thick elytrum coleopterous diet. In 

addition, many of the mandibular characters are very 

similar to the living Chinese Mole- or BurrowingShrew, Anourosorex; thus fossorial habits are implied.

The type of T. mawbyi was described in detail by 

Mawby (1960) with excellent illustrations. He felt 

(p. 954) that a new species was represented but preferred not to name it until other specimens were 

known. Now that these are known, the species is 

named in his honor.

Specimens examined. The type and referred specimens mentioned above, which include all known 

specimens.

Trimylus sansaniensis (Lartet), 1851 

Sorex sansaniensis Lartet, 1851, Annuaire dept. Gers, Auch,

p. 13. 

Trimylus schlosseri Roger, 1885, Naturw., Ver. Schwaben u.

Neuburg Ber., v. 28, p. 106-107. 

Heterosorex delphinensis Gaillard, 1915, Soc. linne"ene Lyon

Annales, v. 62, p. 14. 

Trimylus schlosseri Roger. Stromer, 1928, Bayer. Akad.

Wiss. Abh., Math.-naturw. Abt, v. 32, no. 1, p. 13 ( =

S. sansaniensis and H. delphinensis). 

Heterosorex sansaniensis (Lartet). Viret and Zapfe, 1951,

Eclogae geol. Helvetiae, v. 44, no. 2, p. 419. 

Trimylus sansaniensis (Lartet). Doben-Florin, 1964, Bayer.

Akad. Wiss. Abh., Neue Folge, v. 117, p. 15.

A species of Trimylus with postincisor antemolars 

reduced to three in the lower jaw, of which the most 

anterior is largest (Crusafont Pairo, 1957), and 

tightly compressed between incisor and M^ entoconid 

of MI completely fused to hypolophid; mental foramen below trigonid of M^ size comparable to T. 

roperi; molars not so bulbous and not placed so far 

posteriorly relative to ascending ramus as in T. 

mawbyi; four upper postincisor antemolars (including 

P4 ) are clearly indicated in figure 11 of Viret and 

Zapfe (1951, p. 423). The lower incisor does not have 

subsidiary cusps along its cutting edge (Zapfe, 1951, 

p. 464, fig. 7). Later Miocene (Vindobonian) and 

early Pliocene (early Pontian) of Europe.

No specimens were seen. The Vindobonian specimens from La Grive St. Alban are well described 

and illustrated by Viret and Zapfe (1951, p. 419^24, 

figs. 10, 11), and those from the Vindobonian of 

Valles-Penedes by Crusafont Pairo (1957).
DIAGNOSES AND CONTENTS OF SUBFAMILIES 15

Genus INGENTISOREX Hutchison, 1966

Genotype: Ingentisorex tumididens Hutchison, 1966 

A heterosoricine shrew with bulbous dentition like 

TrimylMS except Mj- missing; P4 very large and larger 

than first postincisor antemolar; lower incisor short, 

not stout at base, one very prominent cusp on its 

cutting edge; labial cingulum of MI and M2 strong 

but not greatly swollen; trigonid of MI and M2 noticeably wider than talonid; mandible slender and not 

massive. Late Miocene (Barstovian) of Oregon. 

Only one species, the genotype, is known.

Specimens examined. The type and referred specimens listed by Hutchison (1966, p. 9).

Subfamily CROCIDUKINAE Milne-Edwards, 1868-1874

P4 has triangular cusp and a sulcus down posterior 

face of cusp, slight or no overhang of labial cingulum 

over root and alveolus, one or two roots; pigmentation 

absent; dental formula 1-6-3/1-4-3 to 1-6-3/1-3-3 in 

late Miocene and 1-3-3/1-2-3 to 1-5-3/1-3-3 in modern forms; articular facets of mandibular condyle 

separated but not widely, articulation surfaces usually 

united either along lingual side of the condyle with 

a labial- interarticular depression or across most of the 

interarticular surface; internal temporal fossa of 

mandible present; mental foramen below P4 in 

Miocene forms and below anterior root of MI in Pliocene to Recent forms; very small to very large size 

(the smallest and largest living shrews are included). 

Middle Miocene to Recent, Europe and Africa; 

early (?) Pliocene to Recent, Asia.

Although the Crocidurinae resemble Domnina in the 

structure of the P4 and are closely paralleled in other 

features by the North American Limnoecinae, there 

are no specimens outside of the Old World that appear 

to represent this subfamily. In many features this 

subfamily is a primitive group, and it is somewhat 

surprising, therefore, to find that the earliest shrews 

of the Old World are only soricine and heterosoricine; 

the earliest crocidurine shrews are from the Burdigalian of Europe (Doben-Florin, 1964, "Sorese" 

pusilliformis, and Soricetta discrepcms] and Africa 

(Butler and Hop wood, 1957, Crocidura sp.). The 

living shrews of Africa, Peninsular India, and Ceylon, 

all belong in the Crocidurinae (including Scutisorex], 

and possibly their earlier record is to be found in this 

area.

Many Old World fossil species of shrews are here 

included in the Crocidurinae and conform to the above 

diagnosis. The two most distinctive crocidurine characters exhibited by all these are the triangular P4 

(shared only with the Heterosoricinae) and the lingual 

union of the mandibular articular facets, with a labial

interarticular depression (shared only with the Limnoecinae). The first species discussed are those included in noncrocidurine genera in previous publications but whose subfamily affinities appear to be corcidurine. These are not reassigned to crocidurine 

genera, however, because the present review has been 

detailed enough to establish subfamily rank only, and 

all available specimens should be examined before 

generic reassignment is made. The remainder of both 

fossil and living species are discussed under their appropriate genera, in alphabetical order.

"Sorex" pusilliformis Doben-Florin, 1964

"Small shrew, tooth cusps unpigmented * * * M2 

about as large as M-L. Incisor toothed [cutting edge 

of lower incisor scalloped], first scallop located far 

from the tooth base., Two antemolars in the lower 

jaw, the posterior one [P4 ] two-rooted, its crown bearing two small secondary cuspids. MI and M2 with 

hypoconulid [entoconid separated from hypolophid], 

talonid of M3 one- to two-cusped * * * Mental foramen 

below the second antemolar [<P4 ]. Condyle with unseparated, Crocidura-like articular surface." (DobenFlorin, 1964, p. 32, part of diagnosis). P4 has a triangular cusp with well-developed posterolabial and 

posterolingual crests and "concave posterior face"; 

small accessory cuspulids are present on each of the 

posterior crests (Doben-Florin, p. 33). The articular 

structure of the mandible is distinctly crocidurine, as 

stated in the original diagnosis and as shown in 

Doben-Florin's illustrations (pi. 4, fig. Ib). The 

species clearly is not /Sorex.

The dental formula is l-?-3/l-2-3, and the species 

differs from Miosorex grwensis also in smaller size 

and cusp height (see Doben-Florin, 1964, p. 41). This 

appears to be one of the oldest crocidurine shrews 

known from Europe and is from the older Burdigalian 

locality of Wintershof-West. Doben-Florin reviewed 

the problem of the identity of Sorex pusttlus von 

Meyer which seems, at present, to be irresolvable. At 

least some of the Aquitanian shrews assigned to this 

species have pigmented teeth and appear to be assignable to the soricine shrew Crocidosorex cmtiquus 

(Pomel) ; the others are not assignable to subfamily 

on the basis of published descriptions.

"Sorex" dehmi Viret and Zapfe, 1951 

Dental formula 1-6-3/1-4-3; P4 has two roots longitudinally placed in mandible, both posteroexternal 

and posterointernal crests of triangular cusp have subsidiary cuspules along them much as in the living 

Sylvisorex, posterolabial cingulum of P4 slightly overhanging posterior root; MI has moderate cusp height,
16 SUBFAMILIES AND GENERA OF THE SORICIDAE

talonid wider than trigonid so that metalophid units 

with protolophid (protoconid-metaconid crest) very 

near to protoconid; M3 has remnant of basin remaining in talonid as in Miosorex grivensis and not trenchant as in modern crocidurines; mental foramen below 

P4 ', dental foramen usually far forward; P* and M* 

have slight emargination of the posterior border quite 

comparable to Miosorex grivensis, according to Mein 

(1958, figs. 20, 21). Late Miocene (Vindobonian) or 

possibly somewhat earlies (late Burdigalian) (Mein, 

1958, p. 106) of Europe.

No specimens were seen. They are, however, well 

figured and described by Viret and Zapfe (1951, p. 

415-417) with a description of the upper dentition by 

Mein (1958, p. 25-26). This species is as distinct 

from other known fossil and living shrews as the 

living genera are distinct from each other. It should 

be placed in a new crocidurine genus or with Miosorex 

grivensis as a separate species. This should not be 

done without study of the large sample now available 

(Mein stated that he has 100 mandibles and 14 maxillaries from Vieux-Collonges). "Sorex" dehmi has an 

interesting combination of features, some of which 

seem more progressive than M. grivensis and most of 

which seem more primitive than its contemporary.

Genus CROCIDURA Wagler, 1832

Figure 8

Genotype: Sorex leucodon Hermann, 1780

Dental formula 1-4-3/1-2-3; lower incisor short to

moderate length and curving upward, cutting edge

slightly serrate to smooth, groove along length of

medial side curves downward below notch in basal

border; first upper antemolar larger than equal-sized 

second and third antemolars, all unicuspid. Late Miocene ( ?) to Recent, Africa; early Pleistocene to Recent, 

Europe; Pleistocene to Recent, Asia.

During this review no characters were noted which 

would serve to separate Crocidura from Suncus on 

the basis of lower dentition or structure of the mandible. Prof. P. M. Butler (written commun. 1964) 

suggested that there appears to be taxonomic value 

in the "cingulumlike groove running along the length 

of the medial side of the first lower incisor. In all 

Suncus except S. lixus, this continues above the notch 

in the basal border of the tooth as a posterolingual 

cingulum. This does not happen in Crocidura." The 

additional upper antemolar in Suncus is, of course, 

diagnostic. Ellerman and Morrison-Scott (1951, p. 

65) stated that in their opinion Suncus is no more than 

a subgenus of Cnocidura. Throughout the Soricidae, 

dental formula is a significant character of generic 

stature, and the opinion held in this review is that 

Crocidura and Suncus must be retained as distinct 

genera on this basis or else an immense amount of 

synonomy is in order throughout all living shrews.

It is not so easy to defend the generic distinctiveness 

of Crocidura and Praesorex by proclaiming custom to 

be guiding dogma of soricid taxonomy. In this comparison the above diagnosis is not diagnostic. Crocidura differs from Praesorex in smaller size (which 

is variable), in having an inflated pterygoid region 

on its skull, in having great interorbital skull breadth 

relative to the rostral breadth at the infraorbital 

foramina, and in having a basined fossa for the inMedial

FIGURE 8. Crocidura russula Hermann, Belgium, author's collection 60186. 

Left mandible and dentition and left upper dentition.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 17

sertion of the superior pterygoid muscle on the mandible (all of which can easily be shown to vary with 

size). There are two characters which may be constant differences between the genera but which have 

not been checked for consistency in this review nor, 

as far as I am aware, in any study because of the 

lack of an adequate sample of Praesorex. These are 

the more anterior position of the mental foramen and 

the shorter, more velvety, fur of Crocidura. In Crocidura the mental foramen lies below the P± to below 

the protoconid of Mlf In the two available specimens 

of Praesorex, it lies below the hypoconid of Mlm According to Meester (1953, p. 213), who had only two 

specimens of Praesorex but a large series of Crocidura, 

the difference in pelage is diagnostic. For this review, 

therefore, Praesorex is retained as a separate genus, 

and the above characters are considered diagnostic of 

Crocidura.

Crocidura sp., Butler and Hopwood, 1957 

This specimen (Butler and Hopwood, 1957, p. 2) 

seems too incomplete to defend the generic assignment. 

Late Miocene, Kenya.

Crocidura wongi Pel, 1936

More primitive than any living crocidurine in that 

the talonid of M3 is not reduced to a single trenchant 

cusp although it is practically so, according to Pei 

(1936, p. 22). No specimen was seen. It is well 

illustrated by Pei (figs. 10,11). Pleistocene of China.

Crocidura taungsensis Broom, 1948

A small species lacking the anterior cusp present on 

the P4 of living species, according to Meester (1955, 

p. 274-275). Early Pleistocene, South Africa.

Crocidura kornfeldi Kormos, 1934

More delicate than living C. suaveolens, according 

to Friant (1947, p. 264). No specimen was seen. 

Early Pleistocene of Europe.

Crocidura obtusa Kretzoi, 1938

Elongate first lower antemolar and "blunt" P4 with 

dimensions comparable to living C. leucodon, according to Kretzoi (1938, p. 92). No specimen was seen. 

Pleistocene of Europe.

Crocidura zorzii Pasa, 1948

Citation from Sulimski (1962, p. 484). No specimen seen. Middle Pleistocene of Italy.

Genus DIPLOMESODON Brandt, 1853 

Figure 9

Genotype: Sorex pulchelbus Lichtenstein, 1823 

Dental formula 1-3-3/1-2-3; first upper antemolar 

larger than second; M2 trapezoidal; lower incisor 

without or with single accessory cuspule on cutting 

edge; P4 slightly larger than preceding antemolar, 

crocidurine posterolingual crest very weak; posterolabial crest virtually lacking; Mv has longitudinally 

shortened talonid, entoconid present and either comMedial

1-3-3 

1-2-3

(JAWS)

FIGURE 9. Diplomesodon pulchellum (Lichtenstein), Kazakhstan, U.S.S.R., 

AMNH 206558. Left mandible and dentition and left upper dentition.
18 SUBFAMILIES AND GENERA OF THE SORICIDAE

pletely fused with hypolophid or very close to this 

lophid, entoconid crest very low, labial cingulum continuous from paraconid to behind hypoconid but not 

inflated or enlarged; MI and P± distinctly anteroposteriorly compressed to adjust to short snout (in upper 

dentition this adjustment is made by loss of antemolars 

rather than by marked anteroposterior compression of 

teeth); trigonid of M3 slightly reduced but talonid 

greatly reduced to single cusp with entoconid crest 

leading directly to metaconid or entirely lost; zygomatic process of maxillary originates just behind parastyle of M2 to opposite metastyle of M-^; mental foramen below protoconid of M± to below anterior tip of 

Mi and posterolabial corner of P±\ ascending ramus 

high, vertical and narrow; internal temporal fossa 

triangular and rather small; mandibular condylar 

process and articular facets somewhat more widely 

separated than in Crocidura to extreme separation for 

subfamily, articular surfaces continuous (or nearly 

so) along lingual margin of interarticular area. 

Kecent of south-central U.S.S.R. (Aral Sea area) and 

early Pleistocene of South Africa.

Specimen examined. American Museum of Natural 

History 206558, a male Diplomesodon pulchellum from 

west Kazakstan, U.S.S.E.

Diplomesodon fossorius Repenning, 1965 

A large species with greatly reduced M-^- and with 

extreme specialization of the mandibular articulation, 

paralleling that of many soricine shrews. Type and 

referred specimens listed by Repenning were seen. 

Early Pleistocene, South Africa.

Genus FEROCULUS Kelaart, 1852

Genotype: Sorex feroculus Kelaart, 1852 

Dental formula 1-5-3/1-2-3; lower incisor with two 

lobes on cutting edges as in Sylvisorex but inclined 

forward rather than rounded; foreclaws enlarged. 

Specimens of this shrew are very scarce. Through 

the kindness of R. W. Hayman I have been able to 

examine photographs of the specimen in the British 

Museum (Nat. History) (BM 28.10.27.6). The similarity to Sylvisorex is striking. From the photographs I am unable to see any differences that would 

appear significant except possibly one less cusp and 

the forward inclination of the cusps on the cutting 

edge of the lower first incisor of Ferocukis. The 

posterolingual emargination of the P4 and M1 is 

broader and deeper in Feroculus. No specimen was 

seen. Recent, Ceylon.

Genus MIOSOREX Kretzoi, 1959 

Figure 10

Genotype: Sorex pusillus var. grivensis Deperet, 1892 

Dental formula 1-6-3/1-3-3; P± has slight overhang of labial cingulum over root and secondary 

cuspule along posterolabial crest; MI with moderate 

cusp height reentrant between the protoconid and 

hypoconid emerges on labial face a short distance 

above cingulum; talonid of MB not as reduced as in 

modern crocidurines but retains slight basin enclosed 

by high labial and very low lingual ridge; P* with 

only slight emargination of posterior border; M1 with 

moderate emargination; mental foramen below root 

of /Y Late Miocene (Vindobonian) of Europe.

Hypolophid Protolophid

Medial

(JAWS)

FIGURE 10. Miosorex grivensis (Deperet), UC 24859 with P4 and incisor 

restored from Viret and Zapfe (1951). Right mandible and dentition.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 19

The slightly reduced and basined talonid of M3, the 

slight emargination of P4 and M1, the retention of 

six upper antemolars, and possibly the lesser height 

of crown of M^ are all that separate this crocidurine 

shrew from related living genera.

Miosorex grivensis (Deperet), 1892

Sorex pusiltus, var. grivensis Deperet, 1892, Mus. d'histoire

nat. Lyon Archives, v. 5, mem. 2, p. 48. 

Sorex (s.l.) grivensis (Deperet). Viret and Zapfe, 1951,

Eclogae geol. Helvetiae, v. 44, no. 2, p. 411-412, figs. 1-3. 

Miosorex grivensis (Deperet). Kretzoi, 1959, Vertebrata hungarica, v. 1, no. 2, p. 248, footnote 1.

This is the only species assigned to the genus.

Specimens examined. University of California 

Museum of Paleontology 42672, a right mandible with 

M2 and Mz, and University of Colorado 24859, a right 

mandible with J/i to Mz from La Grive St. Alban. 

More complete material is well figured and described 

by Viret and Zapfe.

Genus MYOSOREX Gray, 1838 

Figure 11

Genotype: Sorex varius Smuts, 1832 

Dental formula 1-5-3/1-3-3; lower incisor short 

and curving upward, cutting edge smooth or has two 

low cuspulids; first upper antemolar larger than following ones except P4, second very small, third nearly 

as large as first, and fourth minute (lost in Myosorex 

polli Balsac and LaMotte, 1956, p. 155-157), all bicusped with cusps forming parallel longitudinal 

blades; posterior lobe of falciform upper incisor also

bilobed so that two rows of lobate cusps run from 

nipping part of incisor back to P4 ; P4 with accessory 

cuspule on both posterolingual and posterolabial 

crests; P% minute and on the lingual side between the 

two major lower antemolars; early Pleistocene to 

Recent, Africa.

Ellerinan, Morrison-Scott, and Hayman (1953, p. 

19) stated that the Ps is so vestigial as to be very 

rarely present and so considered Myosorex a subgenus 

of Suncus. Eleven specimens of M. varius from Cape 

Province, South Africa (three localities), are in the 

Museum of Vertebrate Zoology, University of California, Berkeley. Of these 11 specimens, 9 have the 

vestigial P3 on both mandibles, 1 (UCMVZ 117080, 

a male) has it on one mandible only, and 1 (UCMVZ 

117085, a very old male) has the Ps missing on both 

mandibles although the upper antemolars are as described above. I am inclined to feel that the tendency 

to have three postincisor lower antemolars is very 

strong in this species and that this tendency, combined with the unique structure of the upper antemolars, is of generic importance and comparable in 

magnitude to differences found between other genera.

Specimens examined. Myosorex warms, 11 specimens in the University of California Museum of 

Vertebrate Zoology. Also, early and late Pleistocene 

specimens of Myosorex in the University of Colorado 

Museum from Cave of Hearths and Limeworks, 

Makapansgat, Transvaal, South Africa.

Medial

(JAWS)

FIGURE 11. Myosorex varius (Smuts), Cape Province, Union of South Africa, 

UCMVZ 17086. Right mandible and dentition and left upper dentition.
20 SUBFAMILIES AND GENERA OF THE SORICIDAE

Myosorex robinsoni Meester, 1955

According to Meester (1955, p. 272-273), this species 

lacks subsidiary cuspules on the posterolabial and 

posterolingual crests of P±. Several specimens were 

examined. Early Pleistocene, South Africa.

Genus PARACEOCIDURA Balsac, 1956

Genotype: Paracrocidura schoutedeni Balsac, 1956 

Dental formula 1-4-3/1-2-3; upper incisor with 

expansion of anterointernal border producing chisellike cutting edge rather than point as in most shrews; 

P4, If1, and M2 with squared posterior borders lacking any emargination; first lower postincisor antemolar very large and longitudinally bicuspid. Kecent, 

Africa.

Relative to soricine shrews, most crocidurine shrews 

tend to have enlarged and elongate first lower postincisor antemolars. The elongation of this tooth in 

Paracrocidura exceeds any shrew examined during 

this review, however, and greatly exceeds the most 

closely comparable genera, Sylvisorex and Feroculus. 

The structure of P4 , M1 , and M2 is unique for the 

subfamily. The development of medial crests on the 

tip of the upper incisors is also unique for the subfamily and appears to be comparable to the fissident 

incisor found in the Heterosoricinae and in most Old 

World genera of the tribe Neomyini (new name; see 

p. 45). The structure of the P4 , which has both posterolabial and posterolingual crests (with prominent 

secondary cuspules), indicates that Paracrocidura is

a crocidurine shrew. The structure of the mandibular 

condyle has not been described. No specimen was 

seen.

Genus PRAESOREX Thomas, 1913 

Figure 12

Genotype: Orocidura goliath Thomas, 1906 

Dental formula 1-4-3/1-2-3; largest known shrew; 

teeth as in Orocidura; pterygoid region of skull not 

inflated; interorbital breadth not great relative to 

rostral breadth at infraorbital foramina; mandibular 

insertion of superior pterygoid muscle not basined and 

pterygoid spicule developed; mental foramen below 

hypoconid of J/i; fur long, coarse, with prominent 

guard hairs. Recent of equatorial Afria.

This ermine- to weasel-sized genus is the largest 

known shrew, living or extinct. It is not, however, 

greatly larger than the giant species Sunous caeruleus. 

Except for size it is hardly more than a species of 

Crocidura, and some species assigned to Orocidktra are 

nearly as large. With the exception of the position 

of the mental foramen and the nature of the pelage, 

all characters listed in the above diagnosis are known 

to vary with size in other shrews. Its similarity to 

Orocidura is discussed under that genus. Balsac and 

LaMotte (1957, p. 29) consider Praesorex goliath, 

the only species included in this genus, to be a species 

of Orocidura.

Specimens examined. Harvard University Museum 

of Comparative Zoology 14718, a female Praesorex

FIGURE 12. Praesorex goliath (Thomas), Cameroons, MCZ 14718. Left mandible 

and dentition and right upper dentition.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 21

goliath from Cameroons; British Museum (Nat. 

History) BM. 11.5.5.7, a male Praesorex goliath from 

Cameroons.

Genus SCUTISOREX Thomas, 1913

Genotype: Sylvisorex somereni Thomas, 1910 

Dental formula 1-5-3/1-2-3; first upper antemolar 

slightly larger than subequal following three; lower 

incisor short, curving upward at tip, with smooth 

upper margin; in dorsal aspect articular facets of 

mandibular condyle much closer to being normal to 

longitudinal axis of the mandible than in other crocidurines; angular process of mandible enlarged, stout, 

and attenuated posteriorly in comparison with other 

crocidurines; vertebral column with remarkable specialization (Alien, 1917). No specimen was seen. It 

is well illustrated by Alien. Kecent, Africa.

Genus SOLISOREX Thomas, 1924

Genotype: Soli&orex pearsoni Thomas, 1924 

Dental formula 1-4-3/1-2-3; lower incisor with 

smooth cutting edge as in Suncus; foreclaws enlarged. 

The more significant parts of the description of Solisorex given by Thomas (1924, p. 94) are as follows: 

"Articulation with lower jaws unusual in various 

aspects. Most noticeable is the great size and transverse extent of the mandibular condyles, each of these 

being broader than the space which separates them 

from each other across the middle line * * * Teeth 

large and heavy * * * Upper 'unicuspids' only three 

in number, of which the anterior one is almost unique 

in the family by being slightly bicuspid, in size the 

first is much the largest, the second about one-sixth 

its bulk, and the third rather more than twice the 

size of the second. First two molars more square 

than in Pachyura [Suncus], * * * with less salient 

posterointernal lobes * * * Lower molars also simplified, without accessory basal lobes or ledges, the 

posterior (M$) remarkably reduced, consisting only 

of a smooth lunate cuspless crest corresponding to the 

tuberculo-sectorial trigonid of the other shrews, the 

unusual heel completely aborted."

Solisorex is distinct from all known shrews. It is 

unique among the crocidurines in the loss of the labial 

cingulum on the lower teeth; the reduction of M3 and 

the specialization of the mandibular articulation is 

matched only by the extinct Diplomesodon f&ssorius; 

the dental formula and other features of the dentition 

are similar to Praesorex.

Through the kindness of R. W. Hayman, I have 

been able to examine photographs of a specimen in 

the British Museum (Nat. History), BM.31.8.14.4. 

No specimen was seen. Recent, Ceylon.

Genus SORICELLA Doben-Florin, 1964

Genotype: Soricella discrepant Doben-Florin, 1964

The following quotations are from Doben-Florin 

(1964, p. 48):

Diagnosis. "Middle-sized soricid, teeth appear unpigmented."

Lower dentition. "Doubly-scalloped incisor, scallops 

inclined forward, the first [most posterior] near the 

tooth base. Three single-cusped antemolars, the last 

[most posterior or P4 ] two rooted, the middle tooth 

smaller. Molars decreasing in size from front to back. 

MI and Mz without hypoconulid [entoconid fused to the 

hypolophid], M% rather large with double-cusped 

talonid. Labial cingulum strong."

Upper dentition. "Incisor with posterior accessory 

cusp, four antemolars, all well visible; the second 

smaller than the rest. Between the last [most posterior] 

and P4 lies a small diastem. Ml and M2 are concavely 

emarginate on their posterior side; their trigonbasins 

closed posteriorly for the most part by a flat enamel 

border."

Condyle. "Rather small with wide, almost straight, 

undivided articular surface."

From Doben-Florin's illustration of the posterior 

view of the mandibular condyle (pi. 5, fig. 5: the condyle 

of specimen 11702, shown on table 34, p. 53, to be a left 

mandible), the articular structure is distinctly crocidurine and has prominent lingual confluence of the 

two articular surfaces. Her figures of the occlusal view 

of P4 are not clear enough for me to be certain that they 

have the crocidurine triangular cusp. The high dental 

formula, 1-5-3/1-3-3, the uniquely sharp cusps on the 

cutting edge of the lower incisor, and the tall, narrow 

coronoid process of the mandible, which is very reminiscent of Diplomesodon, appear to be the most distinctive characters of this genus.

One species, Soricella discrepans Doben-Florin, has 

been described. It is from the older Burdigalian of 

Wintershof-West. No specimen was seen.

Genus SUNCUS Ehrenberg, 1832 

Figure 13

Genotype: Suncus saoer Ehrenberg, 1832 

Dental formula 1-5-3/1-2-3; lower incisor short to 

moderate length and curving upward, cutting edge 

only very slightly serrate at most; groove along length 

of medial side curves above notch in basal border and 

forms posterolingual cingulum; first upper antemolar 

larger than subequal second and third, fourth antemolar 

much smaller, all unicuspid. The genus contains the 

smallest and the next to largest living shrews. Early 

Pleistocene to Recent, Africa; Recent, Europe and Asia.

240-305 O-67 4
22 SUBFAMILIES AND GENERA OF THE SORICIDAE

Posterior

Medial

(JAWS)

FIGURE 13. Suncus caeruleus Kerr, Ceylon UCMVZ 81267. Left mandible 

and dentition and left upper dentition.

