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U. S. DEPARTMENT OF AGRICULTURE
DIVISION OF ORNITHOLOGY AND MAMMALOGY

NORTH AMERICAN FAUNA

No. 8 ^^^>:,^,^..*^

PUBLISHED BY ArTTIORITY OF THE SECRETARY OF AGRICULTURE

[Aftitiil (late of pnhlication, .Tannary 31, 1895]

''^

MONOGRAPHIC REVISION

OF iiii:

Family <iJKOIflHI>JG

(Exclusive of the species of Thomomys)

liV

Dr. C. HART MERRIAM

WASHrXGTOI^

GOVERNMENT PRINTING OFFICE

1895

->f\/T%o b ;j J -^^UJ.J^A- U^>'*>-*-'-*-\j >—

U. S. DEPARTMENT OF AGRICULTURE
DIVISION OF ORNITHOLOGY AND MAMMALOGY

^^ 3 ■ ^

NORTH AMERICAN FAUNA ,. -

No. 8

PUBLISHED BY AUTHORITY OF THE SECRETARY OF AGRICULTURE

[Actual date of publicatiou, January 31, 1895]

MONOGRAPHIC REVISION

OF THE

Family CJKO.^IYIDtE

(Exclusive of the species of Thomomya)

15 Y

Dr. C. HART MERRIAM

WASIIIKaTOX
aoVERNMKNT PliINTH>I(J OFFICE

1805

LETTER OF TRANSMITTx\L.

U. S. Department of Agriculture,

Washington, D. C, September J26, 1894.
Sir : 1 have the honor to transmit herewith, as No. 8 of North Ameri-
can Fauna, a Monograi^hic Kevision of the Family Geomyida', exchisive
of the species of Thomomys.

In preparing a bulletin on the economic relations of the Pocket
Gophers it became necessary to determine the status and geographic
distribution of the various forms. This study developed the fact that
the group was sorely in need of technical revision. The present paper
is the outgrowth of an attempt at such a revision. It has grown so far
beyond the limits originally intended that a large genus {Thomomys) has
been of necessity omitted and will form the subject of a subsequent
paper.

The results of the economic study of the group will appear as a sepa-
rate bulletin prepared by my assistant, Mr. Vernon Bailey.
Respectfully,

C. Hart Merriam,

Chief of Division of
Ornithology and Mammalogy.

Hon. Chas. W. Dabney, Jr.,

Acting Secretary of Agriculture.

CONTENTS.

INTRODUCTION.

PHpe.

Material studied 11

AcknowJedgnieuts 12

Illustrations 12

CHAPTER I.— GENERAL REMARKS.

Adaptation to a subterranean life 15

Progression backward as well as forward 16

The tail an organ of touch 16

Normal position of the fore feet 17

Division of the mouth into two chambers 17

The tongue 18

The cheek pouches 18

How food is put into the cheek pouches 18

Th c f oo d 19

Color phases 19

Sexual variation 20

Individual variation 21

Subdivisions of the family Geomyida^ 22

Key to the genera 23

Phylogenetic tree of the genera 24

List of the genera and species 25

Ccograi>hic distribution of the family and genera 26

Number and distribution of the species 27

The United States species 28

Distribution of the Mexican species 30

Weight of characters 32

List of specimens examined 33

CHAPTER II.— MORPHOLOGY OF THE SKULL.

The cranium as a whole 33

The individual bones 40

Changes with age 61

Coi'ssification of the paired bones 63

Cranial variations — departures from the trunk line 63

CHAPTER III.— THE DENTAL ARMATURE.

THK TEETH.

Incisors 70

Premolars 72

Molars 74

Variation in form of m^ 76

6 CONTENTS.

Page.

Arrangement of the enamel 78

Principal divisions indicated by the enamel plates 79

Normal number of enamel plates 79

Variation in enamel plates of m^ 79

Characters of the unworn teeth 83

Incisors 83

Deciduous premolars 83

Permanent premolars 84

Molars 85

Summary 85

Changes in form and enamel pattern of young teeth with wear 86

The enamel organ 87

The osteodentine 87

MECHANISM AND DYNAMICS OF THE CUTTING MACHINE.

Manner of attachment of the teeth 88

Dynamics of the incisors 89

Dynamics of the molariform teeth 90

(a) Manner of implantation and curvatures 90

(6) Influence of the direction of the jaw movement on the molariform teeth. 92

(1) Effect on the size and curvature of the jirisms 92

(2) Eftect on the proportions of the prisms 93

(3) Ertect on the enamel plates 93

Arrangement and mode of operation of the cutting blades 93

(a) Dominant movement of jaw obliquely transverse 93

(6) Dominant movement of jaw antero-posterior 96

Treatment and course of food 98

Muscles that operate the cutting machine 98

Muscles of the cheek pouches 101

Muscles connecting the head with the neck 102

Aualj'sis of jaw movements 102

Influence of the masseter muscle in molding the skull and modifying the

teeth 104

Efl'ects on the skull 105

Ett'ects on the teeth 106

CHAPTER IV.— SYSTEMATIC DESCRIPTIONS OF THE GENERA AND

SPECIES.

Geumys 109

Pappogtomys 145

CyatoyeumijH 149

Plati/yeomys 162

Orthogeomys 171

Heterogeoiiiy^ 179

Macrogeoinys 185

Zygogcomys 195

Thomomys 198

APPENDIX.

(A.) STATUS OF GEOMYS MEXICANUS AiXT.
(B.) TABLES OF AVERAGE MEASUREMENTS OF THE VARIOUS SPECIES.

1. Of the species of Geomys 202

2. Of the species of Cratogeomya 203

3. Of the species of Platygeontys, Orthogeomys, Ueterogeomya, Pappogeomys, and

Zygogeomys 203

CONTENTS. 7

(C.) TABLES OF CRANIAL MEASUREMENTS.

Table A. Of Geomijs hiirsarius, lufesceiis, hreviceps, sagittaUs, and attwateri .. 204

B. Of Geomys personatus, faUax, iexensis, and areiiariits 206

C. Of Geomys tuza, floridanus, mobilensis, and Zygogeomys trichopud ..... . 208

D. Of the species of Cratogeomys 210

£. Of the species of Platygeomys 212

F. Of thespeciesot' Pappogeomys, Orthogeomys, Macrogeon>ys,audHetero-

geomys ......;.;......;... ^ ^ .. i . i ;...;.;; 214

ILLUSTRATIONS.

PLATES.

Frontispiece, Geomys tuza (Ord).

1. Skull of Geomys biirsarius.

2. Slvull of Cratogeomys mcrriaiiiL

3. Slcull of Platygeomys gymiiuriis.
4 SknU of Heterogeomyshitifidiis.

5. Skull of Macrogeomys dolicliocephahts,

6. Skull of Zygogeomys trichopns.

7. Skulls of Geomys tuza, mobilensis, and floridanus.

8. Skulls of Cratogeomys oreocetes, j)ercgrinns, estor, and perotevsis.

9. Skulls of Geonii/s arenarius, iexensis, attwateri, sagittalis, liitescens, hrevicejys, and

bitrsarius.

10. Lower jaws of Geomys tuza, floridanus, mobilensis, bursarius, Cratogeomys oreocetes,

peregrinus, merriami; Macrogeomys dolichoceplialus, and Platygeomys gymnurns.

11. Skulls of Pappogeomys huUeri, Macrogeomys heterodns, vostaricensis, Platygeomys

fumosus, Orthogeomys latifrons.

12. Skulls of Cratogeomys castanops, fulvescens, and Geomys personatus.

13. Left zygoma, showing variations in jugal bone in the various genera and species.

14. Palatopterygoids, showing variations in the various genera and species.

15. Skulls of Macrogeomys cherriei and Helerogeomys torridus. Occiput of Macrogeomys

dolichoceplialus, Heterogeomys hispidus, Platygeomys bulleri, Cratogeomys merriami,
Platygeomys gymnurns. Uiiper incisors of Macrogeomys dolichocephalus, Crato-
geomys merriami, Zygogeomys trichopns, Geomys bursarius and tuza.

16. Molariform teeth. Heterogeomys torridus, young, showing deciduous premolar in

situ; also upper permanent premolar showing unworn enamel cap; also same
showing permanent enamel pattern.

Geomys bursarius, showing deciduous premolar in situ; also crowns of molari-
form series showing permanent enamel ])attern.

Macrogeomys lieterodus, right upper ijremolar of adult.

Zygogeomys trichopns, crowns of molariform series showing permanent enamel
pattern.

Cratogeomys castanops, enamel pattern of molar crowns in young and adult.

Macrogeomys cherriei, young crowns of molariform series, showing permanent
enamel pattern.

17. Skulls seen from above: vault of cranium cut away, showing floor of brain case

■ in —

Heterogeomys torridus, Zygogeomys trichopus, Geomys bursarius, Platygeomys gymnu-
rus, and Cratogeomys merriami.

18. Vertical median longitudinal section of skull (mesethmoid and right half of

vomer in place) —
Geomys bursarius, Zygogeomys trichopus, Heterogeomys torridus, Cratogeomys mer-
riami, and Platygeomys gymnnrus.

19. Orthogeomys scalops 9 ad- skull from above, and base of cranium. Median longi-

tudinal section of nasal chamber (vomer and mesethmoid removed) showing
turbinated bones in —
Geomys bursarius, Zygogeomys trichopns, Heteroqeomys torridus, Cratogeomys mer-
riami, Platygeomys gymnurus.

8 CONTENTS.

TEXT FIGURES.

1. Face of Oeomj/s bursarius, showing grooved upper incisors and openings of cheek

ponches.

2. Face of Thomomjis talpoUJcs, showing phmo uppei' incisors and openings of cheek

pouches.

3. Left fore foot of Geomyii personatua, s\\ov,i\\g the rows of bristles which form

brushes on the sides of the toes.

4. Side view of skull of Cratogcomiis merriami (zygoma sawed off).

5. Basioccipital of Cratogeomtjs merriami. showing difference in form of upper and

lower surfaces (ankylosed exoccipitals shown also).

6. Variations in interparietal: Plnfi/ficimiis tiilorlnnna (showing changes with age);

Geomys tii:a ^ ad. (Augusta, (ia.); G. mobilensis J yg. ad. and ^ old (Milton,
Fla.). All natural si/e.

7. Longitudinal vertical median section of skull of Cratofjeomijs merriami, showing

interior of brain case and nasal chamber. Vomer and mesethmoid in place.

8. Very young skull of Geomys bursarius from Elk River, Minnesota. Upper sur-

face, showing troutals ankylosed togetlier, and interparietal inseparable from
sniiraoccipital.

9. Young skull of Cratofjeomys merriami, vault of cranium cut away to show floor of

brain case.

10. Vertical median section of front part of skull of Geomys bitrsarins, showing tur-

binated bones, etc. (mesethmoid and vomer removed).

11. Principal types of pa.latoi)terygoids.

12. Inferior surface of young skull of Craiocjeomys merriami.

13. Longitudinal vertical section of nasal chamber of Cratogeomys merriami, showing

vomerine sheath (vomer rcnu)ved).

14. Jugals (showing principal types of form).

15. Three skulls of Zygogeomys trivhopus, showing changes wiih age.

16. Very young skull of Heterogeomys torridiis from Motzorongo, Vera Cruz.

17. Types of frontal: (1) Cratogeomys merriami, (2) Ilelerogcomys torridits, (3) Macro-

(/eomys heierodns, (4) Vrthoijeomys sealops.

18. Outline of skull of Platygeomys gyr/Diurxs, slunviug teeth in situ.

19. Incisors of Platygeomys gymnnru^ sein from behind.

20. Cross section of upjicr incisor in (1) Macroiieomys doliclincephalns; (2) Heterogeo-

mys Itispidiis; (3) M.coslariceiisis; (4) M. vherriei (showing enamel face and sin-
gle sulcus).

21. Cross section of upper incisor in (1) Cratogeomys merriami ; (2) Platygeomys (jym-

nnriis; (3) Cratogeomys perote)isis ; (4) J'appogeomys htiUeri.

22. Cross section of upjier incisor in bisulcate series: (1) Zygogeomys triclioptis; (2)

Geomys hitrsari)is ; (3) Geontys iuza.

23. Cross section of upper incisor of Thomomys doiiglasi, showing shallow sulcus close

to inner side of tooth.

24. Cross section of lower incisor of Cratogeomys merriami.

25. Crowns of upper and lower premolars of Macrogeomys dolichoeephalus.

26. Types of molariform teeth seen in profile: (1) Heterogeomys hispidns ; (2) Crato-

geomys merriami ; (3) Geomys tnza.

27. Principal types of crown pattern of m': (1) Geomys brericeps ; (2) Pappogeomys

huJleri; (3) Platygeomys gymiiurns : (4) Cratogeomys estor : (5) Zygogeomys tricho-
pus ; (6) Macrogeomys dolichoeephalus ; (7) Macrogeomys heterodas.

28. Variations in crown pattern of m^ in Cratogeomys fidveseens.

29. Variations in crown patteru of m^ in Cratogeomys castaiwps.

30. Types of enamel pattern of upper premolar : (1) Cratogeomys merriami ; (2) Hetero-

geomys hispidus.

31. Types of enamel pattern of u])per molariform teetli in the different groups: (1)

Geomys bursarius; (2) Cratogeomys caxlauops; (3) Zygogeomys trichopus; (4)
Macrogeomys cherriei ; (5) Thomomys hulhirorus.

32. Types of enamel pattern of lower molariform teeth: (a) Geo7nys bursarius; (b)

Thomomys bulbirorus. Except in Thomomys tlie enamel pattern is the same
throughout the family.

33. Types of enamel pattern of crown of ni' in the restricted genus Geomys.

.34. Types of enamel pattern of crown of m ■ in the several groups in which this tooth
is a double prism.

35. Variations in form of crown and enamel pattern of m^ in Plattigeomys and Crato-

geomys merriami.

36. Molariform teeth of a A'ery y(Ming Geomys bursarius. showing deciduous and per-

manent premolars in situ, and unworn crown of m' which has not yet reached
plane of crowns of other teeth.

37. Right lower unworn permanent \^xemoh\x oi Heterogeomys torridus: (1) inner or

lingual side;- (2) enamel cap from above.

CONTENTS. 9

ZS. Right last lowci' molar of Hctcror/eomiiK forridns, showing mi worn euamelcap and
rchitioiis oi' enamel and dentine lower down.

3D. Transverse section of skull of J'ttitjif/eomi/s gijmnnriis, showing manner of implan-
tation an I r(dations of molariform teeth.

40. Upper and lower molars of I'lati/gcoiinis </!i)tniunts in normal position, showing

angle of trnneation of crowns necessitating lateral movement in arc of circle.