Ellerman and Morrison-Scott (1951, p. 65) stated 

that in their opinion this genus is no more than a subgenus of Grocidura.

Specimens examined. A small series including 

Suncus caeruleus and S. et-ruscus in the University of 

California Museum of Vertebrate Zoology.

Suncus sp., Meester, 1955

The genus has been reported from several South 

African localities. Several specimens have been seen. 

Early Pleistocene, South Africa.

Genus SURDISOREX Thomas, 1906 

Figure 14

Genotype: Surdisorex norae Thomas, 1906 

Dental formula 1-4^3/1-2-3; upper antemolars as in 

Crocidura except P* distinctly more elongate (also seen 

in Myosorex blarina zinki Balsac and LaMotte, 1956, 

fig. 2); lower antemolars elongate; first lower antemolar with two-cusped crest; P4 two rooted with accessory cuspule on both posterolabial and posterolingual 

crests, posterolingual crest incomplete behind accessory 

cuspule, posterolabial crest normal for subfamily to 

accessory cuspule but behind this cuspule crest swings 

lingually and merges with posterior cingulum at midpoint of tooth or even as far lingually as posterolingual 

corner of tooth; metalophid-protolophid junction of J/i 

placed remarkably close to labial side of tooth; other 

dental characters as in Grocidura; ascending ramus very 

broad; articular condyle of mandible of crocidurine 

pattern but unique in closeness of upper and lower 

articular surfaces and basically a single, triangular,

posteriorly convex articular surface with labial emargination. Recent, Africa.

Although the structure of P± of this shrew closely 

parallels that of the soricine shrews, other features, 

particularly the mandibular condyle, clearly indicate 

that it is a crocidurine shrew. The condyle itself is 

also interesting in that it more closely approaches the 

normal mammalian single condyle than does that of any 

other living shrew examined in this review, although 

in Myosorex the articular surfaces are only slightly 

separated.

Specimens examined. U.S. National Museum 182593, 

a male of /S. norae, from Kenya; American Museum of 

Natural History 187262, a female from Kenya.

Genus SYLYISOBEX Thomas, 1905 

Figure 15

Genotype: Crocidura morio Gray, 1862 

Dental formula 1-5-3/1-2-3; lower incisor long and 

projecting straight anteriorly (except S. megalura, see 

Heim de Balsac and LaMotte, 1957, fig. 10c), cutting 

edge serrate with three prominent notches separating 

rounded cuspate lobes comparable to those of many 

species of Sorex; first upper antemolar behind incisor 

larger than equal-sized second, third, and fourth, all 

unicuspid; P4 with accessory cuspule on the posterolingual crest only. Recent, Africa.

Ellerman, Morrison-Scott, and Hayman (1953, p. 19) 

treated Sylvisorex as a synonym of Myosorex* and 

Myosorex as a subgenus of Suncus. The construction of 

the lower incisor alone would indicate that
DIAGNOSES AND CONTENTS OF SUBFAMILIES 23

FIGURE 14 Surdisorex norae Thomas, Kenya USNM 182593. Left mandible 

and dentition and left upper dentition.

FIGURE 15. Sylvisorex lunaris Thomas, Uganda, CNHM 26260. Left mandible 

and dentition and left upper dentition.

is distinct and possibly closer to Feroculus than to 

Suncus.

Specimen examined. Chicago Natural History Museum 26260, Sylvisorex lunaris, from Uganda.

Crocidurinae ? Incertae Sedis

"limnoecus" micromorphus Doben-Florin, 1964

"Very small shrew, teeth probably pigmented, similar

to L. niobrarensis [on which no pigment is visible] * * *

M3 with a single-cusped talonid. M± and M2 with a

hypoconulid [hypolophid separated from entoconid]

and very strongly developed entoconid. In the J/i the

metaconid stands slightly behind the protoconid. All 

three molars with a delicate, but distinct labial 

cingulum * * * Two antemolars, the posterior one two 

rooted. Mental foramen below the trigonid of MI, 

condyle with undivided articular surfaces, Crociduralike. Mandibular foramen behind the middle of the 

internal temporal fossa." (Doben-Florin, 1964, p. 44). 

In her description of the holotype, Doben-Florin 

further stated (p. 45) that a "pronounced, somewhat 

concave chewing surface, on the three-cornered outline 

[of P±\ slopes from the tooth cusp [apex]." On page 

48, she makes the following comparison: "The chewing
24 SUBFAMILIES AND GENERA OF THE SORICIDAE

surface of the two antemolars are stretched out in L. 

niobrarensis and run to a median, small, posterior cusp. 

In the Wintershof form, however, the chewing surface 

is three cornered, somewhat concavely excavated, the 

elevated posterior margin allows the tooth to appear 

two cusped from the labial side; there is, liowever, no 

corner behind the cusp as formed for example, in the 

Recent 8orex alpinus" From this description, as well 

as from what can be seen from the single illustration 

of the occlusal surface of P4 (pi. 4, fig. 9a), the P4 of 

"Z." micromorpJius appears to be crocidurine rather 

than limnoecine.

Although difficult to judge from figure 9a of plate 4, 

the distance between the protoconid and metaconid of 

MI appears relatively much greater than in any known 

limnoecine shrew.

Doben-Florin's statement about possible pigmentation and the posterior position of the mental foramen 

suggest that the species is not crocidurine, however. It 

seems, therefore, that no conclusive subfamily assignment can be made from the published description. The 

species is here questionably referred to the Crocidurinae 

because of the form of the P4 . The objections to this 

referral may be reduced by noting that Doben-Florin 

was not certain that the teeth were pigmented.

"Z." micromorphu-s is from the older Burdigalian 

fauna of Wintershof-West. No specimen was seen.

"Sorex" collongensis Mein, 1958

To judge from Mein's figure 22, this shrew has a 

crocidurine P4 . Its teeth have no pigmentation, and the 

mental foramen is below P4 , which also suggest the 

crocidurine shrews, in late Miocene or more recent time 

at least. Mein (1958, p. 26) stated that the species also 

has a mandibular articulation that is enormous for the 

size of the mandible but does not figure the articulation 

or further describe it. His description of the internal 

temporal fossa as "semicircular" suggests the Soricinae. 

Based upon the criteria of the present review, this 

shrew is only questionably placed in the Crocidurinae. 

According to Mein, it has an anteriorly placed dental 

foramen comparable to "$." dehmi and four lower 

antemolars. No specimen was seen. Late middle or 

late Miocene of France.

Crocidura?, Mein, 1958

Only two maxillary fragments are reported (Mein 

1958, p. 27). According to Mein, they resemble "/Sorex" 

dehmi from the same fauna but have fewer antemolars. 

Based upon the criteria considered in the present report, 

these specimens are not assignable to a subfamily. No 

specimens were seen. Late middle to early late Miocene, 

France.

Three living genera which are customarily placed in 

the Crocidurinae are not included in this subfamily in 

this review. The main criteria used to remove them 

from the Crocidurinae and to place them in the 

Soricinae are structure of P4 and structure of the 

mandibular condyles. In addition, some have pigmented teeth, at least in some individuals. In one 

genus, pigmentation is only at the very tips of the cusps 

and is removed in early wear; in another, it is of a very 

light color so that it is sometimes unnoticed. One genus 

is unpigmented. In the past few decades many people 

have noticed the fact that some crocidurines have pigmented teeth, and there have been two distinct reactions: One is that the subfamily criteria break down 

and hence the subfamily classification is not real; the 

other is that the shrews in question have pigmented 

teeth and are not crocidurine by definition.

The first reaction is typified by Ellerman and 

Morrison-Scott (1951, p. 41) who made note of the 

inconsistencies in pigmentation and abandoned subfamily classification without further investigation. The 

second reaction is perhaps best typified by Bate (1945, 

p. 761), who removed those shrews with pigmented 

teeth from the Crocidurinae but also cited several other 

skull and skeletal differences that suggest closer relationships to the Soricinae.

Genera which have been included in the Crocidurinae 

(for example, by Simpson, 1945, p. 51-52) but are 

included in the Soricinae in this review are: 

Anourosorex; Chimarrogale; Nectogale.

In addition, North American fossil shrews generally 

assigned to the genus Limnoecus are removed from the 

Crocidurinae and placed in a new subfamily, the 

Limnoecinae, on the basis of the structure of P4 , pigmentation, structure of Ms, and position of the mental 

foramen; the mandibular articulation of the Limnoecinae appears identical to that of the Crocidurinae.

Subfamily LIMNOECINAE Repenning, new name 

P4 with little or no lingual segment or posterolingual 

ridge of primitive triangular cusp, labial segment of 

primitive cusp and its posterolabial crest form a 

medially shifted longitudinal ridge, labial shearing 

blade (as in Soricinae) not developed, posterolingual 

basin (as in Soricinae) little or not developed, single 

root, slight overhang by labial cingulum over root and 

alveolus but never as extreme as in most soricines; MI 

with protoconid and metaconid as close or closer 

together in unworn teeth than in any other known 

shrew, entoconid strong, entoconid crest low; MS lacking 

double-cusped talonid in earliest form and an appreciable talonid basin lost through emphasis of metalophid 

by late Miocene time, becoming very nearly a simple 

trenchant heel like that in modern crocidurine shrews
DIAGNOSES AND CONTENTS OF SUBFAMILIES 25

and more trenchant than in late Miocene crocidurines; 

upper teeth unknown except for incisor and Ma - pigmented heavily in earliest form but only at tips of cusps 

in later forms; lower dental formula 1-3-3 to 1-2 ?-3; 

articular facets of mandibular condyle as in Crocidurinae, internal temporal fossa present; mental foramen below trigonid of MI (more posterior than in most 

crocidurine shrews of comparable age). Early Miocene 

(Arikareean) to middle Pliocene (Hemphillian),3 

North America.

This is a small but nevertheless distinct subfamily. It 

is rather well known in the Miocene and early Pliocene 

faunas of western United States and is represented from 

faunas of both ages in California (Stirton, 1930; James, 

1963, p. 31-47), from the Miocene of Colorado (Wilson, 

1960, p. 33-35), Wyoming (Keed, 1960, p. 6-7), and 

Nebraska (Macdonald, 1947), and from the early 

Pliocene of Nevada (J. E. Mawby, oral commun., 1963) 

and middle Pliocene, Oregon.

Genus ANGUSTIDENS Repenning, n. gen.

Genotype: /Sorex vireti Wilson, 1960 

Diagnosis. P± triangular in occlusal view and 

basically crocidurine in pattern except strong posterolabial crest meets (but does not merge with) cingulum 

distinctly medial of posterolabial corner and posterolingual crest reduced and detached from apex of primary cusp, sulcus on posterior face of cusp between

3 A right mandible of Limnoecus sp. cf. L. tricuspis, bearing unworn Mi to Ms, was collected by the author from the Hemphillian 

fauna at Rome, Oreg. This represents the only known middle Pliocene record.

crests; MI with protoconid and metaconid distinctly 

close together, entoconid strong and detached from 

hypolophid but connected to metaconid by very low 

entoconid crest, reentrant between protoconid and 

hypoconid emerges on labial face of tooth just above 

cingulum, labial cingulum very subdued to absent below 

protoconid of MI but strong both anteriorly and 

posteriorly from this point; M3 with reduced talonid 

but still retaining minute basin and low entoconid 

crest; mental foramen below trigonid of MI ; pigmentation extensive, comparable to Sorex; lower dental formula 1-3-3 or possibly 1-4-3, in which case PS was 

vestigal and represented in type mandible by a minute 

alveolus on anterior margin of alveolus of P4 .

Etymology. Latin: Angustidens, narrow tooth; 

refers to the closeness of the protoconid to the metaconid.

Angustidens vireti (Wilson), 1960 

Figure 16

Sorex vireti Wilson, 1960, Kansas Univ. Paleont. Contr., Vertebrata, art. 7, p. 33-35 and 37-38. 

Limnoecus vireti (Wilson). James, 1963, California Univ.

pub. Geol. Sci., v. 45, p. 36.

A fairly large shrew slightly smaller than living 

Blarina brevicauda. Size, location of the mental foramen in a depression that leads anterodorsally to the 

antemolar region, location of the dental foramen about 

on midline of and well below internal temporal fossa, 

and weakness or absence of the labial cingulum of M\ 

below protoconid may prove to be of specific importance 

when other shrews are found that belong to this genus. 

Early Miocene (Arikareean) of northeastern Colorado.

KU 10037

Medial

FIGURE 16. Angustidens vireti (Wilson), type KU 10037 and referred specimens 

KU 10044 and 10304. Right mandible and lower dentition.
26 SUBFAMILIES AND GENERA OF THE SORICIDAE

James (1963, p. 39) placed this species in the genus 

Limnoecus. The relationship is clear but the differences 

between Limnoecus of the late Miocene and Pliocene 

and this species seems to be greater than comparable 

differences between living shrew genera, as between 

Sorex and Microsorex. Wilson's species, therefore, is 

placed in a new genus. In the form of P* and M$, 

Angustidens vireti is exactly intermediate between the 

condition believed to have been present in a hypothetical 

soricid progenitor (typified in these two teeth, at least, 

by Domnina) and Limnoecus.

Specimens examined. The type, University of 

Kansas Museum of Natural History 10037, and paratypes listed by Wilson (1960, p. 33).

Genus LIMNOECUS Stirton. 1930

Genotype: Limnoecus tricuspis Stirton, 1930 

P* with posterolabial crest of primitive triangular 

/*4 forming medial crest which hooks lingually where 

it merges with posterior cingulum, posterolingual crest 

nearly or entirely lost; reentrant valley between protoconid and hypoconid of MI emerges on labial face 

well above cingulum; talonid of J/3 reduced to 

trenchant heel formed by posteriorly directed metalophid and hypoconid, talonid basin virtually lost 

although faint trace of entoconid crest (not continuous with hypoconid) usually present on lingual edge 

of talonid in unworn teeth; pigment reduced to tips 

of cusps or at most to upper half of paraconid blade 

of Mi] lower dental formula 1-3-3 to 1-2 ?-3.

Limnoecus tricuspis Stirton, 1930 

Figure 17

Medium-sized shrew comparable in size to living 

/Sorex araneus; teeth more slender in occlusal view 

and more delicate than L. niobrarensis ; P* retaining 

slight remnant of posterolingual crest; Ms retaining 

slight talonid basin and entoconid crest with trenchant 

metalophid curving posterolingually to merge with 

hypoconid; lower dental formula 1-2?-3. Late Miocene (Barstovian), early Pliocene (Clarendonian), 

California, and middle Pliocene (Hemphillian), 

Oregon.

This is the genotypic species. In the type description Stirton (1930, p. 218) considered the lack of a 

metaconid on M3 to be a diagnostic feature of 

Limnoecus. James (1963, p. 31-47) reprepared and 

restudied the type and concluded that the metaconid 

had been broken off of the specimen. He also described 

new material from the early Pliocene. Known material is not adequate to rule out the possibility of a 

vestigial P^ being present.

Specimens examined. The type, University of 

California Museum of Paleontology 31047; topotypic 

material in the University of California at Riverside; 

and referred specimen, University of California Museum of Paleontology 55515, from Cuyama Valley, 

Calif. Uncataloged specimen from USGS vertebrate 

loc. M1078, Rome, Oreg.

Anterior view of 

M,\ moderate wear

Medial

FIGURE 17. Limnoecus tricuspis Stirton, type, UCMP 31047. Right mandible

and lower dentition.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 27

Limnoecus niobrarensis Macdonald, 1947

Figure 18

Medium-sized shrew with teeth broader in occlusal 

view, bearing stouter cusps, than those of L. tricuspis; 

PS minute, vestigial, and largely overlapped by P4 ; 

P4 nearly square in occlusal outline with no trace of 

posterolingual crest; Ms with metalophid directed 

posteriorly to hypolophid, very slight remnant of 

entoconid crest but no trace of talonid basin; mandible stouter than in L. tricuspis; lower dental formula 

1-3-3. Late Miocene (Barstovian), Nebraska.

The differences between L. tricuspis and L. niobrarensis are slight but are of the same sort and of same 

magnitude as are observable in the mandibles of living 

species of Sorex, for example, between Sorex cinereus 

and Sorex merriami. Based on this analogy with 

living species, these differences seem to be of specific 

value. Many of the specific characters given by 

Macdonald (1947, p. 124-125) were based upon features that James (1963, p. 32-33, tables 6, 7) pointed 

out resulted from damage of the type of L. tricuspis 

not known to Macdonald. James further concluded 

that the differences in tooth proportions between the 

two species could represent those of sex rather than 

species and synonymizes L. niobrarensis with L. tricuspis. The possibility that the differences are sexual 

rather than specific has not seemed great enough in 

the present review to abandon Macdonald's species, 

and it is here retained until a larger sample of both 

species can shed more light on their variation.

Specimens examined. The type and paratype, University of California Museum of Paleontology 36171 

and 36172.

Subfamily SOBJCINAE Fischer von Waldheim, 1817

P4 with no lingual segment of primitive triangular 

cusp, labial segment of primitive cusp and its posterolabial crest retained as labial shearing blade, in more 

advanced forms this blade is continuous with elevated 

posterior cingulum forming L-shaped crest enclosing 

posterolingual basin (which has evolved from posterior 

sulcus between crests of primitive triangular cusp of 

P4 ), posterolingual crest greatly reduced or absent 

(more usual in advanced forms), cingulum overhangs 

root and lateral surface of mandible below posterolabial corner of P4 in most genera more than in contemporary forms of other subfamilies, single rooted 

except in atypical Miocene form; molars variable; pigmentation usually present, sometimes extreme, but 

absent in some genera of one tribe; dental formula 

1-6-3/1-4-3 in early Miocene forms (higher than in 

Domnina] and 1-6-3/1-2-3 to 1-3-3/1-2-3 in modern 

forms (1-4-2/1-2-2 in Amblycoptus] ; mental foramen 

in many genera farther back than in contemporary 

crocidurines and limnoecines; articular facets of 

condyles little separated (or even joined in earlier 

forms) to extremely separated in most living genera, 

facets are joined along labial 'side of condyle, rather 

than along lingual side as in crocidurine-limnoecine 

pattern, with lingual emargination of interarticular 

area; internal temporal fossa present. Late Oligocene

Anterior view 

t (paratype) 

slight wear

Medial

FIGURE 18. Limnoecus niobrarensis Macdonald, type and MI of paratype, 

UCMP 36171 and 36172. Right mandible and lower dentition.
28 SUBFAMILIES AND GENERA OF THE SORICIDAE

(Stampian) to Recent, Europe; early Miocene (Arikareean) to Recent, North America; early(?) Pliocene 

to Recent, Asia; Recent, South America; not known 

from Africa or peninsular India. Romer (1928, p. 

100) stated that Sorex vulgaris (= Sorex araneus] 

now lives in Mediterranean Algeria but is not known 

in the late Pleistocene deposits of that area; however, 

this is the only publication I have found that mentions 

soricines from Africa.

A diagnosis of this subfamily is difficult to outline 

briefly because of great evolutionary change and 

because of remarkable differences in contemporaneous 

genera of the subfamily, particularly later in the history of the subfamily. Consideration of the subfamily 

as a whole and throughout its history leaves only two 

criteria which are entirely definitive: the structure of 

/*4 and of the articular condyle or condyles. In some 

genera even these are difficult to recognize but never 

has there been any question of subfamily recognition 

when more variable criteria have been used in support 

of the characters of P4 and of the condyles. It should 

be emphasized, however, that other characters vary 

with the stage of evolution and with separate lineages 

within the subfamily Soricinae.

Although the number of living species of crocidurine shrews about equals that of the soricine shrews, 

there are twenty-nine soricine genera recognized in 

this review and only ten crocidurine genera. To review this large subfamily with some system comparable in detail to that used for the other subfamilies, the 

Soricinae have been broken down into three tribes. 

These subdivisions are based on the structure of the 

mandibular articulation and associated musculature 

which appears to remain constant within the tribes 

despite the presence of genera otherwise very diverse 

in form. The subfamily is readily classified into two 

tribes on the basis of the mandibular articulation. A 

third tribe is set up for those few genera (but many 

species) that are of primitive structure in their articulation and cannot easily be assigned to either of the 

two more specialized tribes. In varying degrees other 

features support this tribal grouping.

Within the tribes further subdivision, of a subtribe 

magnitude, is suggested by features found to be exclusive in only part of a tribe, and these features seem 

to have geographic significance, at least on a continental scale. These subtribes are discussed and serve 

as a basis for the order of discussion of genera where 

such lesser subdivisions seem to be present. They are 

not sharply defined, however, and are not formally 

set up as classification units.

The modification of the mandibular articulation is 

developed to increase the strength of the bite. To do

this the lower articular facet, on which the mandible 

is hinged, is moved forward to shorten the lever arm 

of the horizontal ramus. The temporal muscle is also 

strengthened, and its area of origin is spread upward 

and forward over the cranium. Its greater and more 

transverse force increases the tendency for the posterior part of the mandible to be pulled into the side of 

the braincase when biting. Consequently, the upper 

articular facets, the single function of which is to 

counteract the transverse force of the temporal muscle, 

is changed in orientation in order to remain approximately normal to the average force resultant of the 

temporal muscle. As the upper articulation does not 

function as a hinge but rather slides back and forth 

over a strong articular boss on the cranium, this does 

not impair the hinge action of the mandible which is 

controlled by the position of the lower condylar facet.

Primitively the condyloid process supporting the 

upper articular facet is a thin plate of bone alined 

with the plane of the ascending and horizontal rami, 

as in Sorex, As the force of the temporal muscle increases, the plane of the condyloid process curves away 

from the plane of the horizontal ramus and toward the 

braincase in order to parallel the force resultant of 

the temporal muscle to attain greatest possible strength 

in resisting the tendency of this muscle to pull the 

mandible into the braincase. This is actually accomplished by an outward warping of only the ascending 

ramus rather than of the entire proximal part of the 

mandible.

The most readily recognized changes in the proximal part of the mandible resulting from a strengthening of the bite area are, therefore, separation of the 

articular facets by the forward placement of the lower 

facet and outward deflection of the ascending ramus 

to allow the inward curving of that part of the condyloid process that supports the upper facet.

As these changes take place, an emargination of the 

lingual side of the interarticular area develops and 

the insertion of the superior pterygoid muscle on the 

lingual side of the condyloid process, between the two 

articular condyles, becomes basined. This condition is 

found in Neomys and is the articular structure which 

characterized the tribe Neomyini (fig. 1). The action 

of the superior pterygoid muscle upon the mandible 

is directly forward, consequently the lingual condylar 

emargination weakens its insertion and a pterygoid 

spine or boss is developed to strengthen it in many 

forms. The spine is at the dorsal margin of the superior pterygoid fossa just below the upper sigmoid 

notch. The tribe Neomyini is also characterized by a 

bifid upper incisor and a prominent entoconid crest
DIAGNOSES AND CONTENTS OF SUBFAMILIES 29

on J/i of most forms, as well as several other dental 

and cranial features.

In other shrews the insertion of the superior pterygoid muscle is strengthened by the development of a 

bone plate across the lingual condylar emargination 

which also strengthens the condylar process supporting the upper condyle. Except in forms where the 

bone plate is weakly formed, no pterygoid spine is 

required or developed. In posterior view the intercondylar area appears to lack any emargination although the area filled by the bone plate behind the 

basined insertion of the superior pterygoid muscle is 

usually depressed from the rest of the interarticular 

area. This is the condition found in Blarina and is 

the articular structure that characterized the tribe 

Blarinini (fig. 1). The tribe Blarinini is also characterized by an uncleft upper incisor and by no entoconid crest on the M^ as well as several other dental 

and cranial features.

The articular structure is unmodified in the tribe 

Soricini (fig. 1). The articular facets are not far 

apart and the condylar process, the coronoid process, 

and the horizontal ramus all lie in the same plane. 

An entoconid crest is present on J/1? and a bifid upper 

incisor, generally a feature of a variant individual, 

may or may not be present.

Tribe SORICINI Fischer von Waldheim, 1817

Articular facets of mandibular condyle continuous 

or only slightly separated, both facets approximately 

normal to longitudinal axis of cranium; when facets 

are separate, interarticular area broad with no conspicuous lingual emargination seen in posterior view, 

lower facet not offset lingually beyond plane of mandible at lower sigmoid notch or shifted appreciably forward (fig. 1) ; superior pterygoid fossa of the mandible a shallow pit; external temporal fossa of mandible 

extends no lower on ascending ramus than upper sigmoid notch; internal temporal fossa of mandible large 

and triangular; coronoid process not deflected labially 

but in plane of horizontal ramus and of condyloid 

process; zygomatic process of maxillary originates to 

rear of M2 ; M2 rectangular in occlusal view; M^ with 

strong entoconid crest; Mz unreduced or with only 

talonid reduced, roots separate by distance equal to or 

greater than transverse diameter of anterior alveolus; 

teeth pigmented.

This tribe includes four living genera; one is Sorex, 

containing by far the largest number of species of any 

genus of shrew and the only truly holarctic genus now 

living. The tribe also contains the oldest fossil representatives of the subfamily Soricinae because, by 

definition, it has the least specialized mandibular articulation. Among living genera of this tribe, there 

are two distinct lineages. The first lineage, exemplified by the living Sorex, is primitive in nearly all 

respects in addition to the articular structure and has 

a known history dating from the late Oligocene of 

Europe and the early Miocene of North America. 

The second lineage, exemplified by the living Blarinella, is advanced in most respects except in the articular structure and has a known history dating from 

the middle Pliocene of eastern Asia, and from the late 

Pliocene of Europe.

Genus CROCIDOSOREX Lavocat, 1951

Genotype: Crocidosorex piveteaui Lavocat, 1951 

Dental formula 1-6-3/1-4-3, including a minute 

vestigial P3 ; P4 with soricine labial shearing blade 

which bears a secondary cuspule but (in contrast to 

Sorex) does not merge with posterior cingulum and 

does not enclose posterolingual basin, overhang by 

cingulum at posterolabial corner of tooth very slight 

and single root not covered; M\ with cingulum only 

along paraconid blade and with reentrant valley between protoconid and hypoconid emerging on labial 

face of tooth well above enamel base in marked contrast to contemporary Dormiina,; Ms as in /Sorex and 

Donwina with unreduced talonid enclosed lingually 

by entoconid crest; mental foramen below P4 ; mandibular condyle with facets only slightly more separated than in Domnina but lower facet placed more 

ventrally than in heterosoricines and not offset lingually beyond plane of mandible at lower sigmoid 

notch, condyle with soricine pattern of labial connection between upper and lower facets and lingual 

emargination between facets rather than opposite as 

in crocidurines and limnoecines (see pi. 2, fig. 73, in 

Schlosser, 1887, for illustration of condyle); internal 

temporal fossa deep and narrowly triangular; masseteric fossa absent; P4 and M1 with moderate emargination of posterior basal outline between hypoconal 

flange and metastyle.

Crocidosorex piveteaui Lavocat, 1951 

Medium-sized shrew comparable in size to living 

Sorex ara/wus; M\ with prominent entoconid connected to metaconid by relatively low entoconid crest; 

upper teeth unknown; teeth unpigmented according 

to Lavocat (1951, p. 24). No specimen was seen. Late 

Oligocene (upper Stampian) of France according to 

Rene Lavocat (oral commun., 1963).

Crocidosorex antiquus (Fomel), 1853 

Sorex antiquus Pomel, 1853, Annales sci., litt, Indus. [Auvergne], v. 26, p. 81-229. 

Sorex antiquus, Stehlin, 1940, Eclogae geol. Helvetiae, v. 33,

p. 298, figs. Ib, 2b, 3, 4b, and 5b.