41. Cross section of mandible of ridtiif/eovii/s (jymnurns, showing how roots of lUj and

■m:i straddle the incisor.

42. Profile view of lower premolar in Ma(ro(jromiis dolivhocephalus and PJatijgeomys

(/ipinninis.

43. Longitndinal section of molariform teeth of Plati/ficonn/s {lymnurus (diagram-

matic.)

44. Crowns of molariform teeth of riaii/neomijs i/jjmnitrxs.

45. Snperimposed molar series of Platiigeomijs (jymnurns, showing relations of enamel

blades.
4l!. Longitudinal section of molariform teeth of Macrof/eomys dolichoceplialus (dia-
grammatic.)

47. Crowns of molariform teeth of Macroyeo.mys dolicliocephahts.

48. Superimposed molar series of Alacrofjeomys doHchocephahis.

49. Siile view of sknll of Macrogeomys doHcltucephahis.

50. Side view of skull of Platyf/eomyx f/ymnitrits.

51. Hinder part of sknll of Macrogeomya dolichovephahis from above, showing relations

of mandible.
.52. Transverse vertical section of cranium of Macrogeomys dolichocephalns (just in
front of audita! bulhe) with mandible in place — viewed from behind.

53. Hinder part of skull of Platygeoiiiys gymnitrus from above, showing relations of

mandible.

54. Transverse vertical section of cranium of Platygeomtjs giimnnvus (just in front of

audital bulLe) with mandible in place — viewed from behind.

55. Geomys hnrsarius. Side view of skull, zygomatic arch sawed off to show bottom

of orbit.

56. Pappogcomys bullcrt $ . Vault of cranium sawed oft", showing floor of brain case.

From Sierra Nevada de Colima, Mexico.

57. Pappogeomys biiUeri. Vertical longitudinal section of skull, mesethmoid and

vomer in place. (Same specimen as 56.) XlA.

58. Pappogeomys huUeri. . Vertical longitudinal section of skull. Mesethmoid and

vomer removed to show eudoturbinals. (Same specimen as 56.) Xl^.

59. Craiogeomys merv'mmi. Crowns of molariform teeth.

60. Orthogcoiuiis svalojjs. Longitudinal vertical median section of skull. Meseth-

moid and vomer iu place. From Cerro San Felipe, Oaxaca. Xli.

61. Orthogcomys scalops. Same specimen with mesethmoid and vomer removed,

showing eudoturbinals. Xli.

62. Orthogeomys scalopn. Last upper molar showing divided outer enamel plate.

63. Orthogeomys velsoni. From Mount Zempoaltepec, Oaxaca, Mexico. Skull from

above. (Type).

64. Orthogeomys latifrons (type). Crowns of molariform teeth.

65. Heterogeomys hispidiis. l''roin Jico, Vera Cruz.

66. Heterogeomys lorridus. From Motzorougo, Vera Cruz.

67. Macrogeomys costaricensis and cherriei, showing differences in jugal, viewed from

both sides.
68,69,70,71. Tliomomys bidbiroriis. 9 From Salem, Oregon. Sectionized skull.

68. Vertical median longitudinal section; vomer and mesethmoid removed,

showing turbinated bones.

69. Same, mesethmoid and vomer in place.

70. Vault of cranium sawed off, showing floor of brain case.

71. Anterior part of floor of brain case, much enlarged.

Map 1. A Distribution of genus Thomomys.

]i Distribution of genus (ieomys.
Map 2. Distribution of genus Cratogeomys.
Map 3. 1 Distribution of genus Pappogeomys.

2 Distribution of genus Plaiygeomys.

3 Distribution of genus Zygogeomys.

4 Distribution of genus Heterogeomys.

5 Distribution of genus Orthogeomys.

6 Distribution of genus Macrogeomys.

Map 4. Distribution of species of Geomys and Cratogeomys

No. 8. NOETH AMEEIOAN FAUNA. January, 1895.

REVISION OF THE POCKET GOPHERS, FAMILY GEOMYID.E, EXCLUSIVE
OF THE SPECIES OF THOMOMYS.

By Dr. C. Hart Merriam.

INTRODUCTION.

The present paper is based on a study of the rich collection of Pocket
Gophers belonging to the U. S. Department of Agriculture, comprising
upwards of 800 specimens, exclusive of the genus Thomomys. This
material has been supplemented by 110 specimens from my private col-
lection, and a number from the U. S. National Museum,* making a total
of about a thousand specimens, among which are by far the greater num-
ber of actual types known to be extant. The Department collection
contains no less than 200 specimens from Mexico, most of which were
secured by Mr. E. W. Nelson, a field naturalist of the Division. These,
together with a few highly interesting sjiecimens from Costa Eica and
Guatemala in the U. S. National Museum, have enabled me not only
to bring together for actual comparison all of the species i)reviously
described, and to add a considerable number heretofore unknown, but
also to recognize several marked generic types whose existence had
not been suspected.

Critical study of this unparalleled wealth of material has led to the
discovery of some very remarkable dental peculiarities that have been
deemed worthy of detailed description and illustration. Moreover, the
opportunity has been utilized to contribute a chapter on the morphol-
ogy of the skull, which it is hoped will prove of service to those inter-
ested in the craniology of the Rodentia.

It is a matter of deep regret that the magnificent series of specimens
of living forms on which the present paper is based, has not been sup-
plemented by a single fossil ; and it is earnestly hoped that an oppor-
tunity may yet be found to study the remains of the extinct animals
that have been referred to the family — correctly or otherwise — in com-
parison with the rich collection of living types now in our National
Museum. If the theory is correct, that the group has attained its
greatest expansion in the present age, we need not look to the rocks

* Placed at my disposal by the courtesy of Mr. F. W. True, Curator of Mammals-

11

12 NORTH AMERICAN FAUNA. [no. 8.

for additioual liiglily diversified types, but only for the links tbat biud
the several phyla together and connect them with the more primitive
forms from which they came. These would be of the utmost interest.

The author is indebted to Mr. F. W. Tiue, Curator of Mammals in
the U. S. National Museum, for the privilege of describing two species
from Central America; to Dr. J. A. Allen, of the American Museum
of Natural History, New York, for the privilege of examining the type
of Ms Geoniys cherriei; and to Mr. H. P. Attwater, of San Antonio,
Texas, for the loan of a large series of the subspecies here described
as Geomi/s brericeps attwateri. The author is under special obligations
to Mr. Oldfield Thomas, Curator of Mammals in the British Museum,
and to Dr. Paul Matschie, of the Eoyal Museum of Natural History in
Berlin. Mr. Thomas has kindly compared specimens sent him for that
purpose with his own types in the British Museum, and has also con-
tributed measurements and other details of importance. Dr. Matschie
has been good enough to remeasure the original types of Peters'
Geomys heterodus and Lichtenstein's Geomys mexicanus, which speci-
mens are still extant in the Berlin Museum, and has further taken the
trouble to prepare and send me a table of cranial measurements of the
skulls of these same types, with much other information of importance
respecting them. And Dr. F. A. Jentink, the able director of the
Leiden Museum, has done me the favor to send additional particulars
about the Bullock specimen of Geomys hursarius, still extant in the
Leiden Museum, which specimen has given rise to much controversy
and is supposed to be Shaw's original type of the species.