240-305 O-67 5
30 SUBFAMILIES AND GENERA OF THE SORICIDAE

Small shrew, smaller than C. piveteaui and comparable in size to living Microsorex hoyi; MI with 

prominent entoconid connected to metaconid by welldeveloped entoconid crest; teeth pigmented; upper 

teeth and mandible as described for genus. No specimen seen. Excellent material is well illustrated and 

described by Stehlin (1940, p. 298-301). Early Miocene (Aquitanian) of France (Montaigu-le-Blin) and 

Germany (Weisenau).

O. piveteaui and C. antiquus are similar shrews, but 

the differences in pigmentation and in the entoconid 

crest seem to warrant their retention as separate species until such time as the specimens can be compared 

directly. Lavocat (1951, p. 23-24) makes no comparison between the two.

ll/Sorex pusillus von Meyer" of Tobien (1939, p. 165) 

may belong to this genus, but the published description 

is not adequate for certain assignment. Stehlin (1940, 

p. 299, footnote 4) stated his opinion that "Sorex" 

pusillus and "Sorex" a/tribiguus are probably about 

the same as "Sorex" antiquus. The same possibility 

is also expressed by Kretzoi (1959b, p. 247); however, 

Kretzoi also noted that "Sorex" pusillus von Meyer 

was preoccupied by Sorex pusillus Gmelin, and he 

replaced the name with a new genus and species (ibid., 

p. 247, 248, footnote 1) Oligosorex meyeri to apply to 

the shrew from Weisenau (without reference to other 

shrews he would include in the genus) Kretzoi's diagnosis for the genus Oligosorex is "late Oligocene and 

Miocene Soricids of small dimensions, retaining three 

unicuspids between P4 and the relatively small and 

primitive front incisor in the lower jaw"; this diagnosis basically conforms to that here used for Croddosorex. Study of the specimens, however, could 

well lead to the conclusion that the two genera are 

distinct.

Crocidosorex is structurally suitable as an ancestral 

form for all later soricine shrews. It. differs little 

from living Sorex. the differences being those cited in 

the generic diagnosis.

Genus ANTESOREX Repenning, n. gen.

Genotype: /Sorex compressus Wilson, 1960 

Diagnosis. P4 and mandibular condyle as in Soricinae. P4 with weak shearing blade that merges 

with posterior cingulum, shearing blade with slight 

secondary cuspule, posterolingual basin not developed, 

two roots arranged longitudinally, labial cingulum 

slightly overhangs anterior root and not posterior 

root; MI with reentrant valley between the protoconid 

and metaconid emerging on labial face of tooth well 

above oingulum (much higher than in the limnoecine 

shrew Angustidens vireti which was collected in the 

same deposits as the genotype of Antesorex} , welldeveloped entoconid connected to metaconid by a high 

entoconid crest, labial cingulum fairly strong along 

paraconid blade but very weak and difficult to see 

along remainder of labial side of tooth; Ms as in /Sorex 

with talonid basin completely enclosed by the entoconid crest; emargination of posterior basal outline of 

F4 about as in Crocidosorex but possibly slightly 

greater in M1 ; teeth pigmented; mental foramen below posterior root of F4 ; internal temporal fossa 

strong and narrowly triangular as in Crocidosorex',

KU 10051

P4 and Afj-KU 10050 

M-KU 10052

Medial

Posterior

FIGURE 19. Antesorex compressus (Wilson), type KU 10050 and referred 

specimens. Left mandible and lower dentition and referred left upper 

dentition.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 31

masseteric fossa absent; mandibular condyle with 

facets only slightly more separated than in Domnina 

but lower facet placed more ventrally than in heterosoricines, articular surfaces of facets continuous along 

labial side of condyle and not along lingual side of 

condyle as in limnoecines and crocidurines; number 

of antemolars unknown.

Etymology. Latin: Ante, before, and the genus 

Sorex.

Antesorex compressus (Wilson), 1960 

Figure 19

Sorex compressus Wilson, 1960, Kansas Univ. Paleont. Contr., 

Vertebrata, art. 7, p. 35-38, figs. 29-31.

Small shrew comparable in size to Microsorex hoyi; 

P± anteroposteriorly compressed and with constriction 

around crown that may have been deformity during 

growth; MI with protoconid remarkably close to the 

metaconid in unworn teeth. Late early Miocene (late 

Arikareean), northeastern Colorado.

The genus Antesorex from North Americans very 

close to Cr&cidosorex from Europe, and A. compressus 

is most like C. antiquus. Antesorex is slightly more 

advanced in the union of the labial shearing blade to 

the posterior cingulum of F4 , in the continuous (although weak) cingulum along the labial side of J/i, 

and in the somewhat greater emargination of the 

posterior basal outline of M1 . It seems less advanced 

in the retention of two roots on P4 , a feature that may 

be an atavism.

Specimens examined. The type, University of 

Kansas, Museum of Natural History 10050, and paratypes listed by Wilson (1960, p. 35).

Genus SOREX Linnaeus, 1758 

Figure 20

Genotype: Sorex araneus Linnaeus, 1758 

Dental formula 1-6-3/1-2-3; lower incisor usually 

with ventral margin having little dorsal curve toward tip (all Old World species appear to have three 

well-developed scalloped cusps on cutting edge of 

lower incisor, but there are exceptions to this in New 

World species) ; MI with entoconid crest connecting 

entoconid to metaconid; Ms as in Domnina and Crocidosorex with unreduced and basined heel; P4 and 

M1 with strong emargination of posterior basal outline; protocone of P4 not as lingual in position as 

hypoconal flange; teeth pigmented; articular facets of 

mandibular condyle slightly to moderately separated, 

lower facet extends labially to or beyond plane of 

mandible at lower sigmoid notch; interarticular area 

of condyle not appreciably constricted in posterior 

view in most species but extremely variable and approaching the pattern of "other tribes in some species; 

mental foramen usually below protoconid of MI or 

anterior of this point.

The genus is the most generalized and, in many 

respects, the most primitive of living soricine shrews. 

No species are known that are unquestionably of early 

Pliocene age, but there are a large number of fossil 

species included in the genus of later Pliocene and 

Pleistocene age. These have been briefly reviewed 

(mostly from published accounts) for conformity to 

the generic diagnosis but not for synonymy or for

Medial

FIGURE 20. Sorex araneus Linnaeus, Belgium, author's collection 60182. Right 

mandible and lower dentition and left upper dentition.
32 SUBFAMILIES AND GENERA OF THE SORICIDAE

specific diagnosis. The following list is of species

here included in the genus, some questionably: 

Pliocene and earliest Pleistocene species

Sorex dixonensis Hibbard, 1956. To judge from the published 

illustrations, the low-crowned molars, the wide lower mandibular condyle, the narrow interarticular area, and the 

very strong entoconid crest on M2 are exceptional for the 

genus. The generic and tribal assignment is here considered doubtful. No specimen was seen. Earliest Pleistocene 

(late Blancan), Kansas.

Sorex alpinoides Kowalski, 1956. Poland. No specimen was 

seen.

Sorex leahyi Hibbard, 1956. The lower condyle is more elongate lingually than in most species of Sorex. Generic and 

tribal assignment here considered in doubt. No specimen 

was seen. Earliest Pleistocene (late Blancan), Kansas.

Sorex mimitus Linnaeus, 1776, central Europe, late Pliocene 

(Sulimski, 1962, p. 461) to living. Recent specimens seen.

Sorex rexroadensis Hibbard, 1953. No specimen was seen. 

Late Pliocene (early Blancan), Kansas.

Sorex sandersi Hibbard, 1956. No specimen was seen. Earliest 

Pleistocene (late Blancan), Kansas.

Sorex subminutus Sulimski, 1962. No specimen was seen. 

Late Pliocene (pre-Villafranchian), Poland.

Sorex taylori Hibbard, 1937. No specimen was seen. Late 

Pliocene and middle Pleistocene (early Blancan and Irvingtonian), Kansas.

Early Pleistocene species

Sorex araneoides Heller, 1930. Germany. No specimen was 

seen.

Sorex cinereus Kerr, 1792. Kansas (Hibbard, 1944, p. 719- 

720). A living species. Only Recent specimens were examined.

Sorex cudahyensis Hibbard, 1944. Kansas. The type, University of Kansas Museum of Natural History 6513 was 

examined.

Sorex fallax Heller, 1936. Germany. No specimen was seen.

Sorex kennardi Hinton, 1911. (?) Germany (Heller, 1958, p. 

18-19) ; the type, from England, is late Pleistocene. No 

specimen was seen.

Sorex laoustris (Hibbard), 1955 [Neosorex lacustris Hibbard]. 

Kansas. No specimen was seen.

Sorex praealpinus Heller, 1930. Germany and Poland. No 

specimens were seen.

Sorex praearaneus Kormos, 1934. Hungary, Germany, and 

Poland. No specimen was seen.

Sorex runtonensis Hinton, 1911. Austria, Czechoslovakia, England, Germany, and Poland. Two specimens, author's collection 6235 A and C from Czechoslovakia were examined.

Sorex subaraneus Heller, 1959. Germany. No specimen was 

seen.

Species known only from the late Pleistocene

Sorex perminutiis Kretzoi, 1959b, equals Sorex minutissimua 

Heim de Balsac, 1940, France. No specimen seen.

Sorex frankstownensis Peterson, 1926. Pennsylvania. No 

specimen was seen.

Many living species have been reported from late 

Pleistocene deposits.

Genus DREPANOSOREX Kretzoi, 1941

Genotype: Sorex (Drepanosorex] tasnadii 

Kretzoi, 1941

Diagnosis as Sorex except that the upper soricid 

incisor is fissident in all known examples (Miklos 

Kretzoi, oral commun., 1966); lower molars stout with 

heavy cusps that are lower crowned than those of 

Sorex ; teeth with pale-orange-yellow pigment rather 

than dark red; mandibular condyles large with 

greater vertical separation than in Sorex, and dorsal 

condyle extended much farther back approximating

Posterior

FIGURE 21. Microsorex hoyi (Baird), Alaska, author's collection 6059. Left 

mandible and dentition and left upper dentition.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 33

the condition found in Blarinella, Middle Pleistocene of Europe. Four species have been named:

Drepanosorex margaritodon (Kormos), 1935. Middle Pleistocene, Hungary. No specimen seen.

Drepanosorex savini (Hinton), 1911. Middle Pleistocene, Austria, Czechoslovakia, and England. One specimen seen, author's collection 6234 from Czechoslovakia.

Drepanosorex tasnadii (Kretzoi), 1941. Middle Pleistocene, 

Hungary and Germany. No specimen seen.

Drepanosorex pachyodon (Pasa), 1948. Original designation 

Sorex pachyodon. Middle Pleistocene, Italy. No specimen 

seen.

Genus MICROSOREX Baird, 1877 

Figure 21

Genotype: Sorex hoyi Baird, 1858 

Dental formula as in Sorex 1-6-3/1-2-3, but third 

and fifth upper antemolars minute; relative to trigonid, talonid of Ms somewhat more reduced than in 

Sorex; because shortening of upper antemolar arcade 

is reflected in lower antemolars, F4 somewhat broader 

in occlusal view than in Sorex; in contrast to nearly 

all species of Sorex, F4 with curving J-shaped labial 

crest rather than an angulate L-shaped crest, slight 

posterolingual basin; MI as in Sorex; mental foramen rather far to rear and nearly beneath hypoconid 

of MI. Only Recent specimens examined. Two fossil species have been named:

Microsorex pratensis Hibbard, 1944. Middle Pleistocene (Irvingtonian), Kansas. 

Microsorex minutus Brown, 1908. Late Pleistocene (Rancholabrean), Arkansas.

The five genera just discussed are the known representatives of the unspecialized lineage of the tribe

Soricini. In addition to the unspecialized mandibular 

articulation, they have dental characters that do not 

differ greatly between the Oligocene forms and the 

living forms. Nor do the dental characters differ 

greatly from those found in the earliest members of 

other subfamilies, like the heterosoricine Domnina, 

the crocidurine Miosorex grivensis, and the limnoecine Angustidens, although these earliest members of 

the other subfamilies clearly show the beginnings of 

dental modification characteristic of their subfamilies. 

The remaining genera of the tribe Soricini, in contrast, have considerable dental modification from the 

primitive pattern although the mandibular articulation is still clearly primitive.

Genus ALLTTVISOREX Hutchison, 1966

Figure 22

Genotype: Alluvisorex arcadentes Hutchison, 1966 

A soricine shrew with primitive mandibular articulation comparable to that of Antesorex in outline but 

with small interarticular area separating upper and 

lower articular facets; dental formula ?/l-3-3 (in A. 

arcadentes} to 1-6-3/1-2-3 (in A. chasseae) ; cheek 

teeth stout and low cusped; mandible heavy; lower 

incisor with three low rounded cusps on cutting edge; 

F3 minute when present; MI with entoconid very close 

to metaconid and connected to it by a high and prominent entoconid crest, postentoconid valley very wide, 

metalophid joins protolophid close to protoconid, labial 

cingulum prominent but not inflated; occlusal outline 

of Ml and M2 distinctly rectangular; M3 with reduced 

talonid consisting of longitudinal metalophid terMental 

foramen

LACM 4265

Less worn topotypic 

specimen UCR 10032

Medial

FIGURE 22. Alluvisorex chasseae (Tedford), paratype, LACM 4265 and 

topotype UCR 10032. Right mandible and lower dentitions.
34 SUBFAMILIES AND GENERA OF THE SORICIDAE

minating in strong hypoconid, no entoconid, very low 

entoconid crest, more on the lingual side of the talonid 

than on the occlusal surface and missing on one 

specimen of A. chasseae from Nevada (UCR 10237); 

mental foramen below metalophid of M^ upper incisor not fissident; P4 and upper molars intermediate 

between Sorex and Blarinella. Late Miocene (Barstovian) of Oregon and early Pliocene (Clarendonian) 

of Nevada and California. 

Two species have been named:

Alluvisorex arcadentes Hutchison, from the late Miocene of 

southeastern Oregon. This species retains a minute P3 . 

The type and referred material listed by Hutchison (1966, 

p. 18) were seen.

Alluvisorex chasseae (Tedford), originally described from the 

lower Pliocene of California (Tedford, 1961) and subsequently described from the lower Pliocene of Nevada (Suthard, 1966). The para type, Los Angeles County Museum 

4265, and topotypes in the University of California, Riverside, as well as the specimens from Fish Lake Valley, Nev., 

in the same institution were seen.

Genus PETENYIA Kormos, 1934 

Figure 23

Genotype: Petenyia hungarica Kormos, 1934 

Dental formula 1-5-3/1-2-3; lower incisor with two 

low cuspules on its cutting edge; M± with prominent 

entoconid crest about as in living species of Blarinella, 

labial cingulum strong but not inflated and continuous 

to posterior margin of tooth behind hypoconid; 

talonid of M3 greatly reduced and no trace of entoconid crest; third upper antemolar reduced, fourth 

very reduced, fifth upper antemolar absent in Petenyia 

(present but minute in Blarinella quadraticauda) ; P4 

and M1 with slight emargination of posterior basal 

outline; hypoconal crest of M1 and M2 united to protocone to form a continuous endoloph; M2 rectangular 

with zygomatic process of maxillary originating posterior to M2 ; teeth pigmented and stout; mandible 

heavy; mental foramen below hypoconid of MI ; articular structure of mandible as in Blarinella; coronoid 

spicule is well formed and virtually as in living 

Blarinella. Late Pliocene to middle Pleistocene of 

central Europe.

Four species, Petenyia hungarica Kormos, Petenyia 

suavensis Pasa (1948), Petenyia neglecta Kretzoi 

(1943), and Petenyia stehlini Kretzoi (1943) are 

known. The genus is very close to the living Blarinella and differs chiefly in its loss of minute fifth 

upper antemolar.

Specimens examined. Central Geological Survey 

of Czechoslovakia SUtJG, OF 65566, a complete right 

lower jaw, and StJtJG, OF 65565, a complete left 

lower jaw, both from the earliest Pleistocene (Villafranchian) of southern Slovakia.

Genus BLARINELLA Thomas, 1911

Genotype: Sorex quadraticauda Milne-Edwards, 1872 

Dental formula 1-6-3/1-2-3; lower incisor relatively shorter than typical Sorex, upward curving, 

with two low scalloped cusps on upper cutting edge; 

lower cheek teeth stout with deep cingula not inflated, 

MI and M2 with entoconid close to metaconid and 

both cusps connected by high entoconid crest that is 

more prominent than in any other shrew; M3 with 

trigonid unreduced in size but talonid reduced to 

single low nearly conical but slightly bladelike hypoconid; P4 with very slight emargination of posterior 

basal outline and with lingual cingulum very prominent medial to protocone; M1 with very slight posterior

Medial

FIGURE 23. Petenyia hungarica Kormos, StJtJG, OF 65565. Left mandible and

dentition.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 35

emargination; M2 rectangular; teeth heavily pigmented as in Blarlna; zygomatic process of the maxillary originates posterior to M2 ', mental foramen 

beneath middle of 3/j; mandible stout; internal temporal fossa tending to be oval; condylar process and 

articular facets relatively large but otherwise as in 

Soreso.

One fossil and one living species have been named. 

They range from the middle Pliocene to Recent in 

China. The genus Petenyia from the late Pliocene 

and early Pleistocene of Europe is very closely related and differs chiefly in the loss of one upper antemolar.

Blarinella kormosi (Schlosser), 1924

Croddura kormosi Schlosser, 1924, Palaeontologia sinica, ser.

C, v. 1, no. 1, p. 5-6, pi., figs. 1, 2. 

Blarinella kormosi (Schlosser). Miller, 1927, Palaeontologia

sinica, ser. C, v. 5, no. 2, p. 8-9.

From Schlosser's description and illustrations (1924, 

p. 5-6; pi. 1, figs. 1, 2) and from Miller's discussion 

(192Y, p. 8-9), I am not sure that the very distinctive 

entoconid crest of MI and single-cusped talonid of M3 

are on this species. Photographs of Crocidura kormosi 

on file in the U.S. National Museum, which were sent 

by Professor Wiman to Dr. Miller and served as a 

basis for Miller's revision, show that between the time 

of Schlosser's study and the time of Wiman's photograph the specimen had lost its Ms. In addition, 

although the photographs are not clear in this respect, 

the entoconid crests on MI and M2 appear to be rather 

low, if present at all. The species could be a member 

of the tribe Blarinini and related to "/Sorex" kretzoii 

Sulimski, a possibility further discussed under "S."

kretzoii. The mandibular structures shown on Schlosser's illustration seem typical of the genus, as Miller 

pointed out. The talonid of Mj_ does not seem anteroposteriorly shortened as in the Blarinini nor does the 

mental foramen appear to be far enough back on the 

mandible for this tribe. Middle Pliocene (Ertemte 

fauna) of Mongolia. No specimen seen.

Blarinella quadraticauda (Milne-Edwards), 1872 

Figure 24

This living species of Blarinella, bears considerable 

superficial resemblance to Blarina in its dental formula, reduction of upper antemolars, pigmentation, 

stoutness of lower teeth, depth of cingula of lower 

teeth, and in the stoutness of mandible. It clearly 

differs from Blarina in its exceedingly prominent entoconid crest and posterior continuation of the labial 

cingulum to the rear of the hypoconid on Mt and M2, 

manner of reduction of the talonid of Ms, lack of 

anteroposterior shortening of the talonid of MI and 

M2l development of a cingulum medial to the protoconid of /**, rectangular M2, more anterior mental 

foramen, origin of the zygomatic process of the maxillary behind M2, and most importantly it differs from 

Blarina in its Sorex-like mandibular articulation and 

features of the posterior part of the mandible and of 

the cranium related to this type of articular structure.

Specimen examined. Chicago Natural History Museum 3621T, a male from Szechwa,n, China.

Soricini Incertae Sedis 

"Sorex" stehlini Doben-Florin, 1964

"Small Svrex, tooth cusps lightly colored yellowish 

orange. MI M3 =3.13 3.25 mm; Mz somewhat

Medial

FIGURE 24. Blarinella quadraticauda (Milne-Edwards), China, CNHM 36217. 

Left mandible and dentition and left upper dentition.
36 SUBFAMILIES AND GENERA OF THE SORICIDAE

smaller than Mlf I [lower incisor] doubly scalloped, 

first scallop quite close to the tooth base. Three singlerooted antemolars [including P±~\ in the lower jaw, 

the second and third very steeply placed. Molars with 

wide cingulum and massive cusps. Ml and M2 with 

hypoconulid [entoconid separated from hypolophid] 

and a lingual metaconid placed behind the protoconid. 

MS single-cusped in its talonid. Mental foramen beneath 

to behind the protoconid of M^. Condyles with large 

articular surfaces." (Doben-Florin, 1964, p. 41).

The articulation of the mandible is primitive and 

similar to that of Antesorex or Crocidosorex. Hence, 

the species appears to belong in the tribe Soricini. 

Unlike these two earlier Miocene genera, "Sorex" 

stehlini has three quite specialized features: the talonid 

of Ms is reduced to a single cusp, suggestive of the 

talonid of Petenyia, Blarwella, or some genera in the 

tribe Blarinini; the mental foramen is far to the rear 

in comparison to the other two earlier Miocene Soricini 

genera; and the teeth as a whole are massively constructed and have much heavier cingula than in these 

two genera. These specializations, three antemolars, 

and the more primitive mandibular articulation, relative to that on Sorex, all indicate that the species does 

not belong in Sorex. It would be possible to infer an 

ancestral relationship to Petenyia for this species 

except that Doben-Florin's illustrations do not seem 

to indicate the presence of the prominent entoconid 

crest characteristic of Petenyia, and related genera. 

The absence of the entoconid crest would suggest a 

relationship to the largely North American tribe Blarinini and possibly an ancestral relationship to late 

Miocene Adeloblarina through shortening of the relative anteroposterior length of the talonid of M± and 

M2 and specialization of the mandibular articulation.

"Sorex" stehlini is from the older Burdigalian fauna 

of Wintershof-West. No specimen was seen.

"Sorex" hibbardi Sulimski, 1962

Lower incisor has the two strong scalloped cusps 

on upper cutting edge more strongly developed than 

in Blarinella, and the incisor is relatively longer; 

cingula on lower molars strong and of uniform thickness rather than deepened; Ml and M2 with entoconids 

well developed but without very prominent entoconid 

crest of Blarinella and Petenyia; Ms with reduced 

talonid lacking entoconid as in earlier species of Blarwiella; coronoid process low and curved backward at 

tip; condyloid process very heavy with articular structures similar to Blarinella; teeth distinctly pigmented; 

internal temporal fossa large and triangular but with 

well-developed limula; late Pliocene (lower part of 

Breccia), Poland.

This shrew is intermediate between Sorex and Blarinella in almost all its characters. To judge from 

Sulimski's description, the species should be assigned 

to a new genus rather than be used as the basis for 

major enlargement of the definition of either established genus. This genus should not be described, 

however, without study of the specimens. No specimen was seen.

Genus ZELCEINA Sulimski, 1962

Genotype: Neomys soriculoides Sulimski, 1959 

Dental formula 1-5-3/1-2-3; upper incisor not bifid; 

P4 and M1 appear to have moderate posterior emargination; M2 rectangular, and zygomatic process of maxillary originates posterior to M2 ; lower incisor short 

with two low scalloped cusps on cutting edge much as 

in Neomys; MI with strong labial cingulum, but no 

published information indicates presence or absence of 

entoconid crest; Ms described as one-half smaller than 

MI, strongly reduced talonid without entoconid; teeth 

pigmented at tips; mental foramen below hypoconid 

of ML

The F4 is described by Sulimski (1959, p. 150; 1962, 

p. 476) as having a posteriorly projecting protocone 

the only published illustration of this tooth (1959, fig. 

5C) suggests to me that the protocone is anteriorly 

placed close to the parastyle but has a prominent cingular shelf posteromedial to the protocone as in Blarinella. There is no published figure of the occlusal view 

of the lower teeth, and it is not possible to say whether 

the Ms has both talonid and trigonid reduced as in 

some species of SoricuLus or has only the talonid reduced as in Petenyia^-Sulimski's (1959) figures 4a and 

4b of plate 3 seem to show that just the talonid is reduced. The posterior part of the mandible is distinctly 

more like Blarinella than Neomys except that the interarticular area is a little too narrow for Blarmella:, it 

is clearly too broad for Neomys, and in addition to the 

labial edge seems swollen in Sulimski's (1962) figure 

3b of text-plate 2, much as in living Blarinella.

Definite assignment of this genus to any tribe is dependent upon the presence or absence of the entoconid 

crest on MI, the nature of reduction of M%, and considered study of the articular structure of the mandible. Assignment to the tribe Neomyini seems questionable because of the lack of the bifid upper incisor in an 

Old World form of this little specialization and because of the posterior position of the mental foramen 

and the two cusps on the lower incisor. Assignment 

to the tribe Blarinini is not supported by the rectangular M2, the position of the zygomatic process of the
DIAGNOSES AND CONTENTS OF SUBFAMILIES 37

maxillary, and the narrowness of the interarticular 

area of the mandible. Assignment to the tribe Soricini 

near the genus Petenyia is favored by the general similarity of all features that have been described or illustrated but is left uncertain because of those features 

not described or illustrated.

If Zelceina is related to Petenyia, in which there is 

a strong entoconid crest and only the talonid of M3 is 

reduced, it probably should be recognized as a distinct 

genus, at least on the basis of differences in the articular structure of the mandible.

One species has been named Z. soriculoides (Sulimski) from the late Pliocene (lower part of W§ze Breccia), Poland. No specimen was seen.

Tribe BLARININI Stirton, 1930

Articular condyles of the mandible widely separated; 

lingual condylar emargination at least partially filled 

by a bony plate so that interarticular area is broad; 

lower condyle enlarged, offset lingually, moved anteriorly, and facing more ventrally relative to Sorex: 

condylar process supporting upper condyle curved toward skull; coronoid process deflected outward (fig. 1) ; 

superior pterygoid fossa moderately basined but with 

little or no pterygoid spicule; internal temporal fossa of 

the mandible small and oval or with a well-developed 

limula; external temporal fossa of mandible high and 

extending down coronoid process no farther than upper 

sigmoid notch or less, coronoid spicule prominent but 

very high on ascending ramus; mental foramen varies 

in position but usually beneath hypoconid of M± ; zygomatic process of maxillary originates opposite posterior 

part of M2 in all forms; M2 trapezoidal in occlusal view 

in most forms; most conspicuously M± with no trace of 

an entoconid crest but strong entoconid (with one exception), talonid ajiteroposteriorly shortened; entire 

Ms reduced, talonid tending to be further reduced to 

form crescentic crest with V of hypoconid weak or absent and reduced to minute shelf with small cusp in one 

genus; roots of Ms separated by distance less than 

transverse diameter of anterior alveolus; teeth heavily 

pigmented and very stout.

Blarina and Cryptotis are the only living genera included in this tribe. Blarina contains only 2 species 

(Hall and Kelson, 1959, p. 52) confined to the eastern 

part of the United States; Cryptotis contains at least 25 

species (p. 55-56) in the eastern half of North America 

and in Central America. It is the only shrew to have 

entered South America. The tribe contains a fair number of extinct genera in both North America and Eurasia 

and is known back to the late Miocene.

Genus ADELOBLARINA Repenning, n. gen.

Genotype: Adeloblarina berklandi, n. sp.