From time to time during the preparation of the work, collectors
have been sent to special localities from which new or supplemental
material was desired, thus making it possible to settle many points that
were originally in doubt. Much has been learned respecting the habits
and mode of life of the animals from a living Geomys Jutescens sent from
Vernon, Texas, by my field assistant, Mr. J. Alden Loring. This animal
was kept in confinement until sufiQciently tame to permit handling freely
and was the means of the discovery of a surprisingly large number of
interesting facts that otherwise would have escaped detection.

Respecting the illustrations, the frontispiece was drawn by Mr. C. B.
Hudson; plate 1 by Mr. Benjamin Mortimer; text figures 1 and 2 by
Dr. George Marx ; figures 5, 19, 63, 65, and 66 by Dr. James E. McCon-
nell; and all of the outline camera lucida drawings of teeth by myself.
Plates 2 to 19, inclusive, and all of the remaining text figures were drawn
under my constant supervision by Mr. F. Miiller. All of the twenty
full-page plates have been reproduced by photolithography by Mr. Ber-
thold Meisel, of Boston, and the text figures, with two or three excep-
tions, have been electrotyped from the originals by Mr. Harry 0. Jones,
of New York.

It will be observed that the generic names engraved on most of the
plates (pis. 2-6, 8, and 10-16) do not agree with the generic names in the
text. This misfortune is the result of having the plates printed before

JAN., 1895.] INTRODUCTION. 13

the genera were finally segregated. The correct names are given in all
cases on the explanations facing the plates.

The literature relating to the group is rarely referred to in the present
paper, except for original descriptions. The reason is that previous
papers have been based on insufficient material. To use them at all
would necessitate a large amount of explanation and criticism without
corresponding advantage.

All the measurements in the present paper are in millimeters.

CHAPTER I.

GENERAL REMARKS.

The family Geomyid(v, comprising the mammals commonly known as
Pocket Gopliers, is conliued to North America, where it ranges from
the plains of the Saskatchewan in Can-
ada southward to Costa Rica. It attains
its highest development in tlie western
United States and Mexico, and does not
inhabit the region east of the Mississippi
Valley except in the Gulf States, where
it reaches the Atlantic coast in Florida
and Georgia, but does not occur north
of the Savannah River.

The appearance of a characteristic
species is well shown in the frontispiece,
and the peculiar aspect of the face in the
accompanying cut (tigs. 1 and 2), which
shows the openings of the cheek pouches,
wholly outside of the mouth, and also
the pattern of the upper incisor teeth in
two of the commonest genera, Oeomys
and Thomomys.

All the members of the family spend
their entire lives underground, and their
whole organization is modified in accord-
ance with the needs of a subterranean
existence. The species, though n numer-
ous, are very much alike externally. They are short-legged, thickset
animals, without an appreciable neck, without noticeable external
ears, and with very small eyes. The feet are largely developed for
digging. The fore paws in particular are very strong, are armed
with long curved claws,* and the sides of the toes are lined with
rows of bristles that evidently serve in preventing the dirt from

FiG.l. — Faceof Oeomys bur. ■iarius, show-
ing grooved upper incisors and openings
of cheek pouches.

Fi«. 2.— Face of Thomomys talpoides,
showing plane upper incisors and open-
ings of cheek pouclies.

*Tlie relative development of the daws is largely a matter of age and soil. Tbey
continue to increase in size thronghout the life of the individual; their poiiitw are
worn off in hard soil so that the claws become thick and hlunt. In sandy soil they
do not meet enough resistance to ])roduce the usual wear, and, conse(iuentIy, are
longer and more slender than normal,

15

16

NORTH AMERICAN FAUNA.

passiug between the fingers (fig. 3), thus completing a more effective
arrangement for keeping the tunnels clean, and for pushing the earth
out of the openings in the burrows. The tail, which is of moderate
length, is thick, fleshy, and usually devoid of hair, and is endowed
with tactile sensibility.

The Pocket Gophers, in working their way
through the earth in the construction of their tun-
nels, use the j)owerful upper incisors as a pick to
loosen the ground. At the same time the fore feet
are kept in active operation, both in digging and in
pressing the earth back under the body, and the hind
feet are used also in moving it still further backward.
When a sufficient quantity has accumulated behind
the animal, he immediately turns in the burrow and
by bringing the wrists together under the chin,
with the palms of the hands held vertically, forces
himself along by the hind feet, pushing the earth out
in front. When an opening in the tunnel is reached
the earth is discharged through it, forming a little
hillock that resembles in a general way the hills
thrown up by moles. In many species there is a
naked callosity or 'nasal pad' over the anterior half
of the nose, which must be of great assistance in
the construction of the tunnels. When this callos-
ity is largely developed the nasal bones underneath are highly arched or
inflated, as in Heterogcomys hispidus.

PROGRESSION BACKWARD AS WELL AS FORWARD.

The Geomys lutescem already mentioned from Vernon, Texas, which I
kept alive for several months, surprised me very much by running back-
ward as rapidly and easily as forward. This method of progression was
particularly noticeable when the animal was in his own quarters where
he could follow a runway or an accustomed route. When carrying
food to one of his storehouses he rarely turned around, but usually ran
backward to the j)lace of de^wsit, returning for more, and repeating the
operation again and again, the to-and-fro movement suggesting a shut-
tle on its track. The well-known peculiarity of the external genitalia,
which are so hidden and modified that the sexes are determined with
difficulty, is doubtless the result of this habit, protecting the parts
from injury when the animal is moving backward.

THE TAIL AN ORGAN OF TOUCH.

Throughout the family Gcomyidw the tail is rather large and fleshy,
and as a rule is naked or scantily haired; * it varies in length in the

* The tail is naked in most of the southern species and is more or less covered with
hair in the northern species. The latter have much more hair on the tail in winter
than in Rummer,

ria. 3. — Left fore foot of
Geomys personatus, sliow-
iijg the rows of bristles
which form brushes on
the sides of the toes.

JAN , 1895

GENERAL CHARACTERS. 17

various species from about 65 to 115 mm. The functiou of this pecu-
liar appendage had long puzzled me, but by watching- the live Geomys
above mentioned as it ran backward in its runways I saw that it was
used as an organ of touch. It is doubtless endowed with special tactile
sensibility and is evidently of great value in warning the animals of
the presence of an enemy in the rear when they are traveling backward
in their dark tunnels. So far as I am aware this is the only instance in
which the tail of a mammal is used for this purpose.

i POSITION OF THE FORE FEET.

In walking on soft ground the fore feet are usually held in the nor-
mal position, with the soles down, or inclined shghtly inward. In
walking on hard ground, however, the fore feet are turned sideways,
their soles facing one another, so that the claws curve inward, and the
animal walks on the outer or ulnar side of the foot. This method of
using the fore foot in walking on hard substances was frequently
observed, and enables the animal to walk comfortably where the long
curved claws would be in the way A held in the normal position. It
was also frequently noticed that the feet were held in the same jjositioii
(horizontally) when at rest, and when used as a scoop in pushing loads
of earth or sand out of the way. When thus engaged the feet were
drawn back under the breast, the wrists near together and the long
claws turned outward. By moving the body quickly forward the
animal always succeeded in throwing ahead of it a considerable quan-
tity of loose earth.

DIVISION OF THE MOUTH INTO TWO CHAMBERS.