Diagnosis. A soricine shrew with stout molars comparable to Blarina but with mandibular structures more 

like those in Sorex; lower dental formula 1-2 ?-3; MI 

with crown having stout base, dense pigmentation, 

talonid shortened anteroposteriorly, entoconid distinctly separated from hypolophid and relatively close 

to metaconid but separated from metaconid by a pronounced valley, hypolophid extends past the entoconid 

to the lingual cingulum, posterolabial part of labial 

cingulum (near hypoconid) enlarged and attenuate 

posteriorly as in Paracryptotis and not as in Blarina 

so that labial embrasure between M± and M% would be 

less open than in Blarina; alveoli for M& are close together ; mandible moderately stout with depressed area 

that leads anterodorsally from mental foramen to region of the antemolars; mental foramen below middle 

of MI; coronid process narrow, its anterior edge forming a 90° angle with alveolar plane of horizontal ramus 

and its tip not deflected buccally as in Neomys or 

Blarina; inferior condyle oriented 90° from verticle axis 

of horizontal ramus and 45° from longitudinal axis, 

articular facet facing ventrally and offset lingually 

from plane of mandible at inferior sigmoid notch, anterior displacement of inferior condyle not as extreme as 

in Blarina, Cryptotis, or Notiosorex, slightly less than 

in Chimarrogale, about equal to Neomys, greater than 

in Sorex ; interarticular area presumably with moderate 

lingual emargination not filled with bone plate, but 

condyloid process anterior to interarticular area transversely thickened more than in most genera of the 

Neomyini; base of condyloid process, at its junction 

with ascending ramus, vertically narrow in comparison 

with all soricine shrews not included in tribe Soricini; 

superior pterygoid fossa of mandible has very shallow 

basin with poorly developed boss for insertion of superior pterygoid muscle; external temporal fossa of mandible not extended down coronid process as far as superior sigmoid notch; internal temporal fossa of mandible 

narrowly triangular with well-developed limula.

Etymology. Greek: Adelos [aSr/Aos], obscure, and 

BUrina; related to Blarina to an unknown degree.

Adeloblarina berklandi Repenning, n. sp. 

Figure 25

This species was named in honor of James O. 

Berkland, who found the type specimen while collecting with the author in southeastern Oregon during 

the summer of 1963.

Holotype. U.S. National Museum 23098; a right 

mandible lacking the superior mandibular condyle 

and bearing ML Collected by J. O. Berkland.
38 SUBFAMILIES AND GENERA OF THE SORICIDAE

Coronoid spicule 

External temporal fossa

FIGURE 25. Adeloblarina berklandi Repenning, n. gen. and n. sp., USNM 23098.

Right mandible and M\.

Type locality and age. U.S. Geological Survey 

vertebrate loc. Ml 040; in poorly exposed and sparsely 

fossiliferous orange tuffaceous mudstone at the south 

end of the Bowden Hills, Malheur County, Oreg. 

Late Miocene (Barstovian, possibly late Barstovian).

Diagnosis. A species of Adeloblarina of a size 

greater than Cryptotis parva, smaller than Blarina 

b<remcauda, and about equal to Neomys fodiens. Other 

than size, the postmandibular canal connecting the 

bottom of the pocketed internal temporal fossa with 

the dental foramen and the position of the mental 

foramen just behind the anterior root of MI may prove 

to be of specific importance when other shrews are 

found that belong to this genus.

Discussion. At first impression it might seem impossible to identify a fossil shrew from so incomplete 

a specimen as the type of Adeloblarina berklandi. 

The internal temporal fossa, however, shows that it is 

a soricid and not a heterosoricine shrew; the separation of the articular condyles and the degree of pigmentation show that the shrew belongs in the subfamily Soricinae; the anterior placement and lingual 

offset of the lower condyle, as well as the total lack 

of an entoconid crest on MI, eliminate the tribe 

Soricini; the total lack of a entoconid crest on MI, the 

stoutness of M\ in association with a strong cingulum 

and heavy pigmentation, the closeness of the alveoli 

of MS, and the association of moderate articular modification with these features strongly suggest assignment to the tribe Blarinini; the shallow basin of the 

superior pterygoid fossa, the width of the interarticular area, where preserved, just above the lower 

condyle, and the elevation of the bottom of the external temporal fossa above the upper sigmoid notch are 

evolutionary modifications of the Blarinini not found 

in the Neomyini.

Within the tribe Blarinini, Adeloblarina is clearly 

distinct in minimum separation of the articular condyles, in vertical narrowness of the base of the 

condyloid process, in lack of labial deflection of the 

coronoid process, in anterior position of the mental 

foramen, and in slenderness of the horizontal and 

ascending rami. All these features that separate 

Adeloblarina from other genera of the tribe Blarinini 

are clearly to be considered as primitive and approaching the condition observed in more primitive members 

of the tribe Soricini. Conversely, all other genera 

here included in the tribe Blarinini could be derived 

from Adeloblarina by further specialization of these 

same features.

The articular structure of the mandible of Adeloblarina is hardly more modified from the primitive 

articulation of Sorex than is the structure found in 

Blarinella; however, the modifications are those which 

lead to the more typical articulation of the tribe 

Blarinini. It would appear possible, therefore, that
DIAGNOSES AND CONTENTS OF SUBFAMILIES 39

the tribe diverged from a Sorex-\\ke, ancestor shortly 

before the time of Adeloblarina in the latter part of 

the Miocene.

Genus CRYPTOTIS Pomel, 1848

Genotype: Sorex parvus Say, 1823 

Dental formula 1-5-3/1-2-3 in all but one species 

which is 1-6-3/1-2-3; trigonid of M3 not greatly 

reduced relative to MI, but talonid greatly reduced to 

very small heel with single small conical cusp in living 

forms (somewhat less reduced in late Pliocene species) ; P3 much smaller than preceding upper antemolars (except P2 in species still retaining this tooth); 

P4 and M1 have moderate to slight posterior emargination, varying with species, and strong broadly curving hypoconal cingulum; M2 rectangular or very 

nearly so, but zygomatic process of maxillary originates opposite the metacone or metastyle; pigment less 

than in Blarina; mental foramen beneath hypoconid 

of MI; mandibujar articulation as in Adeloblcm/na 

except with lower articular facet placed somewhat 

more forward but well to rear of lower sigmoid notch, 

variable development of bone plate filling the lingual 

excavation of interarticular area but never developed 

to extent found in Blarina, in some species (as G. 

parva) bone plate not noticeably more developed than 

in Adelablarina; coronoid process slightly deflected 

labially and condyloid process supporting upper 

articular facet slightly curved toward cranium; coronoid process broader at level of superior sigmoid notch 

and condyloid process somewhat vertically deeper 

where it merges with ascending ramus than in Adeloblarina; internal temporal fossa triangular about as in

Adeloblarina', external temporal fossa deeper, reaching superior sigmoid notch as in Paraoryptotis. Middle 

Pliocene to Eecent, North America.

In the mandibular structure and in the position of 

the zygomatic process, Cryptotis is a primitive member 

of the Blarinini and is less specialized than all other 

genera of the tribe except Adeloblarina. It appears 

precocious and distinct in that there is extreme reduction of the heel of Ms. The interarticular area of the 

mandible in some species (as C. parva} has a strong 

resemblance to the Neomyini (new name), but this is 

less true of other species in which the bone plate filling 

the lingual emargination of the interarticular area is 

better developed. In addition, the lower articular 

facet is not offset as far lingually as in the Neomyini 

so that there is no groove separating the facet from 

the lower sigmoid notch; the absence of an entoconid 

crest in as unspecialized a shrew as Cryptotis seems 

very contradictory to the condition found in the 

Neomyini or, for that matter, in the Soricini. The 

zygomatic process is farther forward (relative to the 

tooth row) than in comparable shrews of the Neomyini 

or any shrew of the Soricini.

Cryptotis adamsi (Hibbard), 1953 

Figure 26

Blarina adamsi Hibbard, 1953, Jour. Paleont. v. 27, 

no. 1, p. 29-31, figs. 2B, 3C, 4B, C, and D, and 5B.

Assignment of this species to Blarina or Cryptotis 

is a matter of definition. The species is virtually 

identical in mandibular structure and lower and upper 

dentition to living Cryptotis. It does, however, retain 

the minute P2 lost in the living species of the genus

FIGURE 26. Cryptotis adamsi (Hibbard), topotype, UM 28411. Right mandible

and dentition.
40 SUBFAMILIES AND GENERA OF THE SORICIDAE

and, hence, has the dental formula of living Blarina. 

The assignment of this species to Cryptotis in this 

report is based upon the belief that the following 

differences between it and Blarina represent phylogenetically significant characters and that the presence 

of P2 is an expectable primitive feature. From species 

assigned to Blarina, Cryptotis adamsi differs in (1) 

greater anteroposterior shortening of the talonid of 

MI, (2) more posterior placement of the metaconid 

of MI, relative to the position of the protoconid, (3) 

greater reduction of the heel of MS, (4) retention of a 

primitive blarinine mandibular articulation and associated jaw structure, and (5) a rectangular M2. Items 

(4) and (5) could be considered primitive characters 

that represent an ancestral condition in the Blarina 

lineage, but items (1), (2), and (3) are clearly specializations in the late Pliocene species not yet achieved 

in the living species of the Blarina lineage.

A fragmentary specimen assigned to this species has 

been found in the middle Pliocene (Hemphillian) 

Christmas Valley local fauna of Lake County, Oreg. 

(U.S. Geol. Survey vertebrate loc. M1061). The 

geologic range is, therefore, middle Pliocene (Hemphillian) of Oregon and late Pliocene (early Blancan) 

of Kansas.

Specimens examined. Topotypes, University of 

Michigan Museum of Paleontology 28411, 24352, and 

27310, all well-preserved mandibles from the Kexroad 

local fauna, Kansas, and USGS M6511, a fragmentary 

mandible from the Christmas Valley local fauna, 

Oregon.

Cryptotis? meadensis Hibbard, 1953

Less reduced heel on Ms, more lingual metalophidprotolophid junction, and somewhat less forward 

placement of lower articular facet than in living 

Cryptotis.

These features appear less advanced but related to 

those of the living forms. CA meadensis, however, 

is very unusual in not having a distinctly isolated 

entoconid on MI and in having a P4 that is very similar to the P4 in the Limnoecinae. The articular structure of the mandible of C.1 meadensis clearly could 

not belong to a limnoecine shrew. Late Pliocene 

(early Blancan) of Kansas.

Specimen examined. University of Michigan Museum of Paleontology 27266, the type, from the Rexroad fauna of Kansas.

Cryptotis kansasensis Hibbard, 1957

The teeth and mandible, as determined from published descriptions of both species, are similar to those 

in the living species C. magna. No specimens of either 

species have been examined during this review, and I 

am unable to see how the two species differ. From the 

living C'ryptotis parva, they differ in greater size, lack 

of a posterior emargination of the upper molars and 

P4, more trapezoidal form of M2, a more rounded 

anterointernal corner of P4, stouter lower molars, and 

greater filling by bone of the lingual emargination of 

the interarticular area of the mandible. C. kansasensis 

is from the early Pleistocene of Kansas.

Posterior

FIGURE 27. Cryptotis parva (Say), Kansas, UCMVZ 40144. 

and dentition and left upper dentition.

Left mandible
DIAGNOSES AND CONTENTS OF SUBFAMILIES 41

Cryptotis parva (Say), 1823

Figure 27

This living species has also been reported as fossil 

from the middle Pleistocene deposits of Kansas (Hibbard, 1963, p. 203-204). Only Recent specimens were 

seen.

Cryptotis mexicana (Coues), 1877

This living species has been reported as fossil from 

late Pleistocene deposits of Mexico (Findley, 1953, p. 

633-639). No specimen was seen.

Genus PARACRYPTOTIS Hibbard, 1950

Figure 28

Genotype: Paracryptotis rex Hibbard, 1950 

Dental formula 1-5-3/1-2-3; Ms well reduced relative to MI with well-developed crescentic talonid crest; 

M! with deep labial cingulum from protoconid to 

posterior part of tooth (behind hypoconid), where 

it is better developed than in living Blarina, metalophid joins protolophid slightly labial to position of 

this junction in Adeloblarina but well lingual to position of junction in living Blarina, base of crown broad, 

hypolophid does not extend lingually as far as in 

living Blarina, and entoconid is slightly more 

posterior; P3 smaller than preceding upper antemolars; P4 and M1 with no posterior emargination; 

M2 trapezoidal and zygomatic process of maxillary 

originating opposite metacone; pigmentation slightly 

less than in Blarina; mental foramen from beneath 

center of M\ to below posterior root of M\ anterior 

to average position in living Blarina but overlapping

most of range found in living Blarina', mandibular 

articulation as in Blarina with broad interarticular 

area, but inferior facet not placed as far forward and 

slightly visible behind lower sigmoid notch in labial 

view; superior pterygoid fossa a shallow basin with 

no pterygoid spicule; internal temporal fossa small 

and circular; external temporal fossa reaching superior 

sigmoid notch; coronoid process moderately deflected 

labially. Hibbard (1950, p. 122-127) compared this 

genus with Cryptotis and with Blarina. Paracryptotis 

appears to be closely related to Blarina on the basis of 

the reduction in size of the entire M^ and the trapezoidal occlusal outline of M*, in addition to the articular 

structure of the mandible and other similarities to 

Blarina mentioned by Hibbard. In the dental formula 

and possibly in the structure of the talonid of M$, 

Paracryptotis is more closely related to Cryptotis. In 

the lingual position of the metalophid-protolophid 

junction and the development of the cingulum on M^ 

Paracryptotis is less advanced than living Blarina but 

is about equal in this feature to the late Pliocene 

Blarina gidleyi and to Cryptotis. In the mandibular 

articulation, Paracryptotis is somewhat less advanced 

than Blarina but more advanced than Cryptotis.

The genus includes only one species, Paracryptotis 

rex Hibbard, from the late Pliocene (early Blancan) 

Rexroad fauna of Kansas and the middle Pliocene 

(Hemphillian) Rome fauna of Oregon.

Specimens examined. Two topotypic mandibles 

labeled University of Michigan Museum of Paleontology 47760. Uncataloged specimen from USGS 

vertebrate loc. M1078, Rome local fauna, Oregon.

FIGTTKE 28. Paracryptotis rex Hibbard, topotype, UM 47760. Right mandible and dentition.
42 SUBFAMILIES AND GENERA OF THE SORICIDAE

Genus SHIKAMAINO SOREX Hasegawa, 1957

Genotype: Shikamainosorex densicingulata 

Hasegawa, 1957

Articular structure typical of Blarinini; differs from 

Paracryptotis in greater inflation of labial <cingulum of 

MI to J/3, greater reduction of heel of J/3, and greater 

anterior displacement of lower articular facet of 

mandible.

The advancement of the lower articular facet, 

which is hidden behind the mandible in labial view, 

suggests a greater similarity to Blarina. The other 

points of difference between Shikamainosorex and 

Paracryptotis, however, are an accentuation of those 

features which separate Paracryptotis from Blarina. 

In addition, the junction of the metalophid to the 

protolophid is as in Paracryptotis and early Pleistocene Blarina and far more lingually located than in 

living Blarina. Middle Pleistocene, Totigi Prefecture, 

Japan.

The differences between S. densicingulata Hasegawa 

and Paracryptotis rex are of no greater magnitude 

than those between Blarina gidleyi from the late Pliocene of Idaho and the living Blarina. It would seem, 

by comparison, either that Hasegawa's species densicingulata should be included in the genus Paracryptotis 

or that Gazin's species gidleyi should be placed in a 

new genus ancestral to Blarma. This should not be 

done, however, without comparison of the specimens.

Sulimski (1962, p. 472-473) suggested that Shikamainosorex densicingulata may be related to Anourosorex. The very prominent cingula on the lower 

molars, the lesser reduction of M3, as well as the type 

of reduction seen on the talonid of M3 , the height of the 

inferior border of the external temporal fossa of the 

mandible (which is as in Paracryptotis and Blarina)^ 

and most importantly the articular structure of the 

mandible rule out a relationship with Anourosorex. 

Anourosorex, with characteristic development of its 

features, is known from deposits in Japan of the same 

age as those that contained S. densicingulata (Shikama 

and Hasegawa, 1958). No specimen was seen.

"Sorex" dehneli Eowalski, 1956

This shrew seems close to Paracryptotis and to 

Shikamainosorex. As far as can be judged from 

published accounts, the MI is as in Paracryptotis and 

Blarina gidleyi', the M3 is well reduced (at least on 

the holotype) and has a reduced, crescentic talonid; 

the mental foramen is as in the two American genera.4

4 Although Kowalski (1956, p. 346) stated that the mental foramen is below P4, his figures (pi. 1, figs. 5, 6) clearly show it below 

the middle of M,. The talonid of Mi in the holotype is broken, and 

the isolated trigonid resembles, at first glance, a P4 ; this situation 

suggests that Kowalski inadvertently mistook this talonid for P4 while 

describing the position of the mental foramen. He is not alone, for

The mandibular articulation compares favorably with 

Paracryptotis in form and stage of advancement. 

The knoblike structure of the apex of the coronoid 

process seems unique. Because of the mandibular articulation and the tooth structure, the species seems, 

at least, to belong in the tribe Blarinini and not to 

the genus Sorex. Kowalski (1960, p. 170) noted similarities with Blarinoides mariae Sulimski and suggested that they are probably congeneric. The species is known from the late Pliocene of Poland. No 

specimen was seen.

Genus BLARINA Gray, 1838 

Figure 29

Genotype: Sorex talpoides Grapper, 1830 

Dental formula as in Sorex 1-6-3/1-2-3; talonid of 

MI and M2 shortened; M3 least reduced of tribe (possibly excepting Adeloblarina), but with talonid shortened and intraspecifically variable in development of 

crescentic crest, alveoli close together; P4, M1 , and 

M2 with only slight emargination of posterior basal 

outline; M2 trapezoidal and zygomatic process of maxillary originates opposite this tooth; teeth stout and 

heavily pigmented; mandible stout; mental foramen 

varying from beneath protoconid of MI to between 

MI and M2 (farther to the rear than in any other 

shrew besides Trimylus-, articular facets of mandibular condyle widely separated, lower facet placed well 

forward and hidden behind mandible at lower sigmoid notch in labial view, interarticular area broad; 

superior pterygoid fossa as shallow basin with no 

spicule; external temporal fossa extends downward to 

superior sigmoid notch and follows its ventral margin 

to upper condyle.

Of all living shrews for which a reasonable sample was examined during this review, the genus 

Blarina stands out as being the most variable intraspecifically. Most notable are the variations in the 

structure of the talonid of M3 , the position of the 

mental foramen, and the form of the cingulum on the 

lower cheek teeth. In the living species B. Irevicauda, the talonid of M3 varies from a considerably 

reduced and rounded heel that has a well-developed 

crescentic crest similar to Paracryptotis to a rectangular heel that has a reasonably sharp V on the hypoconid, a conspicuous entoconid elongated to form a 

short entoconid crest separated from the hypolophid 

by a valley, and a fairly angulate posterolingual corSulimski, who prepared Kowalski's illustration of the type, apparently took Kowalski's statement at face value without referring to 

the specimen or his own artistry because he (1962, p. 471) made the 

same mistake in thinking the trigonid of Mi was the P4 when he 

compared his "8orea>" kretzoii with "Sorex" dehneli.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 43

Superior

pterygoid

fossa

M, of 6212 \ I 1 ' ,// 

showing variation 

of talonid pattern

Coronoid spicule 

External temporal

Depressed plate posterior to superior 

pterygoid fossa filling lingual 

emargination of interarticular area

FIGUBE 29. Blarina brevicauda (Say), author's collection 6188 from Michigan and 

MS of 6212 from New Jersey. Left mandible and dentition and left P4 .

ner in occlusal view not greatly different from that 

in Sorex. The range in position of the mental foramen in the species Blarina brevicauda is nearly as 

great as that seen within the subfamilies throughout 

their geologic history. Other features, as the position 

of the junction of the metalophid with the protolophid on MI, the posterior emargination of .P4 and 

M1 , or the structure of the mandibular articulation 

seem as stable as in any other shrew genus. Four species have been recognized as fossil.

Blarina gidleyi Gazin, 1933

Figure 30

MS within more /Sorex-like variation of living Blarina ~brevicauda', pigmentation of the teeth and structure of mandibular articulation identical to those features in living species; MI with metalophid-protolophid junction placed far lingual to position in living 

species and similar to Paracryptotis, hypolophid does 

not extend as far lingual as in living species, and 

cingula as in Paracryptotis and Adeloblarina rather

Posterior

Medial

FIGURE 30. Blarina gidleyi Gazin, topotype, UM 3304. Right mandible

and dentition.
44 SUBFAMILIES AND GENERA OF THE SORICIDAE

than reduced in region of hypoconids and deepened 

below protoconids as in living species; upper teeth 

unknown; mental foramen below midpoint of MI; 

horizontal ramus without sharply angulate inferior 

border of living species but possibly stouter throughout.

Most of these characters are clearly primitive, and 

the species is intermediate between the living species 

and Adeloblarina on the basis of these differences. It 

is possible the differences are of generic importance 

and that, therefore, this species should not be included in Blarina.

Late Pliocene (early Blancan) Hagerman fauna of 

Id:Jiu.

Specimen examined. University of Michigan Museum of Paleontology 3304, as well preserved mandible.

The other species of Blarina reported as fossil are 

all from the late Pleistocene. None of these were 

seen.

Blarina fossilis Hibbard, 1943. Kansas.

Blarina simplicidens Cope, 1899. Pennsylvania.

Blarina brevicauda (Say), 1823. A living species in ,the eastern half of North America and reported from the late Pleistocene from Oklahoma and Kansas.

Genus BLARINOIDES Sulimski, 1959 

Genotype: Blarinoides mariae Sulimski, 1959

Dental formula 1-6-3/1-2-3; P4 very large and inflated; MS greatly reduced but talonid not reduced 

relative to trigonid, distinct entoconid; P4 and Ml 

with moderate emargination of posterior basal outline as in Blarina; M2 trapezoidal and zygomatic process originating opposite metacone as in Blarina', teeth 

pigmented and stout; mandible very heavy; mental 

foramen between roots of MI ; articular facets of mandible widely separated and about as in Blarina.

Although very similar to Blarina, according to Sulimski (1959, p. 147) Blarinoides differs in its larger 

size, weaker coronoid spicule, inflated P4 , presence of 

entoconid on M3 of all individuals, and relative sizes 

of the upper antemolars. In labial position of the 

metalophid-protolophid junction and development of 

the cingulum on MI, Blarinoides appears much closer 

to the living Blarina than to the late Pliocene Blarina 

gidleyi.

One species named: Blarinoides mariae Sulimski. 

Known from the late Pliocene of Poland and Hungary 

(Sulimski, 1962, p. 483). Kowalski (1960, p. 170) 

suggested that his species "Sorex" dehneli, from the 

early Pleistocene of Poland, may belong in the genus 

Blarinoides, a possibility that seems logical to me 

from a review of the published accounts. No specimen was seen.

Kretzoi (1962, p. 353) placed this species in the 

genus /Shikamainosorex. Kowalski and Li (1963, p. 

143) did not agree with such a generic assignment and 

pointed out that Shikamainosorex has a greater reduction of MS and a broader cingulum than Blarinoides. These differences appear real to me and in 

addition the MS of Blarinoides clearly retains a small 

entoconid that is not present on the reduced talonid of 

MS of Shikamainosorex. From the published descriptions it appears that Shikamainosorex is clearly 

related to Paracryptotis and that Blarinoides is related more to Blarina, as the authors of both genera 

originally supposed.

Blarinini? Incertae Sedis 

"Sorex' kretzoii Sulimski, 1962

This shrew appears, from Sulimski's description 

and illustrations, to have a heavy lower articular 

facet on the mandible that is basically horizontal and 

notched in its upper edge and is shifted forward only 

slightly from its position in Sorex, a reasonably broad 

interarticular area, an upper articular facet that is 

rotated well toward the longitudinal axis of the mandible, a labially deflected coronoid process, no entoconid crest on the lower molars, and a relatively little 

reduced J/3 . The combination of these features suggests the tribe Blarinini, but unfortunately most of 

the characters cannot be clearly verified in the published description. The vagueness results not so much 

from the fault of the description, which is better than 

the description of many fossil shrews, but from the 

generalized nature of "Sorex" kretsoii. This is particularly so in its mandibular articulation, which 

could be either a Blarinella-tjpe in the tribe Soricini 

or a very primitive articulation in the tribe Blarinini. 

From Sulimski's figures (text-pi. 2, figs. 9, 10), the 

articulation appears to belong in the Blarinini, but if 

so it is closer to the Soricini in lacking a forward 

shift of the lower articular facet than is any member 

of the Blarinini except Adeloblarina from the late 

Miocene.

Tribal assignment must be decided on the basis of 

the lingual offset of the lower articular facet from 

the lower sigmoid notch, rotation of the upper articular facet toward the longitudinal axis of the mandible, degree of curvature toward the skull of the condyloid process supporting the upper facet, labial deflection of the coronoid process, position of the origin 

of zygomatic process of the maxillary, and the occlusal outline of M2. None of these are determinable 

from the published description. One strong point in 

favor of assignment to the Blarinini is the positive 

statement of Sulimski (1962, p. 469) that the ento-
DIAGNOSES AND CONTENTS OF SUBFAMILIES 45

conid crest ("crest between the entoconid and metaconid") is lacking; no shrew assigned to the Soricini 

is yet known that lacks the entoconid crest.

There is a rather striking resemblance between 

Schlosser's (1924) figure 1 of plate 1 of Blarinella 

kormosi (Schlosser) and Sulimski's (1962) figure 9a 

of text-plate 2 of "Sorex" kretzoii Sulimski. If 

Blarinella kormosi does not have the very distinctive 

entoconid crest on J/i and M2 and the reduced talonid 

on M% of Blarinella, as Schlosser's illustrations suggest and as might be implied by Miller's (1927, p. 9) 

words "the characters in which * * * Blarinella kormosi differs from the living Chinese B. wardi appear 

to approach, in every instance, those of Blarina * * *" 

(Blarina does not have the entoconid crest or reduced talonid of Blarinella}, it seems probable that 

Blarinella kormosi is very closely related to "xSWea?" 

kretzoii, and is not Blarinella.

"/Soreoc" kretzoii is from the late Pliocene (lower 

part of the We.ze Breccia), Poland. No specimen was 

seen.

Feisorex pohaiensis Kowalski and Li, 1963

The authors of this genus and species do not mention whether the entoconid crest is present or absent 

on the .3/1-2 of the single known mandible, but it appears to be absent in their illustrations. The unusually specialized manclibular articulation could belong 

to either the tribe Blarinini or the tribe Neomyini, to 

judge from the published description and illustrations. 

The deep pit (superior pterygoid fossa) in the area 

just anterior to the superior condyle of the mandible, 

however, strongly suggests that Peisorex belongs in 

the tribe Blarinini.

According to the authors (Kowalski and Li, 1963, 

p. 142), Peisorex pohaiensis is a large shrew characterized by a connection between the internal temporal fossa (their "fossa pterygoidea") and the mandibular (dental) foramen, by a simple coronoid process lacking any spicule, and by the prominent basin of 

the superior pterygoid fossa. The shrew is known 

from the middle Pleistocene of China. No specimen 

was seen.

Tribe NEOMYINI Repenning, new name 

Diagnosis as for tribe Blarinini except as follows: 

Lingual condylar emargination of mandible open, 

consequently interarticular area very narrow; lower 

condyle elongate, offset lingually from plane of lower 

sigmoid notch to greater extent than in Blarinini, 

more anteriorly placed and facing more ventrally 

than in Soricini, separated in many specimens from 

lower sigmoid notch by groove because of extreme 

lingual offset (fig. 1) ; superior pterygoid fossa deeply

basined with pterygoid spicule in most forms; external temporal fossa of mandible low with ventral 

margin varying from level of superior sigmoid notch 

to level of lower articular facet; coronoid spicule high 

to very low; coronoid process with very strong tendency to curve anteriorly at tip but tip spatulate in 

a few forms; mental foramen beneath middle of J/i 

as in Sorex and not placed so far to rear as in many 

genera of Blarinini; M2 rectangular in occlusal view 

and zygomatic process of maxillary originates posterior to M2 in nearly all forms except those having 

extreme reduction of last molars; protocone of P4 

tends to be farther separated from parastyle and 

placed farther medially than in Sorex; upper incisor 

bifid or fissident, as in Trimylus, in most Old World 

forms; bifid upper incisor not present in New World 

forms; M\ with prominent entoconid crest in most 

forms but weak in some and missing in those having 

extreme reduction of last molars; MS variable and 

may be unreduced as in Sorex, with talonid only 

reduced as in BlarineUa, greatly reduced as in Blarina, 

or entirely lost; labial cingula on lower molars never 

deep as in Blarina, well developed to absent; teeth 

pigmented over less extensive area or lightly pigmented relative to Sorex or colorless, slim to stout.

The Neomyini include nine living genera, more than 

any other tribe of the subfamily Soricinae; however, 

each of these genera include very few living species, 

many only one. The fossil record of the tribe is 

clearly very incomplete. The oldest specimens are of 

early Pliocene age, and some of these are the most 

specialized forms known, a situation illustrating how 

far the history of the Soricidae is from being completely documented. Because the bifid upper incisor 

is also known in the Heterosoricinae, is sometimes 

present in the Soricini, and is found in one crocidurine shrew, it would seem to be a primitive character 

inherited from a common ancestor.

The Neomyini are largely a Palearctic tribe although there are three genera from North America. 

Except for the living Anourosorex and fossil relatives of this genus, most Old World forms have bifid 

upper first incisors; the New World genera do not 

although they are very close to the eastern Asian 

Chodsigoa in many other characters. Two lineages 

are thus suggested. Within the Old World forms no 

suggestion of further phylogeiietic breakdown has 

been noted, but rather, the known forms fit rather 

neatly into a uniformly gradational series from the 

little specialized Neomys to the extremely specialized 

Anourosorex. This graded series is entirely morphologic and is not related to an evolutionary sequence 

insofar as the present fossil record reveals.
46 SUBFAMILIES AND GENERA OF THE SORICIDAE

Coronoid spicule 

External temporal fossa

Boss for insertion of superior 

pterygoid muscle

FIGURE 31. Neomys fodiens (Pennant), France, UCMVZ 30044. Left mandible 

and dentition and left upper dentition.

Genus NEOMYS Kaup, 1829 

Figure 31

Genotype: Sorex daubentonii Erxleben, 1777 

Dental formula 1-5-3/1-2-3; upper incisor bifid in 

about half of the specimens examined; fourth upper 

antemolar small; P4 with protocone shifted more medial from parastyle than in Sorex; lower incisor with 

one weakly defined and irregular cusp on cutting 

edge; labial cingulum strong on lower molars; pigment extensive for tribe but less so than is typical of 

8orex\ pterygoid boss of mandible moderately developed; upper articular facet of mandible oval; lower 

articular facet not placed so far forward that it is 

hidden in labial view; coronoid spicule high; external 

temporal fossa high and terminating at superior sigmoid notch; tip of coronoid process narrow with 

prominent anterior curve. Early Pleistocene to Recent 

of Europe, Kecent of Asia.

Except for dental formula, upper and lower incisor, 

and P4, there appears to be no way to separate Neomys from Sorex on the basis of dentition alone. 

Among European species of Sorex, Neomys appears 

distinguishable by the higher cusps in unworn teeth 

(Miller, 1912, p. 68). 

Five species have been reported as fossil:

Neomys browni Hinton, 1911. Late Pleistocene, England. No 

specimen was seen.

Neomys fodiens (Pennant), 1771. Late Pleistocene, England; 

a living species. Only Recent specimens were seen from 

the Museum of Vertebrate Zoology, University of California.

Neomys newtoni Hinton, 1911. Early Pleistocene, England and 

Czechoslovakia. One specimen was seen, author's collection 

6340, from Czechoslovakia. The specimen was returned to 

the Czechoslovakian Geological Survey since preparation of 

this report; the new catalog number is unknown.

Neomys intermedius Brunner, 1952. Middle Pleistocene, Germany. No specimen seen.

Neomys castellarina Pasa, 1948. Middle Pleistocene, Italy. 

No specimen seen.

One living species other than N. fodiens is recognized by Ellerman and Morrison-Scott (1951, p. 61).

Genus PETENYIELLA Xretzoi, 1956 

Genotype: Sorex gradlis Petenyi, 1864

This genus was named in a footnote (Kretzoi, 1956, 

p. 160), and Sorex gracUis Petenyi (1864, p. 70) was 

cited as the genotype. Kretzoi (1959a, p. 239) gave 

a brief diagnosis.

Sulimski (1962, p. 479^80, 483) recognized the 

genotypic species Petenyiella gradlis and a new species Petenyiella zelcea from the W§ze Breccia on the 

basis of direct comparison with the Hungarian specimens. Unfortunately most of his description is in 

comparison to the genotype, which is poorly described and illustrated in available publications. From 

Sulimski's discussion (1962, pages just cited, and 1959, 

p. 157-159 under the genus Suncus], the following 

features place the genus Petenyiella in the Neomyini.

Condyles fairly widely separated, lower facet elongate, and interarticular area narrow; lower incisor 

with two rather indistinct lobes on its cutting edge;
DIAGNOSES AND CONTENTS OF SUBFAMILIES 47

MI "similar in structure" (Sulimski, 1959, p. 158) to 

Suncus and therefore assumed to have well-developed 

entoconid crest; Ms "strongly reduced talonid with 

poorly recognized hypoconid" also similar in structure 

to Suncus in which the talonid is more reduced than 

in Crocidura in European forms of Suncus and Crocidura (Miller, 1912, p. 84); teeth pigmented at tips; 

size very small.

Kretzoi (1959a, p. 239) gave the following characters in diagnosis of P. gracilis: soricine of Suncus 

size with four upper unicuspids which are comparable 

in form to the unicuspids of Sorex, two-lobed lower 

incisor, and delicate yellow pigmentation.

Kormos (1934, p. 306, fig. 38) gave a good illustration of the mandible under the new name Pachyura 

pannonica. He also suggested that the species might 

be considered a new genus, for which possibility he 

provided the new generic name Allopachyura. Because he did not apply this name to the species, it 

would appear to have no validity, and Petenyiella is 

here used.

As Kretzoi said (1962, p. 353), the genus is close to 

Sorictdus [Episoriculus as here used]. It is a Neomyini shrew characterized by its small size, by its 

rather unspecialized mandibular condyles with the 

small lower condyle farther to the rear than in Neomys, and by a greatly reduced and single-cusped talonid on MBThere appear to be significant differences in the 

development of the upper condylar process in Petenyiella. P. zelcea from the W§ze Breccia (Sulimski,

1959, pi. 3, fig. 2, and pi. 4, fig. 1; 1962, pi 2, fig. 1, 

and text-pi. 2, fig. 1) most closely resembles P. gracilis \P. pann&rdca of Kormos, 1934, fig. 48] and has 

the upper articular facet well above the upper sigmoid notch and the lower facet well posterior to the 

lower sigmoid notch. The specimens which Sulimski 

referred to P. gracilis from the W§ze Breccia (Sulimski, 1959, pi. 3, fig. 1, and pi. 4, fig. 2) have the upper 

articular facet below the level of the upper sigmoid 

notch and the lower facet well forward and almost 

hidden behind the lower sigmoid notch in lateral view. 

This is a much greater difference than has been observed in any other genus of shrew. The possibility 

that two genera are represented and also that Sulimski has referred the wrong one to P. gracilis is here 

suggested.

The genus is known from the late Pliocene of 

Hungary and from the late Pliocene and Pleistocene 

of Poland (mostly from the lower part of the W§ze 

Breccia, according to Sulimski, 1962, p. 483). No 

specimen was seen.

Genus EPISOBICTTLTTS Ellerman and Morrison-Scott, 1951 

Figure 32

Genotype: Sore® caudatus Horsfield, 1851 

Dental formula 1-5-3/1-2-3; upper incisor bifid as 

in Neomys, fourth upper antemolar very small; P* 

with protocone placed close to parastyle as in Sorex-, 

lower incisor as in Neomys but shorter; labial cingulum on lower molars as in Neomys ; Ms with heel very 

slightly reduced relative to Neomys and V of hypoFIGUEE 32. Bpisoriculus caudatus (Horsfield), India, CNHM 35415. Left 

mandible and dentition and left upper dentition.
48 SUBFAMILIES AND GENERA OF THE SORICIDAE

conid not so acute; pigmentation less than in Neomys 

lacking on J/3 and only at very tip of hypoconid on 

talonid of MI and M2 ; pterygoid boss of mandible almost undetectable; upper articular facet of mandible 

oval as in Neomys; lower articular facet placed farther forward than in Neomys and barely visible behind lower sigmoid notch in labial view; coronoid 

spicule lower than in Neomys and directed posteroventrally toward lowest point of upper sigmoid notch 

rather than above it as in Neomys; external temporal fossa slightly lower than in Neomys and extending 

well below upper condyle. Recent of eastern Asia 

and late Pliocene of Europe.

Episoriculus was proposed as a subgenus of Soriculus by Ellerman and Morrison-Scott (1951) because of marked differences between species previously included in Soriculus. They pointed out (p. 57) 

that the external differences between these species are 

greater than those between Sorex and Blarinella. Following Osgood's suggestion of the subgeneric importance of the extra tooth in Chodsigoa, they also considered this genus a subgenus of /Soriculus. In the 

present review, mandibular and dental characters also 

seem to separate these three subgenera to a significant 

degree. Although these differences are not as great 

as those between Sorex and Blarinella, they are as 

great as those that separate many genera of the Neomyini. Because of these differences the three subgenera which Ellerman and Morrison-Scott placed in 

the genus Soriculus are here given full generic status 

for the sake of maintaining the degree of morphologic 

distinctiveness noted between genera of the Neomyini 

and between most genera of the Soricidae.

Episoriculus differs from Chodsigoa in having one 

more upper antemolar, in more extensive pigmentation, in a less anterior pitch of the tip of the coronoid process, in a lesser width of the coronoid process 

at the level of the superior sigmoid notch, and in its 

oval upper articular facet. These differences are not 

great but are of the same sort and magnitude that 

separate Episoriculus from Neomys. If Chodsigoa 

and Episoriculus are to be considered subgenera under one genus, it should be under the genus Neomys', 

the characters of Episoriculus and Chodsigoa differ 

much more from those of Soriculus.

Ellerman and Morrison-Scott recognize two living 

species of Episoriculus.

Specimens exmnined. Two specimens of E. caudatus (Horsfield) : Chicago Natural History Museum 

39632, a male from Szechwan, China, and 35415, a male 

from Sikkim, India. These represent two subspecies.

Episoriculus gibberodon (Petenyi), 1864

Equals Crocidura gibberodon Petenyi, 1864; Soriculus kubinyi Kormos, 1934; Soriculus gibberodon 

(Petenyi) of Kretzoi, 1956; and Asoriculm gibberodon (Petenyi) of Kretzoi, 1959. Elevation of the 

subgenera of Soriculus to generic stature necessitates 

reevaluation of the generic assignment of this fossil 

species. The following characters bar its assignment 

to either Soriculus or to Chodsigoa and indicate its 

affinity with living Episoriculus: dental formula 

1-5-3/1-2-3; lower incisor short; Ms only slightly reduced with distinct entoconid; complete labial cingulum on lower molars; pigment present on tips of 

cuspids of the lower molars; no anterior inclination 

of tip of coronoid process; relative narrowness of 

ascending ramus at level of upper sigmoid notch; 

coronoid spicule higher than in Chodsigoa or Soriculus', lower articular facet not placed as far forward 

as in Soriculus (not as far as in the living Chodsigoa 

or Episoriculus} ; and upper articular facet oval.

Late Pliocene of Hungary and Poland. No specimen was seen.

Genus CHODSIGOA Kastschenko, 1907 

Figures 33, 34

Genotype: Soriculus hypsibius deWinton, 1899 

Dental formula 1-4-3/1-2-3; upper incisor bifid as 

in Neomys ; upper antemolars of uniform size; P4 with 

protocone as in Sorex and with extreme emargination 

of posterior basal outline one species, at least, exceeding all other shrews in this feature except Notiosorex 

jacksoni; emargination of M1 and M2 also extreme; 

lower incisor as in Episoriculus; P± variable in angularity of junction of labial shearing blade with crest 

formed by posterior cingulum; Ms variable in reduction of talonid from nearly unreduced (fig. 34) as in 

Episoriculus to noticeably reduced (fig. 33) with 

talonid shortened, hypolophid crescentic, entoconid lost 

as distinct cuspid, and entoconid crest almost lost but 

present as very low lingual crest; pigmentation very 

reduced but present on tips of antemolars, tip of paracone of P4 and in one species on very tip of protoconid 

of MI ; pterygoid spicule moderately developed on ridge 

running from internal temporal fossa to superior condyle well below upper sigmoid notch; upper articular 

facet triangular as in Anourosorex and Notiosorex,' 

lower articular condyle as in Episoriculus and slightly 

visible behind lower sigmoid notch in labial view; coronoid spicule low entirely at or below level of upper 

sigmoid notch; external temporal fossa lower than in 

Episoriculus and terminating downward closer to level 

of lower condyle than to upper condyle. Pleistocene 

and Recent of eastern Asia.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 49

FIGURE 33. Chodsigoa hypsibia (deWinton), China, CNHM 35760. Left mandible and dentition and left upper dentition.

FIGURE 34. Chodsigoa salenskii (Kastschenko), China, CNHM 39614. Left mandible and dentition and left upper dentition.

Three species in this group are recognized by Ellerman and Morrison-Scott (1951, p. 60-61) of which only 

two have been seen. Chodsigoa hypsibia, with a uniformly curving labial crest of P± and a reduced heel 

on MZ, is almost indistinguishable from the living 

Notiosorex crawfordi of North America on the basis 

of its dentition. Typical of the Old World Neomyini, 

the trigonid of MI is longer relative to the talonid than 

in Notiosorex, and the upper incisor is bifid. There is

pigment on the very tip of the protoconid of M\ in 

N. crawfordi and none on the specimen of G. hypsibia 

available for study, but there is pigment on the tip of 

the MI protoconid of G. salenskii. Both species of Chodsigoa have greater emargination of P4 and the upper 

molars than N. crawfordi but not greater than the late 

Pliocene Notiosorex jacksoni. There is also great similarity in the mandibular structures although the coronoid spicule is clearly lower and the ascending ramus
50 SUBFAMILIES AND GENERA OF THE SORICIDAE

noticeably broader at the level of the upper sigmoid 

notch in Chodsigoa. Hibbard (1950), p. 128) made note 

of this similarity.

Specimens examined. Chicago Natural History Museum 35760, a female C. hypsibia (deWinton) from 

Shansi, China, and 39614, a female C. salensTdi (Kastschenko) from Szechwan, China.

Chodsigoa bohlini (Young), 1934

Neomys bohlini Young, 1934, Palaeontologia sinica, ser. C, v. 8, 

no. 3, p. 16-18, pi. 1, figs. 9-11.

Pei (1936, p. 18-19) stressed the point that this fossil 

differs from Neomys in that pigment is found only on 

the lower incisor and antemolars and not on the lower 

molars and that the talonid of Ms "is reduced to one 

single pointed cusp." From Pei's illustrations the lower 

condyle appears too far forward, the external temporal 

fossa appears much too low, and the coronoid spicule 

also appears too low to be Neomys. The Mz appears 

to match that of Chodsigoa hypsibia. All these features are those that separate Chodsigoa from Neomys, 

and therefore the species is here placed in Chodsigoa. 

The coronoid spicule is too low, the paraconid-metaconid 

distance on MI appears to be too great, and the geographic distribution does not indicate that it should 

be classified as Notiosorex. The species itself could 

stand on the basis of the narrowness of the ascending 

ramus at the level of the upper sigmoid notch. The 

questionably associated upper dentition (figured by 

Pei) seems to be indeterminate; it does not seem to 

belong to either Chodsigoa or Neomys because of the

lack of pigment and the minimal emargination of M1 . 

Kowalski (1958, p. 14) also noted that this species is 

not Neomys.

Pleistocene of China (Choukoutien, loc. 3). No 

specimen was seen.

Genus BEREMENTIA Kormos, 1934 

Figure 35

Genotype: Crossopiis fissidens Petenyi, 1864 

Dental formula 1-5-3/1-2-3; upper incisor bifid; 

third upper antemolar small, fourth very small; P4 

about as in SoricuLus with parastyle placed somewhat 

farther forward than in Neomys, and posterior emargination moderate to strong; lower incisor without irregularities on its cutting edge; P± with somewhat 

inflated cingulum; Mt stout with complete and inflated 

labial cingulum, entoconid crest high and prominent; 

MZ with unreduced trigonid but talonid very reduced, 

lacking entoconid and entoconid crest as in Chodsigoa 

hypsibia', teeth with dark red to black pigment about 

to same extent as Blarinella, about equal in area to 

Soriculus or N&ctogale but much darker in color; upper 

articular facet oval; lower facet placed farther forward 

than in any other genus of Neomyini except Anouro^- 

sorex and its allies and not visible in labial view; interarticular area very broad for Neomyini and very similar 

to Blarina, except superior pterygoid fossa is a deep 

pit and has not filled in with bone to thicken the condyloid process as in Blarina ; coronoid spicule high; external temporal fossa moderately low below upper sigmoid notch and meets superior condyle; anterior curve

Medial

composite of 

SUUG, OF 65563 

and SUUG, OF 65564

FIGURE 35. Beremendia fissidens (Petenyi), SUtJG, OF 65563, 65564, and 732818. 

Right mandible and dentition and left upper dentition.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 51

on tip of coronoid process extreme. Late Pliocene to 

middle Pleistocene of Europe (where it is a very common fossil) and probably Pleistocene of China.

In several features, Beremendia closely parallels 

Blarma and Blarinoides. The stout teeth with very 

dark pigment, the large size, and the articular structures of the mandible seem to show, at first glance, a 

very close relationship between these genera. The bifid 

upper incisor, the entoconid crest on M^ anterior flex 

of the tip of the coronoid, and even details of the condylar structure clearly indicate, however, that Beremendia belongs in the tribe Neomyini. Three species 

have been named:

Beremendia flssidens (PetSnyi), 1864. Europe. The following 

specimens have been examined: Central Geological Survey 

of Czechoslovakia, and StJtJG, OF 732815, 732816, 732817, 

and 732818, all from west Slovakia, and StJtJG, OF 65563 

and 65564 from south Slovakia. These specimens include 

most of the upper and lower dentition with complete mandibles.

Beremendia sinensis (Zdanski), 1928. Choukoutien fauna of 

China (generic assignment by Kretzoi, 1956). No specimen 

was seen.

Beremendia ucrainica (Pidoplichko), 1953. Middle Pleistocene of Europe (generic assignment by Mikl6s Kretzoi, oral 

commun., 1966). No description or specimen seen.

Genus NESIOTITES Bate, 1945

Genotype: Nesiotites hidalgo Bate, 1945

Dental formula 1-5-3/1-2-3 or 1^-3/1-2-3; upper

incisor bifid; fourth upper antemolar minute when

present; P4 and M1 with moderate emargination of

posterior basal outline; lower incisor with one or two

low cusp on cutting edge; P4 paralleling that in Limnoecinae with labial blade centrally located on tooth, 

posterolingual basin present; teeth with or without pigment; articular structure of mandible as in Neomys; 

coronoid process relatively low and neither spatulate nor 

curved forward.

No specimen of this genus has been seen, and this 

diagnosis is based upon Bate's description and illustrations but is not the same as hers. Her diagnosis also 

included the following: Extinct; medium size; deep 

snout; P4 and M1 ' 3 as in Sorex, and M3 larger than in 

Recent Chimarrogale or Soriculus. She made no note 

of the conspicuously less emargination of the posterior 

border of P4 and M1 or of the relatively great lingual 

shift of the protocone of P4 , in comparison with Sorex, 

that seems to be indicated in her figure Tb.

Three species are described, all from the Pleistocene 

of the Mediterranean Islands:

Nesiotites hidalgo Bate, 1945. Mallorca and Minorca. 

Nesiotites corsicanus Bate, 1945. Corsica. 

Nesiotites similis (Hensel), 1855. Sardinia.

Genus NECTOGALE Milne-Edwards, 1870 

Figure 36

Genotype: Nectogale elegans Milne-Edwards, 1870 

Dental formula 1-4-3/1-2-3; upper incisor bifid; 

P4 with protocone fairly close to parastyle; lower teeth 

as in Neomys except stouter cuspids and secondary 

cuspids present on P4 and M±; no cuspids on cutting 

edge of lower incisor; labial cingulum absent on P4 

and present only at base of paraconid blade of lower

FIGURE 36. Nectogale elegans Milne-Edwards, China, USNM 254812. Left mandible and dentition and left upper dentition.
52 SUBFAMILIES AND GENERA OF THE SORICIDAE

molars; entoconid crest continuous between entoconid 

and metaconid on molars; pigmented enamel extreme 

in extent, covering about three-fourths of labial side 

of M\, but very pale flesh color that contrasts very little 

with unpigmented enamel pigment more obvious in 

ultraviolet light; mandibular condyles as in Neomys 

except lower articular facet not as long.

This genus is unique in the great extent of its very 

light pigment, equaled in extent by Beremendia, and 

the accessory cuspids of the P± and lower molars; 

Nectogale, Nettiotites, and Hesperosoresc appear to be 

the only genera in the tribe that do not exhibit either 

a spatulate or anteriorly deflected tip to the coronoid 

process. This is the first genus of the Neomyini discussed which shows a reduced labial cingulum, a feature characteristic of Anourosorex and its allies. There 

is no fossil record of this genus and only one living 

species, Nectogate elegans from eastern Asia, according 

to Ellerman and Morrison-Scott (1951, p. 88).

The genus has been considered a crocidurine shrew.

Specimen examined. U.S. National Museum 254812, 

a male from Szechwan, China.

Genus SORICULUS Blyth, 1854 

Figure 37

Genotype: Cosira nigrescens Gray, 1842 

Dental formula 1-5-3/1-2-3; upper incisor bifid; 

fourth upper antemolar small; P* with protocone 

shifted medially and parastyle shifted anteriorly away 

from paracone to lengthen tooth, posterior emargination moderate; lower incisor with one cusp on cutting

edge; P4 elongate with labial cingulum; labial cingulum of lower molars reduced, moderately strong 

below parastylid, weak below protoconid, and absent 

posterior to protoconid; entoconid crest very low on 

MI but still present (in contrast to tribe Blarinini); 

entire M3 somewhat reduced and talonid further 

reduced as in pattern of Blarina; pigment varying 

from somewhat more extensive than in Neomys to 

undetectable but when present light orange (as in 

Domnina and some other ancient shrews) rather than 

dark reddish brown as in Sorex and Blarina\ no 

pterygoid spicule or boss in specimens seen and superior pterygoid fossa a shallow basin because of 

thickening of condylar process; upper articular facet 

of mandible oval; lower facet barely visible behind 

lower sigmoid notch in labial view; coronoid spicule 

prominent and low on ascending ramus; external 

temporal fossa about as in Chodsigoa but with deep 

basin beneath coronoid spicule; tip of coronoid process 

broadly spatulate with little to no anterior deflection. 

Recent of eastern Asia.

Ellerman and Morrison-Scott (1951, p. 58) recognized only one living species that would be in this 

genus as recognized here. In addition to a reduction 

of the labial cingulum of the lower molars, this genus 

is the first discussed in the tribe Neomyini to show a 

significant reduction of the entire Ms and of the entoconid crest on MI and is the first discussed to show a 

strongly spatulate tip of the coronoid process; all 

these trends reach maximum development in Anourosorex and its allies. There is no known fossil species

FIGUKE 37. Soriculus nigrescens (Gray), India, CNHM 82563 (teeth) and 82565 

(mandible). Left mandible, right lower dentition, and left upper dentition.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 53

Lingual and anterior views of M,

FIGURE 38. Chimarrogale himalayica (Gray), India, UCMVZ 119400. Right 

mandible and lower dentition and left upper dentition.

which could be assigned to this genus as here understood.

/Specimens examined. Soriculus nigrescens (Gray), 

Chicago Natural History Museum 82363, a female 

from Lachen, India; 82564, a male from Lachen, 

India; and 82565, a female from Ghoom, India. These 

specimens represent two subspecies.

Genus CHIMARROGAIE Anderson, 1877 

Figure 38

Genotype: Crossapus Mmalayanus Gray, 1842 

Dental formula 1^-3/1-2-3; upper incisor sometimes bifid; P4 with protocone placed far medially 

from parastyle, posterior emargination strong; M2 

trapezoidal; lower incisor with smooth cutting edge 

but with raised crest of edge in position of cusp of 

N&omys] P± with weak labial cingulum and not elongate; Ml with moderate labial cingulum below paraconid but very thin and weak, almost unnoticeable, 

posterior to protoconid and not present on posterior 

face of tooth by hypoconid; entoconid crest very low 

on Ml about as in KoriculAjbs; entire Mz reduced and 

talonid further reduced to a Blarina-\ik& crescent; 

teeth rather stout and pigment not visible under 

normal light but under either long- or short-wave 

ultraviolet light pigment can be seen to be somewhat 

less extensive than in Neomys; zygomatic process of 

maxillary originates opposite metacone of M^ as in 

Blarinini ; area of insertion of superior pterygoid 

muscle not basined at. all but somewhat swollen 

because of thickening of condylar process (this feature is unique in the Neomyini although approached 

by Soriculus) ; pterygoid boss moderately prominent 

just below upper sigmoid notch; upper articular facet 

oval; lower facet slightly visible in labial view but 

offset lingually from lower sigmoid notch by groove 

as in CJiodsigoa, Notiosorex, and Megasorex; coronoid 

spicule slight and very high on process; external 

temporal fossa fairly low, dropping to about midway between condyles with a relatively deep basin in 

lower part opposite pterygoid insertion on medial surface and partly responsible for lack of basined pterygoid fossa; tip of coronoid process small and knoblike, 

with moderate anterior curve. Kecent of eastern Asia.

Ellerman and Morrison-Scott (1951, p. 87-88) listed 

one living species, CMmarro<gale platycephala (Temminck), 1842, which is possibly better known as 

Chimarrogale himalayica (Gray), 1842. There is no 

known fossil species that could belong to this genus. 

The genus has been considered a crocidurine shrew.

Specimens examined. University of California Museum of Vertebrate Zoology 119400, a male from 

Nepal, and U.S. National Museum 240167, from Yunnan, China.