The lips and thin furry covering of skin are drawn into the broad
space between the incisors and molars, where they meet in a raphe on
the roof of the mouth and then separate again to meet around the under
jaw, forming a diaphragm-like partition between the incisors and
molars. The orifice is small and wholly inferior, and may be completely
closed by the fleshy tongue or by the tailing together of the furry lips,
leaving a vertical slit between. The raphe or line of union of the lips
on the roof of the mouth reaches most of the way from the incisors to
the upper premolars, A narrow band, not covered with fur, connects
the two lips inferiorly, crossing the floor of the mouth near the pos-
terior end of the synqihysis of the mandible. Thence the lips — if the
term lips may be applied to this fold of skin — pass around the lower
incisors, where the skin is attached posteriorly, so that it may be
retracted, leaving a bare space below the point where the tooth pro-
trudes from the alveolus, thus giving greater freedom to the lower
incisors during tlie act of gnawing. During the to-and-fro drill-like
motion of the jaw the skin probably remains nearly stationary, while
the under incisors play rapidly back and forth. The object of the dia-
7i33— No. 8 2

18 NORTH AMERICAN FAUNA. [no. 8.

phragm-like partition which separates the mouth into two chambers is
obviously to prevent dirt and chips from entering the mouth proj)er
during the various subterranean operations of the animal.

THE TONGUE.

The tongue is short, thick, and fleshy. Its principal function doubt-
less is to keep the food between the crowns of the teeth during masti-
cation. At other times it serves as a plug to stop the opening in the
furry diaphragm between the incisors and molars.

THE CHEEK POUCHES.

All of the Pocket Gophers are provided with external cheek pouches,
which open on the sides of the face outside of the mouth, and are cov-
ered with fur inside. These cheek pouches are used exclusively in
carrying food, and not in carting dirt as often erroneously supposed.
The animals are great hoarders and carry away to their storehouses
vastly more than they consume. The cheek pouches reach back as far
as the shoulder and are so attached that they can iiot be completely
everted without rupture of their connections. While the posterior
part of the sack is held back by the muscle which stretches thence to
the lumbar vertebra", the skin of the inner side of the jiouch, which
covers the side of the face below the eye and in front of the ear, may
be everted or prolapsed, hanging down as a flap below the corners of
the mouth. This is probably what happened in the case of snake fright
observed by Dr. A. K. Fisher at Ellis, Kansas, in June, 1893. Dr.
Fisher saw a gopher snake {Pituo2)hi.s) about 5 feet in length hunting
for breakfast. He says: "Presently the snake glided into a gopher
hole. In a few minutes I saw a gopher {Geoniys lutescens) run out as
fast as possible from the other end of the line of hills. I soon caught
up to it. It appeared very much frightened, and its cheek pouches
were hanging out. The gopher evidently had only scented the snake,
for it Avas apparent that the snake had not seen the mammal, as it
came out of the hole by which it entered and glided off deliberately
in another direction."

HOW CIEOMYS PUTS FOOD INTO ITS CHEEK POUCHES.

A live Geoniys from Vernon, Texas, has been carefully observed for the
purpose of ascertaining how the reserve food is placed in the cheek
pouches. The animal soon became sufficiently tame to eat freely from
the hand, and was commonly fed bits of potato, of which he was particu-
larly fond. The manner of eating was peculiar and interesting, and
showed an ability to use the huge fore feet and claws in a way previously
unsuspected. After satisfying the immediate demands of hunger it was
his practice to fill one or both cheek pouclies. His motions were so
swift that it was exceedinglj^ difficult to follow them with sufficient
exactness to see just how the operation Avas performed. If a whole

'AN, 1895] GENERAL CHARACTERS. 19

jiotato was given bim, or apiece too large to go into the pouch, he invari-
ably grasped it between the fore paws and proceeded to pry off a small
Iticce with the long lower incisors. He would then raise himself slightly
on his hind legs and hold the fragment between his fore paws while
eating, for he usually ate a certain quantity before putting any into the
pouches. If small pieces were given him he took tliem prom^itly and
passed them quickly into the pouches. Some pieces were thus disposed
of at once; others were first trimmed by biting off projecting angles.
As a rule one pouch was filled at a time, though not always, and the
hand of the same side was used to push the food in. The usual course
is as follows : A piece of potato, root, or other food is seized between the
incisor teeth, and is immediately transferred to the fore paws, which are
held in a horizontal position, the tips of the claws curving toward one
another. If the food requires reduction in size, the trimming is done
while held in this position. The piece is then passed rapidly across the
side of the face with a sort of wiping motion which forces it into the
open mouth of the pouch. Sometimes a single rapid stroke with one
hand is sufficient; at other times both hands are used, particularly if the
piece is large. In such cases the long claws of one hand are used to
draw down the lower side of the opening, wliile the food is poked in with
the other. It is obviously impossible for the animal to i)ass food from
the mouth to the pouches without the aid of its fore claws.

The most remarkable thing connected, with the use of the pouches is
the way they are emptied. The fore feet are brought back simulta-
neously along the sides of the head until they reach a point opposite the
hinder end of the pouches; they are then pressed firmly against the
head and carried rapidly forward. In this way the contents of the
pouches are promptly dumped in front of the animal. Sometimes several
strokes are necessary. I am not prepared to say that the animal can
not empty the pouches by means of the delicate investing muscles, but
I have never seen them emptied in any other way than that here
described.

THE FOOD.

The food consists chiefly of roots, tubers, and other rather hard veg-
etable substances, though grass and the. succulent parts of plants are
sometimes eaten. In agricultural districts the animals are highly inju-
rious, destroying potatoes and other tubers in large quantities, and
gnawing off" the roots of fruit trees. In fields of grain and fodder they
sometimes do considerable damage by the aggregate area covered by
the little mounds of earth thrown up from the tunnels.

COLOR PHASES.

In- most species of the Geomyidw two color phases occur, a plumbeous
or dusky phase and a chestnut-brown or yellowish-brown phase. The
latter varies greatly in the different species — from pale straw color or

20 NORTH AMERICAN FAUNA. [no. 8.

buffy ochraceous in Tliomoinys perpallidus of the Colorado and Mohave
deserts, to dark liver-brown in Geomys bursarius of the Ui)per Missis-
sippi Valley. Taking the group as a whole, the brown phase is by far
the commonest and may be regarded as normal ; but in certain vSpecies
in nearly all the genera the plumbeous i)hase prevails, as in Thomomys
Orizaba' J Platygcomys fumosus, and Zygogeomys tricJiopus — all from south-
ern Mexico. The plumbeous pelage is commonly more or less metallic
antl sometimes even iridescent. It is rare in the United States species,
though conmion in Thomomys neradensis from central Nevada and
Geomys breviceps from Louisiana. It has not yet been observed in Gra-
iogeomys castaiwps or Geomys lutescens, and the red pelage has not been
observed in Zygogeomys tricliopus. So far as known, only a single color
phase occurs in the genera Heferogeomys and Orthogeomys, both of
which are dark seal brown in fresh pelage and a dull faded brown in
worn iDclage.

Seasonal differences in coloration. — Some of the species vary but
little with season, as Geomys bursarius from the Upper Mississippi Val-
ley; still even this animal is considerably darker in winter than in
summer. Others present two well-marked color phases, according to
season. In the latter category are Geomys lutescens, breviceps, and to a
less degree personatus also. In lutescens the summer pelage differs
from the winter in the absence of the dark dorsal band which is usually
present from October to April, or May, and sometimes even as late as
June. Apparently the absence of this stripe in summer specimens is
sometimes due to wear, the dark tips of the hairs when worn leaving
the pale subapical zone exposed. This can not always be the case,
however, since one specimen from Chadron, Nebraska, collected April
30, has the dorsal stripe plumbeous throughout with but a faint trace
of the pale.subapical zone.

In typical Geomys breviceps, and also in specimens from the western
limit of the range of the species wliere it seems to be shading toward
lutescens and texensis, the same thing occurs, though the renewal of tlie
pelage takes place at a somewhat different date. This is very apparent
in si)ecimens from Gainesville, in the valley of the Ked River in north-
eastern Texas. A specimen taken August 10 has a broad dark dorsal
ban(i, while two specimens taken March 27 and March 29 show no
trace of this band except on the head, the back being a uniform red-
dish brown more or less mixed with dusky.

SEXUAL VARIATION.

Sexual variation is marked.throughout the genus and in some species
is extraordinary. It may "be conveniently discussed under two heads,
(1) difference in size: (2) ditference in cranial characters.

(1) Difference in size. — ^Tlie females are always considerably smaller
than the males; the discrepancy is greater in some species than in
others. Reference to tbe tables of measurements shows that the dif-

JAN.. 1895.1 VARIATIONS. 21

fereuce in total length often amounts to 25 or 30 mm.; in length of tail
to 12 or 15 mm. ; and in hind foot 3 or 5 mm. Tlie difference in the size
of the skull is equally marked, and is well shown in the tables of
cranial measurements.

(2) Difference in cranial characters.* — Independent of the conspicu-
ous differences in size between male and female skulls of the same
species from the same locality, other and more important differences
exist which not infrequently prove troublesome in identifying speci-
mens, particularly if skulls of both sexes are not at hand for compari-
son. The female as a rule has the brain case broader and flatter, the
zygomata narrower and less angular, the jugal narrower anteriorly,
the rostrum and nasals shorter, and the skull as a wliole smoother. In
other words, the cranium of the female is much less specialized than
that of the male and often jioints suggestively to the stock from which
the species was derived. It thus happens in the case of series of
species in which the successive forms in the development of a particu-
lar type are still extant (as in the texensis-hursarius series) that the
female resembles the male of the species next below in the line of
descent more than the male of her own species.

In several forms in wiiich the males have well developed sagittal
crests, the females have a sagittal area bounded by distant temporal
impressions; and in species in which the males have iirominent tem-
poral ribs, the females commonly have more widely separated temporal
impressions which rise as ridges from the outer side but not from the
inner side, the interspace being more or less thickened.

INDIVIDUAL VARIATION.

The family Geomyida' i^resents the usual range of individual varia-
tion, both in size and in cranial characters. While the male and female
skulls of a species agree very well among themselves, showing strong
average characters, there are in every large series one or two skulls
which depart from the type in one or more particulars. These depart-
ures are most common in the form aud manner of ending of the nasals
and ascending branches of the premaxilla. In all such cases sexnal
differences should be carefully eliminated before assuming that the
departure is individual.

Individual variation is always more marked in the secondary or acces-
sory cranial structures than in the more important and less variable
elements. Thus the peripheral processes aud expansions for the attach-
ment of muscles are always more variable than other parts of the skull.
The degree of lateral production of the squamosal, and of the angular
process of the mandible in Platygeomys gymnurus, and the variations in

' The sexual organs are so arranged in the Gcomuidw as to be difficult of determi-
nation in the flesh, except during the rutting season ; hence the sex marks on labels
may be safely ignored if they coullict with the cranial characters.

22 NORTH AMERICAN FAUNA. [no. 8.

detail of the occipital basiu, are illustrations of this kind. Still, in
studying" large series of skulls of a single species, one is inucli more
deeply impressed by the strong tendency toward the development in each
bone of a particular type of form than by the departures therefrom.

The animals continue to grow for several years, so that the majority
of breeding individuals are still far from the full size of their species.
This is very apparent in the skulls, which not only continue to increase
in actual size but also, in many species, in the ratio of zygomatic breadth
to length, and in the development of ridges and processes for muscular
attachments.

SUBDIVISIONS OF THE FAMILY GEOMYID^.

A superficial examination of the skulls of the various species of
Geomyidw is sufficient to show the existence of several widely different
types. Heretofore tlie common practice has been to divide the family
into two genera, Thomomys and Geomys, according to the absence or
presence of distinct grooves in the upper incisors, and to subdivide the
genus Geomys into two series, unisulcate and bisulcate. The number
of grooves was believed to be correlated with certain cranial characters,
the members of the unisulcate series having widely spreading zygomatic
arches, the outer angles of which were broadly exjianded, while the
bisulcate series had narrower arches and lacked the expansion; but no
attempt was made to separate them, even subgenerically. The recent
discovery of a large number of new forms in Mexico and Central Amer-
ica, comprising several highly diversified types, renders this classifica-
tion inadequate. After subtracting the strongly marked genus Thom-
ojiiys, which differs from all the others in numerous important charac-
ters heretofore overlooked, a heterogeneous assemblage remains, com-
prising the animals commonly lumped under the generic name Geomys,
and also the new forms here first described. Of these, the bisulcate
series may be divided into two very distinct and two minor types, while
the unisulcate series contains at least six well-marked subdivisions.

In attempting a logical classification of the group, one is met at the
outset by the difficulty that some of the specialized or peripheral types
are more or less closely connected with the trunk line by existing inter-
mediate forms, making it exceedingly difficult to draw hard and fast
lines without unnecessary subdivision. The genus Geomys as here
restricted is such a case. It comprises two quite distinct branches,
Geomys tuza and G. hursarius, which are connected with one another
and with the trunk line, or something very near it, by a series of gen-
eralized species, the texensis-hreviceps series. In cases of this kind two
courses are open, either to separate the extreme peripheral forms from
the less specialized species leading up to them, or to unite the entire
branch under a single getuis. The latter course has been followed in
the present instance. But each case must be decided on its merits.
One that has been treated differently is the large limb whose ends,
as now" known, are represented by two of Mr. Thomas's species, hulleri

JAN, 1895.] KEY TO GENERA. 23

and merriaml; tlie former is not far removetl from the trauk line of tbe
group; tbe latter is one of the terminal branches. Bat the two forms
differ in cranial and dental characters of great weight, and are further-
more separated by an enormo.us gap which is not bridged at any point
by any of the species yet discovered. For these reasons they are treated
as independent genera. Still another reason for this course, if another
were needed, is the circumstance that the branch ending in merriami is
only one of four- equally specialized terminal boughs, all apparently
springing from and bearing the same relation to the single limb or main
stem whose base is marked by b idler i.

In dividing the family into genera the aim has been to select as types
the most specialized j)eripheral forms and to assemble around them the
less specialized species. A study of the enamel pattern of the molari-
form teeth shows that the Geomijidcc maybe divided primarily into five
groups, several of which are of supergeneric value, and a study of the
fundamental cranial characters leads to the recognition of nine genera.
By means of the following brief key, any of the species now known may
be easily referred to its proper genus without cutting the skull:

KEY TO GENERA.
d) NO ENAMEL ON POSTEKIOK FACE OF UPPER PREMOLAR.

posterior enamel plate present on first and second upper molars.

Upper incisor bisulcate Geomys.

Upper iucisor uuisulcate

Frontal strongly constricted (biconcave) between orbits Pappogeomys.

Fiontal not constricted between orbits; very broad *Orthogeomys.

Posterior enamel jtlate absent In first and second upper molars.