Genus ANOUROSOBEX Milne-Edwards, 1870 

Figure 39

Genotype: Anourosorex squamipes

Milne-Edwards, 1872

Dental formula 1-3-3/1-2-3; upper incisor not bified 

in specimen seen; first upper antemolar larger than 

second; P4 with protocone shifted medially and para-
54 SUBFAMILIES AND GENERA OF THE SORICIDAE

Medial

FIGURE 39. Anourosorex squamipes Milne-Edwards, China, CNHM 36970. Left 

mandible and lower dentition and left upper dentition.

style subdued, consequently anterior and medial basal 

outlines form approximately a right angle, emargination of posterior basal outline very slight; M1 with 

exaggerated parastyle and subdued mesostyle; M2 

reduced and almost triangular; MB minute; lower 

incisor with smooth or very slightly irregular upper 

cutting edge; M1 with exceptionally elongate trigonid, 

entoconid crest lacking, metalophid-protolophid junction located very labially and comparable to Blarina, 

labial cingulum totally lacking as on all lower teeth; 

Mz reduced; Ms minute with very slight talonid remaining; teeth unpigmented and bulbous; zygomatic 

process of maxillary originates opposite M2 ', mental 

foramen in depression below middle of M^ coronoid 

process broad and spatulate, very broad at level of 

upper condyle; coronoid spicule strong and fairly low; 

external temporal fossa of mandible low, extending 

about to level of lower condyle; superior pterygoid 

fossa of mandible as deep basin with prominent pterygoid spicule; upper articular facet triangular as in 

Chodsigoa; interarticular area very narrow; lower 

condyle placed well forward of lower sigmoid notch. 

Middle Pliocene to Eecent, eastern Asia.

In dental and mandible structures, this is the most 

specialized of living shrews. Nectogale, Soricuhis, 

and Chimarrogale all have features that suggest a 

relationship to Anourosorex, but it does not seem possible to decide which is most closely related from the 

characters considered in this review. Although all 

these are eastern Asian shrews of the modern fauna, 

close relatives are also found in the Pliocene record of

North America (J. H. Hutchison, written commun. 

1964) and Europe.

Ellerman and Morrison-Scott (1951, p. 87) recognized only one species, Anourosorex squamipes MilneEdwards, although some authors appear to still consider A. assamensis Anderson as a distinct species.

Specimen examined. Chicago Natural History Museum 36970, a male from Szechwan, China.

There are two fossil species included in the genus:

Anourosorex inexpectatus (Schlosser), 1924. Placed in the 

genus Neomys by Schlosser and compared with Beremendia 

flssidens from Europe. As Miller (1927, p. 10) noted, the 

species agrees well with living Anourosorex. From Schlosser's illustration (1924, pi. 1, fig. 4), the presence of Ms is 

indicated by its alveolus and thus eliminates assignment of 

the species to the genus AmblycopPus. Middle Pliocene of 

Mongolia (Ertemte fauna). No specimen was seen.

Anourosorex japonicus Shikama and Hasegawa, 1958. Mz 

triangular; M3 and Ms further reduced than in living species. Middle Pleistocene of Japan (Siraiwa fauna). No 

specimen was seen.

Genus AMBLYCOPTTTS Kormos, 1926 

Genotype: Amblycoptus oligodon Kormos, 1962 

Dental formula 1-1-2/1-2-2; differs from Anourosorex in the following characteristics: One more upper 

antemolar; M1 with greater emphasis of parastyle; 

M2 completely reduced to triangle and resembling M3 

of other shrews; M\ missing; jaw stouter; interarticular area somewhat broader with oval upper 

articular facet. Early Pliocene (Polgardi fissureHipparion fauna), Hungary. One species named,
DIAGNOSES AND CONTENTS OF SUBFAMILIES 55

Amblycoptus oligodon Kormos. No specimen was 

seen.

Genus HESPEROSOREX Hibbard, 1957

Genotype: Hesperosorex l&vei Hibbard, 1957 

Labial cingulum of M± prominent, somewhat inflated, not deepened, and continuous around hypoconid 

to posterior border of tooth; entoconid of MI prominent, close to metaconid (closer than on M2 ) and connected to metaconid by an entoconid crest of moderate 

height; hypolophid of M± (inadvertently called the 

paraconid by Hibbard) well developed and extending 

past the entoconid to the lingual cingulum (not so 

extensive lingually on M2 to judge from Hibbard's 

figure); MI and M2 stout and "tips of the molars show 

evidence of pigmentation," suggesting noticeably less 

pigmentation than in Blarina; mental foramen below 

middle of M^ coronoid spicule 5 not well developed 

and fairly low, as in Chodsigoa; tip of coronoid 

process narrow, not curved anteriorly and not spatulate; articular structures most generalized of tribe; 

upper facet narrowly oval; lower facet broadly oval 

and placed only moderately forward but offset far 

lingually (characteristic of Neomyini) and separated 

from lower sigmoid notch by a groove; interarticular 

area broad relative to all other genera of Neomyini 

except Beremendia and comparable in general development to specialized Soricini as Blarinella, or generalized Blarinini as Cryptotis and Adeloblfirina,. Middle 

Pliocene (Hemphillian) of North America.

This genus is somewhat questionably referred to the 

Neomyini because the genotypic specimen, a mandible 

with MI and M2 , was not available for examination. 

Classification with the Soricini seems to be eliminated 

by the condyloid and coronoid structures but not by 

great or obvious differences. The tribe Blarinini is 

eliminated by the entoconid crest of M± and by the 

extreme offset of the lower condyle. The absence of a 

lingual emargination of the interarticular area is the 

chief objection to placing Hesperosorex in the tribe 

Neomyini, but this objection seems to be outweighted 

by the distinctive lingual offset of the lower condyle 

and the resulting groove between the condyle and the 

lower sigmoid notch. The absence of the emargination 

of the interarticular area seems to be due to a minimal 

lingual shift of the superior condyle resulting from 

minimal curvature toward the cranium of the condyloid process supporting the superior condyle and thus 

suggests that the coronoid process is not deflected 

labially as in more typical members of the Neomyini. 

If these suggestions are true, the mandibular struc5 As here used; Hibbard reserved the term, possibly correctly, in 

the sense used by Gaugrahn, for a needlelike projection rather than a 

ridge.

tures could represent a primitive condition, not far 

removed from the Soricini, that could be expected in 

the Neomyini. These suggestions, however, cannot be 

evaluated from the published descriptions.

The genotypic species, Hesperosorex lo'vei Hibbard, 

is from the middle Pliocene Teewinot Formation of 

Wyoming. No specimen was seen.

Genus NOTIOSOREX Baird, 1877

Figure 40

Genotype: Sorex (Notiosorex} crawfordi Coues, 1877 

Dental formula l-4r-3/1-2-3; upper incisor not 

bifid; P4 with protocone about as in Sorex, posterior 

emargination strong to extreme; M2 rectangular; lower 

incisor short with one low cusp on cutting edge; P4 

with curving junction of labial blade and crest of 

posterior cingulum; Mi with metaconid and paraconid 

close together relative to Chodsigoa, labial cingulum 

strong, entoconid and entoconid crest strong; J/3 with 

reduced heel, entoconid lost, entoconid crest suppressed, V of hypoconid not developed, and metalophid 

and hypoconid form curved trenchant crest; tip of 

protoconid of M^ of paracone of P*, and tips of more 

anterior teeth pigmented; teeth slender; zygomatic 

process of maxillary originates opposite metastyle of 

M2 ; mental foramen below middle of M\\ tip of 

coronoid process narrow and inclined slightly forward; 

coronoid spicule moderately low; external temporal 

fossa of mandible moderately low and extending down 

to about midway between condyles; superior pterygoid 

fossa of mandible deeply basined, pterygoid spicule 

strong or barlike; condyloid structure as in Chodsigoa 

with triangular upper facet and groove labial to 

lower facet. Late Pliocene (early Blancan) to Recent, 

North America.

One living species, Notiosorex crawfordi (Coues), is 

recognized and is also known from the late Pleistocene 

of Kansas and California. The one extinct species, 

Notiosorex jacksoni Hibbard, is known from the late 

Pliocene (early Blancan) of Kansas (Hibbard, 1950) 

and possibly from the middle Pleistocene (Irvingtonian) of California (Vallecito Creek fauna, unpub. 

record, J. A. White, oral commun., 1963) and differs 

in being intermediate in size, bet ween living Notiosorex 

and Megasorex and in having extreme emargination 

of the posterior basal outline of P4 and the upper 

molars, thus resembling Chodsigoa salenskii.

Specimens examined. Four specimens of the living 

species from Texas and California in the University 

of California Museum of Vertebrate Zoology; one 

specimen possibly of N. jacksoni from the Pleistocene 

of California in the Los Angeles County Museum.
56 SUBFAMILIES AND GENERA OF THE SORICIDAE

FIGURE 40. Notiosorex crawfordi (Coues), Texas, UCMVZ 98040. 

and dentition and left upper dentition.

Left mandible

FIGURE 41. Megasorex gigas (Merriam), Mexico, LACM 13568. Left mandible 

and dentition and left upper dentition.

Genus MEGASOREX Hibbard, 1950 

Figure 41

Genotype: Notiosorex gigas Merriam, 1897 

Dental formula 1-4-3/1-2-3; upper incisor not 

bifid; P4 about as in Notiosorex but with only slight 

posterior emargination; M2 rectangular; lower incisor 

as in Notiosorex', P4 with more angulate junction of 

labial shearing blade with crest of posterior cingulum 

than in Notiosorex and more like Chodsigoa salenskii;

entoconid crest of M\ very low; talonid of Ms little 

reduced and as in Chodsigoa scdenskii; zygomatic 

process of maxillary originates behind J/2 ; mandibular 

structures virtually as in Notiosorex and Chodsigoa 

except tip of coronoid more inclined anteriorly and 

more spatulate than in Notiosorex; one living species, 

Megasorex gigas (Merriam), Recent of Mexico.

Specimen examined. Los Angeles County Museum 

13568, a female from Colima, Mexico.
DIAGNOSES AND CONTENTS OF SUBFAMILIES 57

Neomyini? Incertae Sedis 

Deinsdorfia franconia Heller, 1963

This genus and species was defined (Heller, 1963, 

p. 5-7) on the basis of a toothless right mandible from 

the Pleistocene fauna of Deinsdorf. In Heller's 

photographs it is characterized by rather extreme 

posterior placement of the mandibular condyles but 

otherwise appears not unlike Neomys. Heller's (fig. 

2c) drawing of the posterior view of the condyles, 

however, does not appear similar to Neomys at all. 

Fejfar (1966, p. 243) interpreted Heller's drawing as 

indicative of the tribe Soricini, but I have some question about the accuracy of the drawing. If the drawing is correct then I suspect that Deinsdorfia belongs 

in the Blarinini. In any event, I am unable to place 

the genus in any soricid tribe from published descriptions but feel that it most likely belongs in the 

Neomyini.

Subfamily ALLOSORICINAE Fejfar, 1966

This subfamily was recently established by Fejfar to 

include a most unusual shrew, Allosorex stenodus. In 

his discussion of this shrew, Fejfar (1966, particularly 

p. 240-241) pointed out the similarities of Allosorex to 

"Sorex" gracilidens and suggested that the latter could 

well be the late Miocene ancestor of Allosorex. He did 

not, however, include "/Sorex" gracilidens in his new 

subfamily. To me, the similarities are strong, however, and, being faced with a problem of subfamily 

assignment that Fejfar did not have, I include "Sorex" 

gracilidens in the family Allosoricinae, rather than 

in "Soricidae, Incertae Sedis," for it clearly does not 

belong in any other subfamily of shrews. The following diagnosis of the subfamily Allosoricinae is revised 

from Fejfar (p. 223) with modifications to allow inclusion of "Sorex" gracilidens.

Lower antemolars high cusped with conspicuous 

sectorial adaptation, P4 apparently derived from a 

crocidurinelike triangular crown but a clear tendency 

to form a high sharp anteroposterior shearing blade 

with no overhang of alveoli by enamel cap; lower 

molars narrow and high cusped, very carnivorelike, 

trigonid much longer than talonid, paralophid blade 

knife edged and conspicuously oriented anteroposteriorly in comparison to most shrews, metaconid close to 

and much lower than protoconid, metalophicl united 

to protoconid and forming a continuous crest with 

hypolophid, hypolophid united with entoconid, entoconid crest absent and talonid basin open lingually 

between entoconid and metaconid particularly in Allosorex, cingulum weak to not present; coronoid process 

low and anteroposteriorly wide; mandibular condyles 

separated and upper one placed far posteriorad; upper

incisor fissident, teeth not pigmented. The unpigmented, carnivorelike lower teeth are most characteristic of this subfamily.

"Sorex" gracilidens Viret and Zapfe, 1951

P4 with single, narrow, high cusp that has narrowly 

triangular occlusal section surrounded by a basal 

cingulum, anteroposteriorly two rooted; MI very suggestive of F4 in Martes: narrow, paralophid nearly 

parallel to axis of mandible, metaconid reduced and 

fused to protoconid; metalophid nearly parallel to 

axis of mandible, hypoconid and entoconid strong 

and forming square corners on posterior border of 

short talonid; MB also similar but reduced and talonid 

reduced to rounded heel apparently with only one 

cusp; mental foramen below P4 , internal temporal 

fossa well developed; upper and lower articular facets 

clearly formed and fairly widely separated, but there 

appears to be no interarticular excavation. Late Miocene (Vindobonian) of Vienna Basin.

From published descriptions, this shrew does not 

clearly belong to any subfamily except the Allosoricinae. No specimen was seen. Fejfar (1966, p. 240- 

241, figs. 13, 14) described new material and discussed 

the possible ancestral position of "Sorex" gracilidens 

to the genus Allosorex.

Genus ALLOSOREX Fejfar, 1966

Genotype: Allosorex stenodus Fejfar, 1966 

Large; dental formula l-?-?/l-2-3; sectorial development of cheek teeth extreme, paralophid and 

metalophid parallel to axis of mandible; lower incisor 

with hooked tip and trough along lingual side of 

cutting edge; upper incisor very weakly fissident; 

coronoid process very low; condyles very large and 

widely separated so that they ar.e not connected by an 

interarticular area. Late Pliocene, Czechoslovakia.

This genus is, easily, the most specialized and unusual shrew known.

Specimen seen. STOG, OF 652366, a toothless 

mandible, and STOG, OF 652370, a mandibular fragment with ./I/i, both topotypes from Ivanovce, near 

Trencin, Czechoslovakia.

SORICIDAE Incertae Sedis 

Podihik kura Deraniyagala, 1958

This living shrew was defined on the basis of two 

specimens from Ceylon and was considered to be a 

soricine shrew because the teeth were pigmented pale 

reddish gray. I am informed by Dr. Deraniyagala 

(written commun., 1965), however, that the coloration 

of the teeth has entirely disappeared in both preserved 

specimens. This suggests to me that the reddish color 

was a stain rather than actual tooth pigment, which
58 SUBFAMILIES AND GENERA OF THE SORICIDAE

is very stable. From published descriptions and 

supplemental notes provided by Dr. Deraniyagala I 

am unable to assign this shrew to a subfamily. If it is 

a soricine shrew, its occurrence would be most unusual, 

and the only one in peninsular India or Ceylon and 

the only one south of the ancient Tethys seaway.

Names of fossil genera not used in text

Allopachyura Kormos, 1934 ; included in Petenyiella.

Asoriculus Kretzoi, 1959 ; included in Episoriculus.

Heterosorex Gaillard, 1915 ; included in Trimylus.

Miothen Cope, 1873; included in Domnina.

Oligosorex Kretzoi, 1959 (Oligosorex meyeri) ; possibly valid;

in Crocidosorex ( Crocidosorex antiquus ). 

Plesiosorex Pomel, 1848, not a soricid. 

Protosorex Scott, 1895; included in Domnina.

ANCESTRY OF THE SORICIDAE

In this review it has been shown that the Soricidae 

is a family of diverse phylogeny and that subfamilies 

can be recognized by consistent morphologic differences which are traceable back through the late and 

middle Tertiary toward a common ancestry. The 

family is, however, distinctly differentiated from all 

other mammals. There is no shrew known that is 

sufficiently generalized morphologically to be this common ancestor. The dental and mandibular characters 

of the oldest known member of each of the four subfamilies and the directions of modification of these 

characters in younger forms suggest what this original shrew should look like. Its diagnosis may be 

about as follows:

Dental formula 1-6-3/1-5-3; upper incisor falciform and bifid; upper antemolars except P4 small, 

simple in form, equal in size or anterior ones larger, 

only slightly crowded; P4 nearly equilaterally triangular in occlusal view, protocone close to anterior 

apex of triangle and close to incipient parastyle at 

anterior apex, posterior emargination very slight or 

lacking; J/1 square in occlusal view with W-shaped 

ectoloph, protocone strong, hypoconal shelf and cingulum strong, emargination of posterior outline slight 

or lacking, no protoconule or metaconule; M2 square 

and similar to Ml ; M3 triangular, metastyle and hypoconal shelf lacking; lower incisor large with extreme 

procumbency typical of Soricidae, possibly with two 

rounded cuspules on upper cutting edge; lower antemolars with a single triangular cusp, the first and last 

(P4 ) largest, only slight tendency to be crowded; 

P4 not even slightly molariform and clearly single 

cusped, posterior and lateral cingula prominent, two 

rooted; MI with trigonid and talonid well developed, 

metaconid and paraconid more widely separated than 

on M2 , metalophid meeting protolophid at midline of

tooth or possibly closer to the metaconid, entoconid 

clearly separated from hypolophid and connected to 

metaconid by a high and well-formed entoconid crest, 

no hypoconulid, labial cingulum probably strong, labial reentrant between the protoconid and hypoconid 

emerges at level of cingulum; M2 like MI but paraconid closer to metaconid; M3 with trigonid like that 

on Ma but talonid reduced (to condition as in Sorex}, 

entoconid connected to hypolophid; all teeth pigmented light orange; zygomatic process of maxillary 

originates behind M2 ] zygomatic arch probably incomplete but maxillary process strong; mental foramen below P± or /*3 ; internal temporal fossa of mandible strong but not pocketed; external temporal 

fossa of mandible weak and confined to tip of coronoid process, its posterior margin above upper sigmoid notch; masseteric fossa of mandible moderately 

strong, insertion on angular process marked by crest; 

superior pterygoid fossa of mandible on anterior side 

of condylar process; condylar process delicate; condyle small and single, articular facet probably covers dorsal, posterior, and ventral sides of condyle to 

fit into pocketed glenoid; small size.

This hypothetical ancestor would unquestionably 

resemble a shrew despite the lack of the pocketed internal temporal fossa, of the doubled mandibular condyle, of the emargination of the posterior border of 

P* and the upper molars, and of the reduced number 

of antemolars. The characters that would make it 

appear shrewlike, together with the occurrence of these 

characters in mammals with similar dentition, are as 

follows:

1. Procumbent and enlarged lower incisor. Perhaps 

this is the most typically soricid feature. It is 

shared with no similar mammal.

2. Falciform upper incisor. This is shared to a slight 

degree by some erinaceoids, as Saturninia, but still 

is a typical soricid feature. In connection with this 

tooth, it should be noted that the bifid upper incisor 

found in the Heterosoricinae, in the tribe Neomyini, 

and in some members of the tribe Soricini is also 

found in some Chiroptera but has not been noted 

in any other mammal with closely comparable dentition.

3. Entoconid separated from the hypolophid (except 

on M3 ). This is a typical soricid feature but is 

found in a few Chiroptera and also in erinaceoids 

when the hypoconulid breaks the continuity of the 

hypolophid. In the erinaceoids, however, the presence of the hypoconulid is a far greater dissimilarity 

than the separation of the entoconid is a similarity 

to the shrews.
PALEOZOOGEOGRAPHY AND CORRELATION 59

4. Entoconid crest on MI (and M2). This is a typical 

soricid feature but is shared with most Chiroptera 

and with Nesophontes.

5. A strong hypoconal shelf and cingulum depressed 

from the level of protocone and no distinct hypocone on M1 and M2. This is a typical soricid feature that is shared in typical development only by 

the Chiroptera. The Talpidae either lack hypoconal structures or have a hypoconal shelf that is on 

a level with the protocone and has a fairly distinct 

hypocone. The same can be said for most forms included in the Erinaceoidea.

6. Protoconule and hypoconule absent. This is a 

character that is shared with the Chiroptera, Talpidae, and Nesophontes.

7. W-shaped ectoloph. This is a character that is 

shared with the Chiroptera, Talpidae, and perhaps 

Nesophontes, as well as many other dialambdont 

insectivores.

8. M3 reduced by loss of metastyle and hypoconal 

structures. This is a character shared with the 

Chiroptera and Talpidae but not with typical Erinaceidae.

9. P4 not molariform. This is a character shared 

with the Chiroptera, Talpidae, and Nesophontes.

10. No hypoconulid on MI and M2 . This is a character shared with the Chiroptera and Talpidae, as 

well as with some erinaceoids and possibly Nesophontes.

11. Zygomatic arch (or process of the maxillary) originates behind M2 . This is a character shared with 

the Chiroptera and the Talpidae.

12. Metalophid-protolophid junction near the metaconid of MI and M2 . This is a character shared 

and exceeded by the Talpidae and shared by the 

Chiroptera.

13. Metaconid farther from paraconid on MI than on 

M2 . This character is found in some forms of most 

insectivore groups. It is mentioned here because it 

has not been seen in the Chiroptera during this 

review.

Although it represents only an approximation based 

upon the inferred dental characters of the hypothetical 

ancestor of the four subfamilies of the Soricidae, it is 

interesting to note the relative degree of resem^ance 

of the shrews to other insectivores and bats. Of the 

13 characters listed as shrewlike, the Chiroptera share 

11, the Talpidae share 7, the Erinaceoids share 4+, 

and the problematic sub-Recent genus Nesophontes 

from the West Indies shares 4+. Of course consideration of other morphologic characters, as in the 

forelimbs of bats, may outweigh dental similarities.

It appears, therefore, that Soricidae were certainly 

represented in the Oligocene by the subfamilies Heterosoricinae and Soricinae. The subfamily Crocidurinae and the Limnoecinae have features that could 

not have been entirely derived from either the Soriciniae or the Heterosoricinae, as these subfamilies are 

here understood from their earliest known records ; the features appear to have been inherited from' 

an ancestral form possessing some characters of both 

Oligocene subfamilies. Therefore it has to be assumed that the Crocidurinae and Limnoecinae were 

also present in the Oligocene or that an unknown 

group was present from which both diverged. The 

geographic isolation of these two subfamilies, the 

Crocidurinae in the Old World and perhaps only 

south of the Tethys seaway before the Miocene (Repenning, 1965, p. 195) and the Limnoecinae in the 

New World after the early Miocene, suggests that they 

were derived from the native soricid stock sometime 

before the earliest record of the soricines. This would 

certainly be in the Oligocene for the origin of the 

Crocidurinae of the Old World and probably in the 

Oligocene for the origin of the Limnoecinae in the 

New World.

The Oligocene subfamilies of the Soricidae diverged 

from an unknown, preearly Oligocene, very shrewlike soricid ancestor. This common ancestor appears 

to have evolved, along with the Chiroptera and the 

Talpidae, from an unknown soricoid. If one assumes 

that the slow rate of morphologic change evidenced 

in the middle to late Tertiary fossil record of the 

shrews reflects earlier rates of change, divergence of 

these groups from the unknown soricoid must have 

taken place before the Eocene. It would seem the 

Soricoidea-Chiroptera stock could not have diverged 

from the remaining insectivores after these had a 

molariform P4 , and this may well take the ancestry 

of the Soricoidea (at this point including the Chiroptera) back to the Mesozoic and possibly to the origins 

of placental mammals.

PALEOZOOGEOGRAPHY AND CORRELATION

Before the late Pliocene the fossil record of the 

shrews is quite inadequate. Even after that time the 

known specimens are not abundant except from a few 

localities where either the manner of fossil accumulation or the method of collection has been favorable. 

Furthermore, most shrews are highly endemic animals not only geographically limited in both living 

and fossil species but tending to remain so throughout appreciable spans of their evolutionary histories. 

Finally, rates of evolution vary greatly as illustrated 

by the conservatism of the tribe Soricini contrasted
60 SUBFAMILIES AND GENERA OF THE SORICIDAE

with the exorbitant adaptive modification and variety seen in the tribe Neomyini, both within the subfamily Soricinae. These conditions all indicate that 

interpretation of the significance of a fossil shrew 

should be made with caution.

The small subfamily Heterosoricinae contains only 

four known genera and is characterized by little ramification of its phylogeny. Domnina°is known only 

from North America, where it ranges through the 

Oligocene and early Miocene. During this time it 

tends to increase the crowding of its antemolars and 

possibly to reduce its pigmentation, but stages of evolution cannot be clearly defined on the basis of the 

known material. Unless a specimen can be identified 

with a known species, it appears that it may be useful in establishing geologic age only in terms of 

closeness to one limit of the temporal range of the 

genus rather than to the other.

The genera Paradomnina and Ingentisoreso are 

known only from the late Miocene of North America.

Trimylus is better represented in number of species 

and in greater temporal and geographic range. In 

North America and possibly in Europe the evolution of 

this genus appears to be almost straight line with a 

uniform rate of change. It seems quite possible at 

present to recognize middle or late Oligocene and early, 

middle, and late Miocene stages of evolution in North 

American Trimylus. The possibility of similar usefulness in the European forms cannot be evaluated without examination of the specimens.

Although Trimylus is known only back to the middle 

Oligocene, it seems reasonable to assume that the genus 

became a part of the fauna of both North America and 

Europe during the early Oligocene mammalian interchange. Oligocene species show a greater similarity 

than do more recent species of both hemispheres; since 

the beginning of the Miocene, possibly since the early 

Oligocene, endemic Old and New World lineages appear to have been in existence. Trimylus roperi from 

the early Miocene of Colorado may represent an immigration of the European stock during the early Miocene 

mammalian exchange or may be a part of the American 

stock that has an otherwise unrecognized variability in 

the position of the mental foramen. The Heterosoricinae 

is unique in several features and seems much more distantly related to the other soricid subfamilies than all 

the other subfamilies are to each other.

The fossil record of the Crocidurinae is very meager. 

The oldest specimens known are of middle Miocene age 

from Europe (Doben-Florin, 1964) and Africa (Butler 

and Hop wood, 1957, p. 2). The subfamily is characterized by the retention of primitive characters, and the 

modern forms differ from those of the late Miocene in

the loss of one, two, or three upper and lower antemolars, in greater reduction of the talonid of Ms, and 

in a greater emargination of the posterior basal outline 

of P4 and M1 . In all other features, morphologic differences seem to be greater between living crocidurines 

than between living forms and late Miocene forms.

The Crocidurinae are largely tropical Old World 

forms and are the only shrews, fossil or living, known 

from Africa, peninsular India, and Ceylon. In Africa 

the subfamily shows its greatest diversity in living 

genera. The apparently late appearance of the crocidurines in the Tertiary scene in Eurasia and their obvious African affinities suggest that they may have 

evolved largely in Africa. Their distinctly soricid features, however, do not suggest that they are derived 

from independent African sources but rather that they 

are descendant from the same stem as all other shrews 

and in many features are not far removed, morphologically, from this ancestry.

Closely paralleling the Crocidurinae in lack of specialized morphology but entirely distinct in distribution, the small New World subfamily Limnoecinae is 

known by a few species from the early Miocene to the 

middle Pliocene. The earliest known species is associated with soricine shrews from which it is clearly distinct on the basis of all subfamily criteria. These 

earliest forms are clearly intermediate between the late 

Miocene to middle Pliocene Limnoecus and the hypothetical soricid ancestor. Thus, stage of evolution 

within the Limnoecinae is recognizable to the extent of 

distinguishing a late Miocene to middle Pliocene species from an early to middle Miocene one. Assuming 

comparable rates of evolution, earlier (but now unknown) forms should be equally as definitive of geologic age, although perhaps not distinguishable from 

the Crocidurinae except in their geographic distribution.