Breadth of cranium across squamosals much less than zygomatic
breadth ; lambdoid crest not sinuous (simply convex pos-
teriorly) ; angle of mandible short • Cratogeomys.

Breadth of cranium across S([uamosals greater than zygomatic
breadth; lambdoid crest strongly sinuous; angle of man-
dible very long Plutygeomys.

(2) ENAMEL PRESENT OX POSTERIOR FACE OF UPPER PREMOLAR.

Posterior enamel plate of upper premolar restricted to inner side.
Posterior enamel plate present and complete on first and second upper
molars.
Frontal not constricted between orbits; very broad; pterygoids

long * Orthogeomys.

Frontal strongly constricted between orbits ; pterygoids short.
Postorbital process absent ; palatopterygoids long and slen-
der (pterygoid part narrow) Heterogeomys.

Postorbital process strongly marked ; palatopterygoids short

and broad (pterygoid part very broad) Macrogeomys.

* Orthogeomys presents an exceptional condition of the enamel pattern of the upper
premolar. The posterior enamel plate of this tooth is evidently disappearing; it is
present on the inner side in O. latifrons, hut is altogether absent or reduced to a
very narrow strip in O. grandis and acalops.

24

NORTH AMERICAN FAUNA.

Posterior enamel plate of upper premolar complete.

Posterior enamel plate i^rescnt on inner {liniinal) sich- onhj of first and
second upper molars.
Zygomatic arch couiplete without jugal (jugal inferior); incisors

bisulcate Zygogeomys.

Posterior enamel j^l'ite present and complete on first, second, and third
upper molars.
Incisors not grooved, or with a single fine sulcus on inner side. . Tliomomys.

PHYLOGENETIC TREE OF THE GENERA.

The accompauymg- pliylogeiietic tree is iutended to represent the
author's conception of the interrelations of the nine living genera of

\ HlacrogeorTLys

Phylogenetic tree of tho GeoiiiyidiT>.

the Geomyidcc now known. It is introduced with a full knowledge of
the modern tendency to disregard and even belittle such attempts; but
I am aware of no way in whicli tlie results of painstaking research
respecting the afitinities of organisms may be expressed so graphically.
Apparently there were four forks to the early Paleo-Geomine phylum:
one running into Tliomomys, another producing the bisulcate series of
Geomys, beginning with texensis or arenarius and ending in bursarius;
the third developing the anomalous bisulcate Zygofjeomys; the fourth,
a strictly unisulcate series, of which hulleri and albinasus are the least
specialized forms now known, splitting into four very distinct branches,
each of which now forms a well-niarked genus. In the case of the

JAN., 1895.] LIST OF GENERA AND SPECIES. 25

branch leading np to Geomys biirsarius the series of living forms is
practically complete; in the case of the other branches the connecting
links are unknown. It is evident that both Pappogeomys bulleri and
Geomys texensis branched off from i^oints not very remote from the
l^lace where Thomomys left the trunk line, and that they have under-
gone relatively little modification since.

The evolution of some types takes place in a very direct way, appar-
ent?ly by uninterrupted progress in a definite direction, and the species
comprising such a series, as texensis, breviceps, luteseens, and bursariiis,
may be looked upon as stages in the evolution of the type. The origin
of other types is more circuitous and less easily understood. For-
tuitous variations lead to the appearance of numerous side branches,
most of which abort before developing any very pronounced individ-
uality. Others are moi-e fortunate. Chancing to fit some phase of the
environment previously unutilized, they go on until a maximum of
departure compatible with the balance of the organism as a whole is
attained. There are several of these highly specialized departures
from the main stem in the GeomyidWj such as Cratogeomys, Platygeomys ,
Macrogeomys, and Zygogeomys.

LIST OF THE GENERA AND SPECIES.

Genus Gkomys Rafiuesque.

Name of species. Type locality.

Geomys tuza (Ord) Augusta, Georgia.

tiiza floridanns (Aud. and Bach.) St. Augustine, Florida.

tuza mohilensis'Siihs]). nov Mobile Bay, Alabama.

hursariits (Shaw) Minnesota?

lutescen sMevii-din Western Nebraska.

hreviceps Baird Mer Rouge, Louisiana.

hrevicepssagittalis subsp. nov Galveston Bay, Texas.

breviceps attwaieri siibsp. nov Rockport, Aransas County, Texas.

teidensis sp. nov Mason, Texas.

arenarius sp. nov El Paso, Texas.

personatiis True Padre Island, Texas.

personatus fallax suhs]). nov Corpus Christi, Texas.

Genus Pappogeomys nob.

Pappogcomys huJIeri (Thomas) Talpa, Mascota, .lalisco, Mexico.

alhiiiasns sji. uov Guadalajara, Jalisco, Mexico.

Genus Cratogeomys nob.

Cratogeomys merriami (Thomas) Valley of Mexico.

perotcnsis &i>. nov Cofre de Perote, Mexico.

estor sp. nov Las Vigas, Vera Cruz, Mexico.

peregrinus sp. nov Mount Iztaccihuatl, Mexico.

oreocetes sp. nov Mount Popocatapetl, Mexico.

castanops (Baird) Las Animas, Colorado.

castavops goldmaui subsp. nov. . .Canitas, Zacatecas, Mexico.
fulvescens sp. uov Chalchiconmla, Puebla, Mexico.

26 NORTH AMERICAN FAUNA. [no. 8

Genus Platygkomys nob.

Platijgeomijs gymnurus Merriam , Zapotlan, Jalisco, Mexico.

fjllorhinus sp. nov Tula, Hidalgo, Mexico.

planiceps sp. nov Northern slope Volcau Toluca, Mexico.

fumosHs Merriam Colima City, Colima, Mexico.

Genus Ohthogeomys nob.

Orthogeomys scaJops (Thomas) Teliuantepec, Mexico.

grandis (Thomas) Duefiaa, Guatemala.

latifrons sp. nov Guatemala.

nelsoni sp. nov Mt. Zempoaltepec, Oaxaca, Mexico.

Genus Hkterogeomy'S nob.

Heterogeomys hispidus (LeConte) Near Jalapa, Vera Cruz, Mexico.

torridus sp. nov Chichicaxtle, Vera Cruz, Mexico.

Genus Macrogeomys nob.

Macrogeomys heterodus (Peters) Costa Rica.

dolichocephalns sp. nov San Jose, Costa Rica.

costaricensis sp. nov Pacuare, Costa Rica.

chcrriei (Allen) Santa Clara, Costa Rica.

Genus Zy^gogeomys nob.

Zygogeomys trichoptis sp. nov Nahuatzin, Michoacan, Mexico.

GEOGRAPHIC DISTRIBUTION OF THE FAMILY AND GENERA.

The area inhabited by the family Geomyidw stretches from the dry
interior of British Columbia and the Plains of the Saskatchewan south-
ward to Costa Rica. In an east and west direction the grouj) covers
the continent from ocean to ocean, except that it is absent from the
region north of the Savannah River and east of the Mississippi Valley,
as shown by the accompanying maps (maps 1, 13, and 3). The group is
clearly of Sonoran origin and reaches its highest development on the
sonthern r>art of the table-land of Mexico. The great majority of the
species inhabit the upi)er and lower Sonoran zones, though a few
specially modified forms range upward on favorable mountain sides
through the Transition and even into the lower edge of the Boreal zone.
On the other hand, two species inhabit the tropical belt of Mexico.