The similarity of the dental and mandibular characters of the Crocidurinae and the Limnoecinae result 

largely from the retention of those features believed 

to have been typical of the hypothetical soricid 

ancestor. It appears to be reasonable that both of 

these subfamilies evolved in isolation, removed from 

the competitive pressure of the closely related Soricinae. Although the earliest known limnoecine genus, 

Angustidens, is associated with the earliest known 

North American soricine genus, Antesorex, it seems 

likely both that the Limnoecinae had an older history 

in North America and that Antesorex represented a 

recently introduced stock closely similar to Crocidosorex of comparable and even greater age in Europe. 

Furthermore, the extinction of the North American 

Limnoecinae in the middle Pliocene was at the time
PALEOZOOGEOGRAPHY AND CORRELATION 61

CROCIDURINAE

Crocidura

Diplomesodon

Feroculus

Myosorex

Praesorex

Scutisorex

Solisorex

Suncus

Surdisorex

Sylvisorex

Crocidura 

Myosorex

Crocidura 

Suncus

Shikamainosor >,x

Myosorex 

Crocidura 

Suncus 

Diplomesodon

Blarina 

Blarinoides

\

Crocidura 

Soricella

Pigment lostto Africa

Unknown form with internal 

temporal fossa of mandible not pocketed

SORICINAE

NEOMYINI

Anourosorex 

Chimarrogale 

Chodsigoa 

Episoriculus

Nectogale 

: Neomys 

 Notiosorex 

'. Soriculus

{ .Neomys

: : Notiosorex-/

':' ' Nesiotites : 

: Notiosorex '. 

' Chodsigoa ' 

Anourosorex 

' Beremedia

Neomys 

Episoriculus '

Beremendia 

Notiosorex

Hesperosorex. 

 Anourosorex 

 ; ;. '.' '.:'. Amblycoptus[:]' ::':" .'. ' .' ' :' 

'. '. ' : ' aff. Anourosorex ; :". '. '. '/:'

-Pigment reduced or 

lost in a few genera

Bifid upper incisor 

retained in most Old World forms

Pigment reduced- to North America

Pigment retained

Unknown form with internal temporal 

fossa pocketed into mandible

Hypothetical soricide ancestor

FIGURE 42. A phylogeny of the Soricidae.
62 SUBFAMILIES AND GENERA OF THE SORICIDAE

of apparently explosive diversification of the Soricinae. This may also coincide with the approximate 

time of extinction of the Heterosoricinae in both the 

Old and New Worlds, although there is no record of 

this subfamily younger than late Miocene in North 

America. In the Old World the Crocidurinae have 

survived. With the exception of a few species of 

Suncus and Crocidura, however, the Crocidurinae of 

Africa and peninsular India and the Soricinae of 

Palearctica (excluding Mediterranean Africa and including northern India, Burma, and Indochina) are 

mutually exclusive in their ranges. Those species of 

Suncm and Crocidura which have successfully invaded 

Palearctica seem to have done so by being adaptable to 

a great diversity of environments from dry grasslands 

to damp forests and shores; many are commensal with 

humans.

The boundary between those areas occupied entirely 

by the living crocidurine shrews and those areas occupied largely by the living soricine shrews coincides 

with the position of the Miocene Tethys seaway to a 

remarkable degree and suggests an incredible degree 

of endemism with these shrews.

The Soricinae is the largest subfamily of the shrews 

and has a fossil record known from the late Oligocene 

of Europe and from the early Miocene of North 

America. The earliest members of the subfamily are 

very generalized, and this primitive condition persists 

today in Sorex and closely related genera, here included in the tribe Soricini. Two other tribal lineages 

diverge from this unspecialized stem: the tribe 

Neomyini, apparently of Old World origin and still 

largely of Old World distribution, and the tribe 

Blarinini, apparently of New World origin and now 

confined to this area. Morphologically this subfamily 

is so diverse that the stage-of-evolution concept has 

no meaning at more than tribal level and in many 

groupings is apparent only in smaller taxonomic units, 

here informally referred to as generic groups. In the 

tribe Neomyini the fossil record is inadequate to 

chronologically define any evolutionary picture although the tribe is rather well represented in the fossil 

record.

The tribe Soricini includes two generic groups: the 

genus Sorex and closely related forms and the genus 

BlarineUa with fossil genera related to it. The Sorex 

group is conspicuously unspecialized and uniquely 

successful among all shrews, as exemplified by /Sorex, 

the only living holarctic genus. Except for the Heterosoricinae, the earliest known shrews belong in the 

Sorex group by virtue of their lack of specialization.

These early soricines (Orocidosorex and Antesoreae) 

differ from the living Sorex in their greater number of 

teeth, more anterior position of the mental foramen, a 

more primitive P4 , less emargination of the posterior 

border of P4, M1 , and Mz, and in the slightly less 

separation of the articular facets on the mandibular 

condyle. These early soricines are also believed to be 

ancestral to other shrews of the Soricinae.

The Blwrinella generic group diverged from those 

with this primitive pattern presumably in the Miocene 

by enlargement of the mandibular condyle (but with 

no relative increase in separation of the facets as was 

done in other tribes), accentuation of the entoconid 

crest of MI and M2, and reduction of the talonid of 

M3 . This generic group is represented in the late 

Miocene and early Pliocene of North America, in the 

middle Pliocene of China, and the late Pliocene of 

Europe (where Petenyid continues until the middle 

Pleistocene), and now lives in the Chinese highlands.

The tribe Blarinini is first known from the late 

Miocene in the genus Adeloblarina of North America. 

This genus is not far removed from the basic stock of 

the Soricinae, which is first known in North America 

from the early Miocene, but has several features 

clearly indicative of the Blarinini. The middle Pliocene to living Cryptotis is not much advanced over this 

stage in several respects, but the living Blarina is 

greatly specialized in most characters except for its 

retention of five upper antemolars. The number of 

upper antemolars, the nature of reduction of the M$, 

and the degree of specialization of the mandibular 

articulation seem to indicate that the tribe is divisible 

into three generic groups: Two are represented by the 

living genera Blarma and Cryptotis and the third by 

the extinct Paracryptotis from the Pliocene.

Blarina and related shrews are derived from a form 

similar to Adeloblarina by retention of five upper 

unicuspids, by lack of reduction of the talonid relative 

to the trigonid of M3 (although the entire tooth may 

be reduced), and by extreme separation of the mandibular condyles. Only two genera have this characterization: Blarinoides from the late Pliocene of 

Europe and Blarina from the late Pliocene to Kecent 

of North America.

Cryptotis and Paracryptotis are derived also from 

a form similar to Adeloblarina by the retention of 

only four upper unicuspids except the earliest known 

species and by the reduction of the talonid of M3 

relative to the trigonid. In Cryptotis, reduction of 

the M3 talonid is greatest but separation of the mandibular condyles is relatively slight. The genus is
KEY TO LIVING AND FOSSIL GENERA OF THE SORICIDAE 63

known from the middle Pliocene to the Kecent in 

North America. Paracryptotis has a somewhat less 

reduced talonid on Ms but has much greater separation of the mandibular condyles, nearly as extreme as 

in Blarina, and is known from the middle and late 

Pliocene of North America. /Shikamainosoreos from 

the middle Pleistocene of Japan is very closely related 

to Paracryptotis but is somewhat more advanced in 

reduction of the talonid of J/3 and in separation of 

the mandibular condyles. "/Soreiv" kretzoii and "Sorex" 

dehneli from the late Pliocene of Poland may be related to Cryptotis or Paracryptotis, but published information is not sufficient to clearly indicate that these 

generic groups are represented in the Cenozoic of 

Europe.

Although largely a North American tribe, the Blarinini clearly were introduced once, and possibly twice, 

into the fauna of the Old World, where they do not 

seem to have been too successful. Possibly they found 

the Old World Neomyini strong competition. Cryptotis is the only shrew which has entered South America.

The tribe Neomyini is first known from the early 

Pliocene of Europe and North America. The earliest 

known European form, Amblycoptus, is the most specialized shrew of the tribe. Therefore, it appears that 

no pattern of evolutionary stages is detectable from 

the information available for this review, but more 

likely that the diversification in morphologic types of 

the tribe Neomyini was as great in the early Pliocene 

as it is today.

The tribe is also represented in North America by 

Hesperosorex from the middle Pliocene, by Notiosorex 

from the late Pliocene to the Kecent, and by the living Megasorex. These genera differ from most of the 

Old World Neomyini in their lack of a bifid upper 

incisor. Inasmuch as the bifid falciform upper incisor 

is also found in the Heterosoricinae, it is considered 

to be a primitive feature inherited from a common 

ancestor. It therefore seems more reasonable to postulate that the Eurasian Neomyini, with its great diversity of forms, represents the parental stock and that a 

few forms entered North America before the early 

Pliocene and were also separated from the Eurasian 

population before that time.

In establishing the subfamily Heterosoricinae, Viret 

and Zapfe (1951, p. 426) suggested that the genus 

Amblycoptus may belong to that subfamily because 

of mandibular similarities. Although clearly not a 

heterosoricine shrew, Amblycoptus and fossil and living Anourosorex do closely parallel Trimylus in the 

massive masticatory structures. The suggested similarity of habits and the presence of this specialized 

member of the Neomyini in the lower Pliocene of the 

Old World, at the approximate time of the extinction 

of Trimylus, seems to provide a plausible explanation 

for the extinction of the Heterosoricinae in that area. 

Within the Crocidurinae the extinct Diplomesodon 

fossorius and to some extent the living Solisorex have 

also made comparable adaptations of their masticatory structures.

In summary, the shrews are useful in correlation of 

geologic age, but the degree of their usefulness depends on the lineage represented and on the geographic considerations involved in such correlation. 

The subfamily Limnoecinae clearly originated in 

North America. The subfamily Crocidurinae almost 

certainly is of Indo-African origin although it must 

have been derived from a soricid stock of holarctic 

distribution which was also the source of the Limnoecinae and the Soricinae. The large subfamily Soricinae seems to be of Eurasian origin although diversification of the subfamily into tribes took place 

throughout Holarctica: the tribe Blarinini of New 

World origin, the tribe of Neomyini of Old World 

origin, and the tribe Soricini representing the primitive Old World stock.

The Soricidae offer, with varying certainty, evidence 

of intercontinental faunal exchanges. Tentatively, 

these may be listed as follows:

1. Late Eocene or earliest Oligocene. Dispersal of 

Trimylus between North America and Eurasia; possible dispersal of ancestral Limnoecinae from Eurasia to North America; possible dispersal of ancestral Crocidurinae from Eurasia to Africa.

2. Early Miocene. Dispersal of Soricinae from Eurasia to North America; possible dispersal of Trimylus to North America.

3. Middle Miocene. Dispersal of some Crocidurinae 

from Africa to Eurasia.

4. Late(?) Miocene. Dispersal of Neomyini from 

Eurasia to North America.

5. Middle and late Pliocene. Dispersal of Blarinini 

from North America to Eurasia.

6. Late Pleistocene or Recent. Dispersal of Blarinini 

from North America into adjacent South America; 

introduction of some crocidurines to Pacific islands 

by man.

KEY TO IIVING AND FOSSIL GENERA OF THE SORICIDAE 

[Based on mandible and lower dentition]

la. Internal temporal fossa of the mandible not pocketed

(Subfamily HETEROSORICINAE), p. 7 

2a. Entoconid crest present; teeth delicate. Domnina, p. 7
64 SUBFAMILIES AND GENERA OF THE SOKICIDAE

la. Internal temporal fossa of the mandible not pocketed Con. 

2b. Entoconid crest absent.

3a. Teeth delicate, cingulum not inflated; talonid of 

MS not reduced and with rectangular crest 

through hypoconid and entoconid

Paradomnina, p. 10

3b. Teeth stout to bulbous, cingulum inflated; talonid 

of Ms reduced with crescentic crest through hypoconid ____________________ Trimylus, p. 10

3c. Teeth stout, cingulum strong but not inflated; Ms 

lost_______________________ Ingentisorex, p. 15

Ib. Internal temporal fossa of the mandible pocketed.

4a. Interarticular area of mandible prominent lingually, 

or articular facets confluent lingually, and excavated 

labially.

5a. Teeth not pigmented; P4 tetrahedral with prominent anterior, posterolateral, and posterolingual 

crests radiating from apex of principal cusp; 

range Old World

(Subfamily CROCIDURINAE), p. 15 

6a. Mandibular dental formula 1-4-3.

"Sorex" dehmi, p. 15 

6b. Mandibular dental formula 1-3-3.

7a. PS nearly as large as P4 ; talonid of Ms 

unreduced, double-cusped with welldeveloped basin; incisor short, curved, 

with two sharp, anteriorly inclined cusps 

on its cutting edge___ Soricella, p. 21 

7b. PS minute; talonid of Ms slightly reduced, 

single-cusped but with entoconid crest 

and weak basin; incisor with two very 

low, rounded cusps on cutting edge

Miosorex, p. 18

7c. PS minute; talonid of Ms reduced to single 

cusp; incisor short.____ Myosorex, p. 19

6c. Mandibular dental formula 1-2-3.

8a. Mental foramen beneath P4 to paraconid of MI. 

9a. Incisor with prominent scallops on

cutting edge.

lOa. Three rounded scallops as in Sorex 

Sylvisorex, p. 22

lOb. Two anteriorly inclined pointe 

cusps as in Soricella

Feroculus, p. 18 

9b. Incisor not prominently scalloped.

11. Talonid of Ms virtually lost, no 

cingulum on molars, condyles 

very specialized

Solisorex, p. 21

12. Pt very elongate, condyles united 

Surdisorex, p. 22

13. Talonid of MI anterosposteriorly 

compressed, minute PS present 

in some individuals

Myosorex, p. 19

14. First antemolar behind incisor 

elongate relative to P* as in 

Sylvisorex...-Scutisorex, p. 21

15. P* two-rooted

"Sorex" pusilliformis, p. 15

Ib. Internal temporal fossa of the mandible pocketed Con. 

4a. Interarticular area of mandible prominent lingually Con. 

5a. Teeth not pigmented Continued

6c. Mandibular dental formula 1-2-3 Continued

8b. Mental foramen beneath protoconid of M\.

16a. Cingulumlike groove along medial

side of incisor continues above

notch in basal border of tooth

Suncus, p. 21

16b. Cingulumlike groove along medial 

side of incisor continues below 

notch in basal border of tooth.

17. Talonid of MI moderately compressed anteroposteriorly; coronoid process broad at base 

Crocidura, p. 16

18. Talonid of M\ greatly compressed 

anteroposteriorly; coronoid 

process narrow at base

Diplomesodon, p. 17

19. First post-incisor antemolar very 

elongate and anteroposteriorly 

double-cusped

Paracrocidura, p. 20

8c. Mental foramen beneath hypoconid of MI, 

size gigantic _________ Praesorex, p. 20

5b. Teeth pigmented at tips; P4 a longitudinal blade or 

tending that way, posterolingual crest from cusp 

apex reduced or lost; range North America

(Subfamily LIMNOECINAE), p. 24

20a. Posterolabial crest of P4 terminates against

posterior cingulum midway between posterolabial corner of tooth and midline of

tooth-----.-..--------Angustidens, p. 25

20b. Posterolabial crest of P4 terminates against 

posterior cingulum at midline of tooth

Limnoecus, p. 26

4b. Interarticular area of mandible prominent labially or 

facets confluent labially, interarticular area excavated 

lingually; P4 with prominent anterior and posterolabial crests radiating from apex of principal cusp but 

posterolingual crest essentially lacking, posterolabial 

crest curves lingually and merges with posterior 

cingulum of tooth enclosing a posterolingual basin

(Subfamily SORICINAE), p. 27

21 a. Condyles of mandible not widely separated and 

lower condyle not moved greatly forward on 

the mandible; entoconid crest present on 

MU -_-_-------_-(Tribe SORICINI), p. 29

22a. Mandibular condyles confluent.

23a. Posterolabial crest of P4 does not merge 

with posterior cingulum

Crocidosorex, p. 29

23b. Posterolabial crest of P4 merges with posterior cingulum.-----Antesorex, p. 30

22b. Mandibular condyles separated..

24a. Ms unspecialized, virtually as in Domnina 

with distinct entoconid on rectangular 

hypolophid; pigmented dark red; considerable variation in pattern of 

mandibular condyles ...-Sorex, p. 31
KEY TO LIVING AND FOSSIL GENERA OF THE SORICIDAE 65

lb. Internal temporal fossa of the mandible pocketed Con. 

4b. Interarticular area of mandible prominent labially or

facets confluent labially Continued 

21a. Condyles of mandible not widely separated and 

lower condyle not moved greatly forward on the 

mandible Continued

22b. Mandilar condyles separated Continued 

24b. Ma unspecialized, teeth heavy and low 

cusped; pigment orange; pattern of 

mandibular condyles as in Sorex 

araneus but large, stout, and dorsal 

condyle extended further back

Drepanosorex, p. 32

24c. M3 slightly reduced with entoconid 

lacking and crescentic hypolophid; 

pigmented dark red; Pt somewhat 

compressed anteroposteriorly; condyles 

relatively large and close together

Microsorex, p. 33

24d. M3 with greatly reduced talonid; teeth 

stout, entoconid crest very prominent 

and postentoconid valley very wide 

on M\.

25a. Metalophid of Ms forms median 

crest with only minor remnant of 

entoconid crest; mandibular condyles separated but pattern not 

greatly different than in Antesorex 

Alluvisorex, p. 33

25b. Metalophid of Ms forms single 

elongate cusp on talonid, faint 

trace of entoconid crest; labial 

cingulum on MI deep below protoconid; mandibular condyles well 

separated___ -.-Petenyia, p. 34 

25c. Metalophid of M3 forms single 

conical cusp on talonid, no trace 

of entoconid crest; labial cingulum 

on MI deep below protoconid; 

mandibular condyles widely separated. ___-.-.-Blarinella, p. 34 

21b. Condyles of the mandible widely separated and 

lower condyle moved conspicuously forward 

on the mandible, interarticular area broad; 

entoconid crest lacking on MI

(Tribe BLARININI), p. 37 

26a. Lower condyle not hidden behind lower

sigmoid notch in labial aspect. 

27a. Mental foramen below protoconid of 

MI_________-__ Adeloblarina, p. 37

27b. Mental foramen below hypoconid of 

MI-- _____________ Cryptotis, p. 39

26b. Lower condyle meets lower sigmoid notch 

and is barely visible in labial aspect 

"/Sorex" dekneli, p. 42, and

Paracryptotis, p. 41

26c. Lower condyle entirely anterior to lower 

sigmoid notch and not visible in labial 

aspect.

28a. ?4 bulbous; entoconid present on MS; 

labial cingulum on M\ inflated

Blarinoides, p. 44

lb. Internal temporal fossa of the mandible pocketed Con. 

4b. Interarticular area of mandible prominent labially or

facets confluent labially Continued 

21b. Condyles of the mandible widely separated Con. 

26c. Lower condyle entirely anterior to lower 

sigmoid notch and not visible in labial 

aspect Continued

28b. PI not bulbous; entoconid present or absent 

on My when absent hypolophid crescentic; labial cingulum on M\ deep 

but not inflated-____ Blarina, p. 42 

28c. Pt not bulbous; entoconid and hypolophid 

absent on M3 talonid reduced to metaconid crest; labial cingulum on M\ 

inflated. _____ Shikamainosorex, p. 42 

21c. Condyles of mandible widely separated and lower 

condyle moved conspicuously forward on the 

mandible and located lingually so that groove 

separates lower condyle from lower sigmoid 

notch, interarticular area narrow; entoconid 

crest present on M\ except in Anourosorex and 

Amblycoptus___ (Tribe NEOMYINI), p. 45 

29a. Upper mandibular articulation with oval

facet.

30a. Ms unreduced, entoconid present, hypolophid rectangular.

31. Incisor long; pigment conspicuous

Neomys, p. 46

32. Incisor short; pigment slight

Episoriculus, p. 47

33. Incisor short; pigment variable and 

light orange; entoconid crest very 

low on M\; labial cingulum not continuous around hypoconid on MI

Soriculiis, p. 52

34. Incisor short to intermediate; pigment 

slight to absent; labial shearing 

blade of Pt on longitudinal midline 

of tooth_________ Nesiotites, p. 51

35. Incisor long; pigment very light but 

widespread; labial cingulum present 

only below paraconid of MI

Nectogale, p. 51

30b. Ms reduced with crescentic hypolophid. 

36a. Pigment not visible in ordinary light; 

labial cingulum weak on MI behind protoconid

Chimarrogale, p. 53

36b. Pigment prominent; labial cingulum 

strong and deep around MI

Beremendia, p. 50

30c. M3 reduced with single cusp on talonid

Petenyietta, p. 46

29b. Upper mandibular articulation with triangular facet. 

37a. M 3 unreduced.

38a. Pigment prominent; entoconid crest

of MI strong_ ..-Neomys, p. 46

38b. Pigment slight; entoconid crest of

MI very low._ Megasorex, p. 56
66 SUBFAMILIES AND GENERA OF THE SORICIDAE

Ib. Internal temporal fossa of the mandible pocketed Con.

4b. Interarticular area of mandible prominent labially or

facets confluent labially Continued 

21c. Condyles of mandibles widely separated Con. 

29b. Upper mandibular articulation with triangular

facet Continued 

37a. M3 unreduced Continued

38c. Pigment slight; entoconid crest of M i 

strong_____._ --..Chodsigoa, p. 48

37b. M3 reduced.

39a. Entoconid lacking but entoconid 

crest and talonid basin present 

on M3 ; very little pigment. 

40a. Tip of coronoid process massive, coronoid spicule large

Chodsigoa, p. 48

40b. Tip of coronoid process slight, 

coronoid spicule small

Notiosorex, p. 55

39b. Entire talonid of M 3 virtually lacking. __________Anourosorex, p. 53

39c. M 3 lacking.-.--.Amblycoptus, p. 54 

4c. Mandibular articulations widely separated and upper 

articulation placed very far to the rear; teeth unpigmented and high cusped with carnivorelike appearance; 

MI with trigonid very long, talonid very short, paralophid oriented essentially anteroposteriorly, metaconid 

close to protoconid, metalophid unites protoconid to 

hypoconid, hypolophid rectangular with no post entoconid valley, entoconid crest lacking and talonid basin 

opens lingually between metaconid. and entoconid 

(Subfamily ALLOSORICINAE), p. 57 

41a. Mandibular condyles not extremely separated; 

labial cingulum present on cheek teeth

"Sorex" gracilidens, p. 57

41b. Mandibular condyles extremely separated; labial 

cingulum lacking on cheek teeth

Allosorex, p. 57 

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europaischen Tertians und deren Beziehungen zu ihren 

lebenden und fossilen aus europaischen Verwandten: 

Palaontologie Geologic Osterreich Ungarns Beitr. v. 6, 

pts. 1 and 2, p. 1-227, 9 pis.

 1924, Tertiary vertebrates from Mongolia: Palaeontologia sinica, ser. C, v. 1, no. 1, p. 1-119, 5 figs., 6 pis.

Shikama, Tokio, and Hasegawa, Yoshikazu, 1958, On a new Anourosorex from the Ryugasi formation (fissure deposits) in 

Japan: Yokohama Natl. Univ. Sci. Rept., sec. 2, no. 7, p. 

105-112, pi. 16, 1 map.

Simpson, G. G., 1941, A new Oligocene insectivore: Am. Mus. 

Nat. History Novitates, no. 1150, p. 1-3, 1 fig.

 1945, The principals of classification and a classification

of mammals: Am. Mus. Nat. History Bull., v. 85. 350 p. 

Stehlin, H. G., 1940, Zur Stammesgeschichte der Soriciden:

Eclogae geol. Helvetiae, v. 33, p. 298-306, 5 figs. 

Stirton, R. A., 1930, A new genus of Soricidae from the Barstow

Miocene of California: California Univ. Dept. Geol. Sci.

Bull., v. 19, no. 8, p. 217-228, 2 figs. 

Stromer, Ernst, 1928, Wirbeltiere im obermiocanen Flinz Miinchens: Bayer. Akad. Wiss. Abh., Math. nurw. Abt., v. 32,

no. 1, p. 1-71, 3 figs., 3 pis. 

Sulimski, Andrzej, 1959, Pliocene insectivores from We.ze: Acta

palaeont. polonica, v. 4, no. 2, p. 119-175, 5 figs., 4 pis.

 1962, Supplementary studies on the insectivores from 

We.2e I (Poland): Acta palaeont. polonica, v. 7, nos. 3-4, 

p. 441-502, 2 figs., 2 pis.

Suthard, J. A., 1966, Stratigraphy and paleontology in Fish 

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Riverside, M.A. thesis.

Tedford, R. H., 1961, Clarendonian Insectivora from the Ricardo 

Formation, Kern County, California: Southern California 

Acad. Sci. Bull., v. 60, pt. 2, p. 57-76, 4 figs.

Thomas, Oldfield, 1906, New African mammals of the genera 

Ceropithecus, Scotophilus, Miniopterus, Crocidura, Georychus, 

and Helipohobius: Annals Mag. Nat. History, ser. 7, v. 17, 

p. 173-179.

 1913, On African bats and shrews: Annals Mag. Nat. 

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-1924, A new genus and species of shrew from Ceylon:

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Tobien, Heinz, 1939, Die Insektenfresser und Nagetiere aus der

aquitanen Spaltenfullung bei Tomerdingen (Ulmer Alb):

Naturf. Gesell. Freiburg Ber., v. 36, p. 159-180, 1 pi. 

Viret, Jean, and Zapfe, Helmuth, 1951, Sur quelques Soricides

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12 figs. 

Wilson, R. W., 1960, Early Miocene rodents and insectivores

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2 figs.

Young, C. C., 1934, On the Insectivora, Chiroptera, Rodentia, 

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REFERENCES 69

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Zdansky, Otto, 1928, Die Saiigetiere der Quartarfauna von

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palaeontologischen Kenntnis der Cyrenenmergelschichten im 

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no. 42, p. 87-146, 3 figs., 4 pis.

INDEX

[Italic page numbers indicate both major references and descriptions]

A Page 

adamsi, Cryptotis...................... 39,40,flg. 26

Adeloblarina.... 36, 57, 38, 39, 41, 42, 43, 44, 55, 62

berklandi......................... 37.38, fig. 25

Africa.-- . 2, (50,62

Allopachyura................................. 47,58

Allosorei--................................. __ 67

stenodus---............................... 6,57

Allosoricinae.. ______________ 2,5,57

Alluvisorex....................... .......... S3

arcadentes................................. 33,34

chasseae........_____...._.. 33, 34, fig. 22

alpinoides, Sorex.............................. 32

alpinus, Sorex................................ 24

ambiguus, Sorex............................... 30

Amblycoptus.-............................. 27,54,63

oligodon ................................ 54,55

Amphisorex. See Sorex or Neomys.

Ancestry of the Soricidae.__________.. 58

Angustidens.-. ________________ 25,33,60

vireti.......................... 25, 26, 30, fig. 16

Anotus. See Blarina.

Anourosorex...- 2, 14, 24, 42, 45, 48, 50, 52, 63, 54

assamensis................................ 54

inexpectatus............................... 54

japonicus................................. 54

squamipes____________.. _ 53,54, fig. 39 

Antemolar, definition...________..__ 3 

Antesorex......................... 30,31,33,36,60,62

compressus.-....................._ 31, fig. 19

antiguus, Crocidosorex................... 15,30,31,58

Sorex..................................... 29.30

araneoides, Sorex.............................. 32

araneus, Sorex................... 26,28,29,31,fig. 20

arcadentes, Alluvisorex........................ 33,34

Archaeolagus............. ..______..__.. 9

Arkansas. .._____.___________.. 33 

Asoriculus.................................... 58

gibberodon................................ 48

assamensis, Anourosorex...................... 54

Atavism.. -.. -- ..__........ . 3,31

Atophyrax. See Sorex.

Austria-__....._ . 32,33

Beremendia............................. 50,51,52,55

fissidens .................... 51, 54, fig. 35

sinmsit ............................. 51

ucrainica................................. 51

berklandi, Adeloblarina................. 37,38, fig. 25

Blarina...................................... 29,35,

37,39,40,41, 42, 44,45,50,51,52,53,54,55,62,63 

brericauda 25,38,42,43,44, fig. 29

fossilis......_____._____ __ 44

Oidleyi....................... 41,42,43,44,fig. 30

simplicidens.............................. 44

blarina zinki, Myosorex........................ 22

Blarinella...... 29,33, 34, 35,36,38,44,45,48,50,55,62

kormosi................................... 35,45

quadraticauda____________ 34,35, fig. 24 

wardi..................................... 45

Blarinini- . . -- 29,35,

36,37,38,39,42,44,45,52,53,55,57,62,63

incertae sedis______________. 44

Blarinoides................................ 44,51,62

mariae.................................... 42,44

bohlini, diodsigm............................. 50

Neomys..-............................... 50

Brachysorex. See Cryptotis.

breaicauda, Blarina............ 25,38,42,43,44, fig. 29

browni, Neomys. ........................... 46

Burma.. 62

caeruleus, Suncus...................... 20,22,fig. 13

California-.- 9,25,26,34,55

castellarina, Neomys.......................... 46

caudatus, Episoriculus.................... 48,fig. 32

Sorex.............................____ 47

Ceylon.. . 2,18,21,57,60

chasseae, Alluvisorex................... 33,34,fig. 22

Chimarrogale........................ 24,37,51,53,54

himalayica............................ 53,fig. 38

platycephala.-............................ 53

China--._ 17,35,45,48,50,51,52,53,54,62 

Chiroptera..-_-_. ._. __ 5,58,59 

Chodsigoa................ 45,48,49,50,52,53,54,55,56

bohlini..__________________ 50 

hypabia........................... 49,50,fig. 33

salentkii..................... 49,50,55,56,fig. 34

cinereus, Sorex. __________________ 27,32 

collongensis, Sorex -........................... 24

Colorado.- 8,11,13,25,31

compressa, Domnina.. _____________ 11 

compressus, Antesorez..................... 31,fig. 19

Heterosorex..-......... __________ 11

Sorex............................... . 30,31

Trimylus- _____......__.. 11,12,13, fig. 5

Condyle, mandibular___________._ 5 

basic patterns..-_________..-. 6,fig. 1 

function________________ 28

Congosorex. See Myosorex.

Coronoidspicule--..____ 37,45,55, figs. 25, 29, 31

Correlation of the Soricidae____.__.... 69

Corsica_____________________ 51 

corsicanus, Nesiotites......................-... 51

Cosira. See Sorex.

nigrescent................................. 52

crawfordi, Notiosorex................... 49, 55, fig. 40

Sorex (Notiosorex)------.................. 55

Crocidosorex................... 29,30,31,36,58,60,62

antiquus__________._____ 15,30,31,58 

piveteaui................................ 2,29,30

Crocidura............ 2,6,15, 16, 17,18,20,22,23,47,62

gibberodon.---............................ 48

goliath.................................... 20

kormosi________________...... 2,35

kornfeldi_______________...-..-. 17 

leucodon..____________._..._ 17 

morio_____________________ 22 

obtusa.-.................... . 17

russula-. __________________... fig. 8 

suaveolens.-.-.....______...___ 17

taungsensis... _______________.. 17 

wongi............... ____________.. 17

zorzii-.................................... 17

sp_ ....... ...... 15,17,24

Crocidurinae.. . 1,2,3,5,6,7, 16, 24,25,59,60,62,63 

African aflBnities ____.... _... 15,60

incertae sedis.___________ .... 23

Crossogale. See Chimarrogale. 

Crossopus. See Neomys.

fissidens.............________.__. 50

himalayanus.............................. 53

Page 

Cryptotis--.................... 37,39,40,41,55,62,63

adamsi.________________ 39,40, fig. 26 

kansasensis-........... __________. 40

magna.................................... 40

meadensis .................._____ 40

mexicana..-- _______________ 41 

parva...................... 38, 39, 40, 41, fig. 27

cudahyensis, Sorex............................ 32

Cusps, height. -----________-_____ 6

Czechoslovakia- ... 32,33,34,46,51,57

D

dakotensis, Trimylus- ________ __ 12,13, fig. 6 

daubentonii, Sorex......... ___________ 46

dehmi, "Sorex"........................... 15,16,24

dehneli, "Sorex"........................... 42,44,63

Deinsdorfia.......-_______________ 57

delphinensis, Heterosorex..............____ 14

densicingulata, Shikamainosorex...... _____ 42

Dental formulae of the soricids. -.-_____ 2 

Diplomesodon................................. 17,21

fossorius............................. 6,18,21,63

pulchellum............................ 18, fig. 9

discrepans, Soricella........................... 15,21

.dixonensis, Sorex.-.....................___ 32

Domnina---......................._____.. 3,

5, 6, 7, 8, 9, 10, 11, 13, 15, 26, 27, 29, 

31, 33, 52, 58, 60. 

compressa................................. 11

gradata.. .. 3, 5, 8, 9, 10, 11, 13, 14, fig. 3 

greeni ...............________.--.-. 9

thompsoni.............. 5, 8, 9, 10, 12, 13, fig. 2

n. sy................................ 9,10, fig.4

Drepanosorex. -............................... 32

margaritodon.............................. 33

pachyodon.-..________.... 33

savini..................................... 33

tasnadii................................... 33

(Drepanosorex) tasnadii, Sorex................. 32

E 

elegans, Nectogale...................... 51,52, fig. 36

Embryologic studies .. . 2

Endemism.... ......__.. 59,60,62

England. 32,33,46

Entoconidcrest..----.. --. 7, 10, 28, 29, figs. 2,3

Episoriculus......______._______ 47,48,58

caudatus.. -...................... 48, fig. 32

gibberodon---............................. 48

Erinaceoidea..._______.. .. 59 

etruscus, Suncus........_________.... 22

Eurasia.._____________________ 60 

Europe._....____._-_-_..._.. 60,62,63

Evaluation of characters____________ 3 

External temporal fossa_- 29, 37, 45, figs. 25, 29, 31

fallax, Sorex 32

Faunal exchanges, intercontinental. _____ 63 

Feroculus................................ 2,18,20,23

feroculus, Sorex.. _________. 18

fissidens, Beremendia.................. 51, 54, fig. 35

Crossopus................................ 50

fodiens, Neomys........................ 38,46, fig. 31

Fossa pterygoidea. ________ 2,5

71
72 INDEX

Fossil localities: page 

Beatty Butte, Oreg ______ ..... 13

Bowden Hills, Oreg...__.____..._ 38 

Choukoutien, China.... 50,51

Christmas Valley, Oreg-----_______ 40

Cuyama Valley, Calif .________. 26 

Ertemte, Mongolia.____ 35,54

Flint Hill, S. Dak. .. . 12 

Quano Ranch, Oreg-___________ 13 

Hagennan, Idaho..._______--___ 44 

Ivanovce, Czechoslovakia.-..._____ 57 

La Qrive St. Alban, France.--..._-.-.. 14,19 

Montaigu-le-Blin, France_________ 30 

Polgardi, Hungary..-_____ ... 54 

Rexroad, Kansas...______.____ 40,41 

Rome, Oreg--_______________ 26,41 

Snyder Creek, Oreg--_._________ 13 

Vallecito Creek, Calif.-- .. 55 

Valles-Penedes, Spain.................... 14

Vieux-Collonges, France____------__ 16

Weisenau, Germany___________ 12,30 

We.ze Breccia, Poland-,__.... 36,37,45,46,47

Wintershof-West, Germany.-... 12,15,21,24.36 

fossilis, Blarina. _________________ 44 

fossorius, Diplomesodon. ............__- 6,18,21,63

France--. . .... 14,16,19,29,30,32

frankstownensis, Sorex_____..._.___. 32

G

Germany ___ -...--. 12,15,21,24,30,32,33,36,46,57

gibber odan, Asoriculus _____________ . 48

Crocidura. ...... .... .... ...... .... ...... .. 48

Episoriculus .............................. 48

Soriculus. ................................ 48

gidleyi, Blarina.................. 41, 42, 43, 44, fig. 30

gigas, Megasorex....... ...... . ........... - 56, fig. 41

Noliosorex... ............................. 56

goliath, Crocidura.. ........................... 20

Praesorex.. .... . .......... ......... 20,21,fig. 12

gracilidens, "Sorex". .......................... 57

gracilis, Petenyiella ............................ 46,47

Saturninia ......................... ___ 4

Sorex ................. .. ........... ___ 46

gradata. Domnina. ........ 3,5,8,9,10, 11, 13, 14, fig. 3

greeni, Domnina. _______________ 9 

grieensis. Miosorex... ........... . 15, 16, 19, 33, fig. 10

Sorex.... ... . ..... . ...... . ..... . ..... . .... 19

pusittus. .............................. 18,19

H

Heel of -Ma-.-- 6

Heliosorex. See Crocidura.

Hesperosorex ________________ . 52,55,63 

lovei. ..................................... 55

Heterosorex.- ............................. . 10,12,58

compressus . . . . ........................... 11

delphinensis ............................... 14

neumayrianus. .. ______________ 11, 12 

roperi ..................................... 13

sansaniensis. . . . ......... ____ ____ 14

sp.-- . ... _ _..-.. ___ .--....... 13

Heterosoricinae ________________ 2,

3, 5, 6, 7, 8, 15, 20, 45, 58, 59, 60, 62, 63

hibbardi, Sorex. ............................... 35

hidalgo, Nesiotites. ________________ 51 

himalayanus, Crossopus ............. . ......... 53

himalayica, Chimarrogale .... . ............. 53, fig. 38

Hipparion .................................... 54

Holarctica ___________________ 63

Homalurus. See Sorex.

hoyi, Microsorex ..... . ... ... ..... ... .... 30,31, fig. 21

Sorex ..................................... 33

hungarica, Petenyia. ...................... 34, fig. 23

Hungary-.. _________ ..... 32,33,44,47,48,54

Hydrogale. See Neomys.

Hydrosorex. See Neomys.

Hypothetical Soricid ancestor _ ________ 58

hj/psitria, Chodsigoa ........... .......... 49, 50, fig. 33

hypsifnus, Boriculus ........................... 48

I Page 

Idaho 44

Illustrations, contents and preparation.-.__ 2 

India 48,53,60,62

Indochina.___________________ 62 

inexpectatus, Anourosorex..................... 54

Inferior pterygoid fossa____________fig. 11 

Inferior sigmoid notch___________... fig. 8 

Ingentisorex............_.-..._-..-..-- 3,7,15,60

tumidtdens................................ 15

intermedius, Neomys.......................... 46

Internal temporal fossa of the mandible...__ 2,

5, figs. 2,10

Intertemporal fossa... . . 2,5 

Italy--.-...... 17,33,46

jacksoni, Notiosorex......................... 48,49,55

Japan.--........ . . 42,54,63

japonicus, Anourosorex...................._. 54

K

Kansas... -. . - 32,33,40,41,44,55 

kansasensis, Cryptotis......................... 40

kennardi, Sorex............................... 32

Kenya.... 17,22

kinkelini, "Sorex"............................. 7,12

kormosi, Blarinella............................ 35,45

Crocidura................................. 2,35

kornfeldi, Crocidura........................... 17

kretzoii, "Sorex"..................... 35,42,44,45,63

kubinyi, Soriculus____________..._ 4§ 

kura, Podihik................................. 57

lacustris, Neosorex. .................. ......... 32

Sorex. ..... . .... . ................ . ........ 32

leahyi, Sorex. ................................. 32

Leptacodon... .... ---------------------------- 4

Leucodmi. See Crocidura.

leucodon, Crocidura--. __ 17

Sorex. _____________________ 16

Leucorrhynchus. See Neomys.

Limnoecinae- - - . 2, 3, 5, 6, 15, S4, 40, 59, 60, 63 

parallel evolution _____________ 60

Limnoecus ................................. 24,26,60

micromorphus ..... _________ . __ 23,24

niobrarensis ..... .... ... 5, 6, 23, 24, 26, 27, fig. 18

tricuspis. .............. ........ 25, 26, 27, fig. 17

vireti.... ...... ............. ... . ..... ...... 25

sp.. _____ .... ... ..... ... 25

lixus, Suncus.. ............................... 16

lovei, Hesperosorex.--..-. ... ... ... .... ... ..... 55

lunaris, Sylvisorei ... ... ... ... ... .......... 23, fig. 15

M

M' and P*, structure. _____________ 7 

magna, Cryptotis. ... .......... ... . ...... ...... 40

Mallorca.-- __ .. ... ... _ . 51 

Mamblarinaus. See Blarina.

margaritodon, Drepanosorex _ _ . ________ 33 

mariae, Blarinoides.... ... ........ .......... .. 42,44

Maries.. ... ........ . ___________ ___ 57

martini, Mydecodon.. .... . .................... 5

Masseteric fossa.- _ --- - 9, 58, fig. 2 

mawbyi, Trimylus... ........ .. ...... 12, 13, 14, fig. 7

meadensis, Cryptotis.... ........ .. ... ..... .... 40

megalura, Sylvisorex. ................ .......... 22

Megasorex. ......... ................. . 53,55,56,63

gigas....... ........................... 56, fig. 41

Mental foramen, position... ____ . __ __ 2,6 

merriami, Sorex ............................... 27

Metaconid, closeness on Mi to protoconid. . - . 7 

mexicana, Cryptotis-. . .... .. .... ...... ... ..... 41

Mexico... . 41,56

meyeri, Oligosorex. ....... ___ . _ -... 30,58

micromorphus, "Limnoecus"... ..... .. ........ 23,24

Page 

Microsorex.................................... 26,33

hoyi............................... 30,31, fig. 21

minutus-................................. 33

pratensis ............................... 33

Miniopterus.................................. 5

Minorca____________________ 51 

minutissimus, Sorex........................... 32

minutus, Microsorex.......................... 33

Sorex..................................... 32

Miosorex...................................... 18

grivensis--- ............... 15,16,19,33, Bg. 10

Miothen...................................... 58

Mongolia ...___._____-...._.__ 35,54

Montana____________________ 8 

mono, Crocidura.............................. 22

Morphologic characteristics, Allosoricinae. _ 57 

Blarinini................................. 37,62

Crocidurinae-. -___..-.-..__... 2,15,60 

Heterosoricinae______________ 7,60 

hypothetical shrew ancestor..___ ... 58 

Limnoecinae-_______________ 24,60 

Neomyini...^_______________ 45,63 

shared with insectivores and bats 59 

Soricinae. _________....___.. 27

Soricini......... ...... __.. 29,62

subfamilies-..._ ____... 3 

Musaraneus. See Sorex. 

Mydecodon martini............................ 5

Myosictis. See Neomys.

Mysorex-..................................... 19,22

blarina zinki.............................. 22

potti...................................... 19

robinsoni.................................. 20

varius................................ 19, fig. 11

Mystipterus................................... 5

N 

Nebraska 8,11,25,27

Nectogale.......................... 5,24,50,51,52,54

elegans............................ 51,52,fig. 36

neglecta, Petenyia............................. 34

Neomyini 20,28,36,37,38,39,

46, 46,47,48,49,50,51,52,53,55,57,58,60,62,63

incertae sedis___________ 57

Neomys... 13,28,36,37,45, 46, 47,48,50,51,52,53,54,57

bohlini ................................. 50

browni .................................. 46

castellarina-.............................. 46

fodiens .......................... 38,46, fig. 31

intermedius............................... 46

newtoni.......................... ... .. 46

soriculoides-. .......................... 36

Neosorex. See Sorex.

lacustris---............................... 32

Nesiotites..................................... 61,52

corsicanus-............................... 51

hidalgo.......... -~~ 51

similis.................................... 51

Nesophontes..- .- .---- 59

neumayrianus, Heterosorex.................... 11,12

Sorer 12

Trimylus .. ..... 7,11,12,13

Neurotrichus........................- . .-.. 5

Nevada---- .. - 25,34

New World distribution, Blarinini-. 62 

Limnoecinae--------------__---_----- 60

newtoni, Neomys...................... ... 46

nigrescens, Cosira.................... 52

Soriculus.......... ...... 53, fig. 37

niobrarensis, Limnoecus...... 5,6,23,24,26,27,fig. 18

norae, Surdisorex.... ... ..... 22, fig. 14

North America _ _ - 2,60,62,63 

Notiosorex.................. 37,48,49,50,53,55,56,63

crawfordi--.- 49, 55, fig. 40

gigas...... ...... . .. 56

jacksoni... 48,49,55

(Notiosorei) crawfordi, Sorex.............. . 55
INDEX 73

0

obtusa, Crocidura.--..___ . 

Oklahoma __ -__ . 

Old World distribution, Crocidurinae. 

Neomyini..__.___.. . 

Old World fossil shrews...___ ...

Page 

17 

44

1

62

2 

ctigodon, Amblycoptus......................... 54,55

Oligosarex ........................... .... 30,58

meyeri..................____- ..... 30,58

Oregon 10,13,15,25,26,34,37,38,40,41

Otiosorex. See Sorei. 

Oi-yrhin. See Sorer.

P 

Pt, structure..... --.... . . 2,3, fig. l

P*and M1 , structure.._________.__ 7 

Pachyura. [See Suncus.]..................... 21

pannonica.----------------............... 47

pachyodon, Drepanosorex.......... .... 33

Sorer.___.___---.__.--. .-__. 33 

Palearctica.- . .__. . . .. 2,62 

Paleozoogeography. -.____.... ... 59

pannonica, Pachyura-....... . -... 47

Paracrocidura...__._____.... --.-. 20

schoutedeni_____________...__ 20 

Paracryptotis 37,39,4*, 42,43,44,62,63

rez . 41,42,flg. 28

Paradomnina............................... 3, 10, 60

relictus................................... 10

Paradoxodon. See Suncus.

Parahippus................................... 9,

parva, Cryptotis................. 38, 39, 40, 41, fig. 27

pareus, Sorex................................. 39

Paurodus. See Crocidura.

pearsoni, Solisorex. ......... 21

Peisorex......................---------------- 45

pohaiensis................................ 44,45

Pennsylvania_...._______.... _.. 32,44

perminutus, Sorex............................. 32

Petenyia...... ........__......... 34,35,36,37,62

hungarica.......___.. 34,flg. 23

neglecta .................................. 34

stehlini-..................... ..... 34

suavensis................................. 34

PmtmyieOa.............................. 46,47,58

gracilits.................................... 46,47

zelcea....................... ... _ 46,47

Pigmentation_._________.--.-__ 5

Pinalia. See Neomys.

piveteaui, Crocidosorez...... ....._ .. 2,29,30

Plerodus. See Suncus.

Plesiosorex_____________......._.. 58

Podihik hum.................................. 57

pohaiensis, Peisorex. _________....._.. 44,45

Poland................. 32,36,37,42,44,45,46,47,48

polli, Myosorex.---.__________.. _.. 19

Posterointernal ramal fossa. .-.---. .._ 2,5 

praelpinus, Sorex..._________.__.. 32 

praearaneus, Sorex... ......__________ 32

Praesorex.............................. 16,17,20,21

goliath............................ 20,21, fig. 12

pratensis, Microsorex.......................... 33

Protoconid, closeness on Mi to metaconid.... 7

Protosorex.--------........................... 58

Pterygoid spicule (or spine)__ 28,37,45, figs. 33,34 

pulchellum, Diplomesodon................. 18, fig. 9

pulchellus, Sorex.............................. 17

pusiUiformis, Sorex............................ 15

pusillus, Sorex_____ _______ ____._ 15,30

grivensis, Sorex___________. ____.. 18,19

Pygmura. See Anourosorex.

Q

quadraticauda, Blarinella________ 34,35, fig. 24 

Sorex. _____________________ 34

R 

relictus, Paradomnina.-.______ ________ 10

rex, Paracryptotis.---....._____. 41,42,fig. 28

rexroadensis, Sorex. ______________ 32

Rhinomus. See Crocidura. Page 

robinsoni, Myosorex-.._____________ 20 

roperi, Heterosorex----.............. ....... 13

Trimylus ................ 11,12,13,14,60

runtonensis, Sorea..._______________ 32 

russula, Crocidura---. __._..----______.. fig. 8

sacer, Suncus---_________________ 21 

salenskii, Chodsigoa. ............. 49,50,55,56, fig. 34

sandersi, Sorex.-- ........................ 32

sansaniensis, Heterosorex... ___________ 14 

Sorex.. .. .. 10,14 

Trimylus-. ............ . -- 10,13,14

Sardinia . 51

Saturninia..............................___ 4,58

gracilis..........________________ 4

savini, Drepanosorex-............. ___________. 33

schlosseri, Trimylus............ _________ 10,14

schoutedeni, Paracrocidura___________ 20

Scutisorex................................... 1,15,2.

Shikamainosorex...______________ 42,44,63 

densicingulata............................. 42

similis, Nesiotites- ............... __ 51

simplicidens, Blarina.......................... 44

sinensis, Beremendia.......... __ 51

Solisorex.................................... 2,2^,63

pearsoni-........................_____ 21

somereni, Sylvisorex........................... 21

Sorea; 2,15,22,25,26,27,29,30, SI, 32,

33,34,35,36,37,38,39,42,43,44,45,46, 

47,48,51,52,55,58,62. 

alpinoides................................. 32

alpinus................................... 24

ambiguus................................. 30

antiquus.................................. 29,30

araneoides. .................... __ 32

araneus.-................... 26,28,29,31,fig. 20

caudatus ................................ 47

cinereus................................... 27,32

collongensis. ____________...__ 24 

compressus.. ________________ 30,31 

cudahyensis.-. ....-...-................. 32

daubentonii.............. _________ 46

dekmi.................................. 15,16,24

dehndi........................ ...... 42,44,63

dixonensis................................. 32

faUax..................................... 32

feroculus- ............................... 18

frankstownensis.__________.__ 32 

gracilidens...........................___ 57

gracilis............................. .-... 46

grivensis-..........................___ 19

hibbardi-................................. 35

lioyi...... ............................ 33

hennardi-......................-..-....... 32

kinkelini. ............................ 7, 12

kretzoii........................... 35,42,44,45,63

lacustris.................................. 32

leahyi..................................... 32

leucodon-....................... ._ __ 16

merriami---- ............... ... 27

minutissimus............................. 32

minutus...........................____ 32

neumayrianus............................ 12

pachyodon.--............................. 33

parvus .................................. 39

perminutus..................... ..____ 32

praealpinus............................... 32

praearaneus.....................-.-..__ 32

pulchellus............................... 17

pusittifofmis-.........................-... 15

pusillus...................... .......... 15,30

grivensis-.______ ... ..... 18,19

quadraticauda_____________..._ 34 

rexroadensis...______.... . . 32

runtonensis____________...__. 32 

sandersi..............___________ 32

sansaniensis-.-.__________. .-.-. 10,14

Sorex Continued

stehlini---- -.--..

subaraneus........__.

subminutus.______.

talpoides................

taylori.- ________.

varius . .

vireti- .........

vvlgaris-..........__.

(Drepanosorex) tasnadii-

(Notiosorex) crawfordi... 

Soricella..___ ___.

 .... 36

 32

 . ... 32

 - .-... 42 

. ..... 32

 ..... 19

. . .. 25 

.- -.-... 28 

. ..... 32

. ..... 55

.. ..-... SI 

discrepans.-. -______________ 15,21 

Soricidae...___..__ 1,2,3,5,7,16,45,48,58,59,63 

incertae sedis.______________ 57 

Soricidus. See Sorex.

Soricinae.-.__ _____________ 1, 

2,3,5,6,7,24,27,28,29,30,38,45.59,60, 

62,63.

ancestral characteristics_...______ 62 

Soricini __-.-.- ______________ 29, 

33,36,37,38,39,44,45,55,57,58,59,62, 

63.

incertae sedis._______________ 35 

Soriciscus. See Cryptotis.

Soricoidea..__________________ 59 

soriculoides, Neomys.......................... 36

Zelceina.................................. 37

Soriculus................... 36,47,48,50,51,52,53,54

gibberodon..-............................. 48

hypsibius.-............................... 48

kubinyi................................... 48

nigrescens............................ 53, fig. 37

SouthAfrica. ..... 17,18,19,20,22

South America... ______________ 63 

South Dakota...____.._____....__. 9,12

Spain........ .. 14

s, Anourosorex................ 53,54,flg. 39

Stability of morphologic characteristics. ___ 3 

stehlini, Petenyia......................... 34

Sorez.____...__._ ... 36 

stenodus, Allosorex...........................- 6,57

suavensis, Petenyia............................ 34

suaveolens, Crocidura__________.. .._ 17

subaraneus, Sorex............................. 32

Subfamilies.. - _ 2 

origins...... _ . 63

subminutus, Sorex-...__._.. . 32 

Suncus.... ... ... ... ... 6,16,19, SI, 22,23,46,47,62

caeruleus.......................... 20,22,fig. 13

etruscus ............. 22

lixus ---------.--...---. - ---- -- 16

sacer.................... 21

sp.. ........ ..... 22

Sunkus. See Suncus.

Superior pterygoid fossa..___.- figs. 11,29,31 

Superior sigmoid notch.__ fig- 8 

Suprageneric taxa, characteristics.- -. 2 

Surdisorex- .-.. 22 

norae................................. 22, fig. 14

Switzerland... . 11 

Sylvisorex.................... . - 15,18,20,22

lunaris.-.- . --- 23, fig. 15 

megalura.--- ---------------- 22

somereni.... 21

Talonid of Ms.. 6

Talpidae.--.. . -... ------ ------ 59

talpoides, Sorex..-. 42

Talposorex. See Blarina.

tasnadii, Drepanosorex- ._ .. ------ 33

Sorex (Drepanosorex)........ ......... 32

taungsensis, Crocidura---- -------------------- 17

taylori, Sorex. . 32 

Tethys seaway.. S9, 62 

thompsoni, Domnina......... - 5,8,9,10.12,13, fig. 2

tricuspis, Limnoecus. - - 25,26,27,flg. 17
74 INDEX

Page

Trimylus . . . . - 3,5,6,7,8, 10,

11,12,13,14,15,42,45,58,60,63

compressus____________ 11,12,13, fig. 5

dakotensis._..------------..___ 12,13,fig. 6

mawbyi. . 12,13,14,fig. 7

neumayrianus __.---__------- 7,11,12,13

roperi . - .-. .---- 11,12,13,14,60

sansaniensis--......................... 10,13,14

schlosseri....________.______... 10,14

habits. --.-. - - -- --- 14

two lineages. __________ _____ 10

tumididens, Ingentisorex....................... 15

ucrainica, Beremendia. 

Uganda._______ 

U.8.8.R..............

Page 

51 

23

18

Variation of rates of evolution...... __... _ 59

varius, Myosorex....................._.. 19,fig. 11

Sorex.. ...._........_._-------_.- 19

vireti, Angustidens--. ........... 25, 26, 30, fig. 16

Hmnoecus_______________ 25

Sarex... ............................... 25

vulgaris, Sorex-.- ____.___________ 28

W Page

wardi, Blarinella_._____________. 45 

West Indies.._________________ 59 

wongi, Crocidura_______ ________ 17 

Wyoming.................................. 8,25,55

Z

zelcea, Petenyietta________________ 46,47 

Zelceina. ___...____________.. 36, 37

soriculoides ......______________ 37

zinki, Myosorex Marina...____...._.__ 22

zorzii, Crocidura----- ___________. 17

Zygomatic process.._________ 7,9,29,37,45,58

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