Distribution by (jcnera. — The present distribution of the genera coin-
cides very nicely with their systematic relations.

The genus Thomomys (ma]) 1, A) has by far the most extended range
of any single genus, inhabitating suitable localities from the valley of
Mexico and Mount Orizaba northward to British Columbia and the
North Saskatchewan river, and from the Pacific coast eastward to the
Great Plains.

The genus Geomys (map 1, B and B' ) inhabits a broad belt across the
middle part of the United States, from the Red River Valley in north-
western Minnesota and northeastern North Dakota southward to the

JAN., 1895.] DISTRIBUTION. 27

Mexicau boundary along- the liio Grande; and also the southern half
of Alabama and Georgia, and the northern half of Florida. The genus
does not occur west of eastern Wyoming, east-central Colorado, and
the Rio Grande Yalley in New Mexico. (See also map 4.)

The genus Gratogeomys (map 2) inhabits the Gre^it Plains of the United
States from the Arkansas River in eastern Colorado southward, and
the eastern table-land region of Mexico to its extreme southern edge in
the States of Mexico and Puebla.

The genus Pappoyeomy.s (map .'i') is known only from the State of
Jalisco in Mexico.

The genus Platyf/eomys (map 3^) inhabits a rather narrow belt along
the southern border of the Mexican table-land in the States of Jalisco,
Colima, Michoacan, Mexico, and Hidalgo.

The genus Orthogeomy.s (map 3'') inhabits elevated parts of the States
of Oaxaca and Chiapas, in extreme southern Mexico and adjacent parts
of Guatemala.

The genus Heterogeomys (map 3^) inhabits the tropical plains of Vera
Cruz, below the edge of the table-land, and extends thence southerly to
Coban in Guatemala, probably following the low coastal plain of
Tabasco to the Rio Usumacinta and thence up tlie valleys of the San
Pedro and its tributaries to the interior of Guatemala.*

The genus Macrogeomys (map 3'') inhabits the highlands and moun-
tains of Costa Rica and is not known elsewhere.

The genus Zygogeomys (map 3 ') inhabits the Sierra Madre of the State
of Michoacan on the southern ijart of the table-land of Mexico.

NUMBER AND DISTRIBUTION OF THK SPECIES.

Omitting the genus Thomomys, the number of species recognized by
Baird in 1857 was 7, as follows : 0. hursarim, hrevicepSy pinctis [ = tuza],
clarlcil, casfmiops, hispidus, and mcxicanus. The number recognized by
Cones twenty years later, in 1877, was 5, as follows : G. bursarius, tuza,
castaiiops, hispidus, and mexicanus. Cones degraded U of Baird's spe-
cies to synonomy, uniting hreviceps with bursarius, and clarkii with
castanops. The same fate overtook G. heterodus of Peters, described
111 the interval between Baird and Cones ; it was made a synonym of
hispidus.

The number of species and subspecies recognized in the present
paper is 37, of which 21 are described as new. The remaining 16 are
accounted for as follows : Four out of the 5 admitted by Coues are
retained, namely, bursarius, tuza, castanops, and hispidus, but the tilth,
mexicanus, is rejected as preoccupied by an unidentifiable species (see

' While this paper is passing through the press, a specimen of Heterogeomi/s has
been received from Mr. Nelson, collected by him at Reyes, about 50 miles north of
the city of Oaxaca, in the State of the same name, and 33 miles south of the bound-
ary of Vera Cruz and Puebla.

28 NORTH AMERICAN FAUNA. [no. 8.

postca, p. 200). Baird's hrevleeps and Peters's heterodus are rein-
stated as valid species, and jioridanus of Audubon and Bacliman is
admitted as a subspecies of tuza. Tlie remaining 9 have been described
since the publication of Coues's Monograph — in fact, during* the past
five years — and no less than G of them are from Mexico and Guate-
malii. These species are: personatns oiTv\m; biiUeri* grandis, scalops,
and merriami of Thomas; lutescenSy fumosus, and gymnurus of Mer-
riam, and dierrici of Allen. Of the 21 new forms here described, 6
are from the southern United States (1 from Alabama and 5 from
Texas), 12 from southern Mexico, 2 from Costa Eica, and 1 from Guate-
mala. Of the total number here recognized (37), 10 are restricted to
the United States; 2 (probably 3t) are common to the Unitetl States
and northern JMexico; 17 are restricted to the southern half of Mexico;
2 are common to southeastern JVIexico and adjacent parts of Guate-
mala, and o are known froui Guatemala and Costa Eica only. Thus
no less than 24 species, representing, as will be shown later, 7 distinct
groups or genera, are absolutely contincd to southern Mexico and north-
ern Central America. The extraordinary and unexpected richness of
this part of tropical America iu members of the group, | and the even
more remarkable div^ersity of structure presented by the various types,
are of the utmost interest in view of the time and place of origin of
the family to which they belong.

UNITED S'J'ATES SPECIES.

The Pocket Gophers of the United States fall naturally into two prin-
cipal subdivisions, (1) those having the upi)er incisors deeply marked
by a median longitudinal furrow {unisulcatc series), and (2) those having
the upper incisors double grooved, a narrow sulcus on the inner margin
of the tooth and a larger and deeper one near the middle (bisulcate
series). The unisulcate series is represented by a single species, casta-
nops of Baird, which inhabits the western plains from middle Colorado
southward into Mexico. The members of the bisulcate series iuhabit-

* G. hulhri was descriljed ahiiost siuuiltaneonsly by Mr. Tlionias and myself, but
Mr. Thomas's descriptiou was issued tirst aud liis iiainc btilhrl Las i>riority over my
name iichoni.

I These are Geomijs arenarius, which is common on both sides of the Rio Grande
at El Paso, Texas, and Juarez, Mexico, aud Crato(jeomijs castunops, which inhabits
extensive areas in western Texas and Chihuahua. A third species, Geomys pemo-
natus, inhabits the lower Rio Grande region iu Texas and iu all probability occurs
on the Mexican side also (in the state of Tamaulipas).

iWheu it is remembered that only about half a dozen specimens, all told, have
been examined from Costa Rica and Guatemala, as compared with 200 front Mexico,
it must be evident that the possibilities of Central America have been by no nutans
exhausted. Furthermore, no specimens have been seen from Yucatan, though the
lamily is represented there by at least one species. (Biologia Ccntrali-Americana,
Mammalia, 1880, p. IGO.)

JAN.

,is:n.| UNITED STATES SPECIES. 29

iiig the United States are 12 in uiimber. These, with tlieir type locali-
tie;', are as follows:

Gcomi/s liiza (Ord) Augusta, Georgia.

tuzafloridanus Bach St. Augustine, Florida.

tuza mohilensis subsp. iiov Mobile Bay, Alabama.

bursarius (Shaw) Miuuesota?.

lutescens Merriam Birdwood Creek, western Nel)ra8ka.

hreviceps Baird Mer Rouge, Louisiana.

hreviceps sagittalis subsp. nov Galveston Bay, Texas.^

hreviceps atlwateri subsp. nov Roekport, Aransas County, Texas.

/exojsis sp. nov Mason, Texas.

rt) enarius sp. nov El Paso, Texas.

pemonatus True Padre Island, Texas.

personatus faUaX. subsp. nov Corpus Christi, Texas.

Geomijs bursarius is thecoiiiinoii Pocket G-opherof the northern Mis-
sissippi Valley, from eastern North Dakota and western Minnesota
south to southeastern Missouri. It is a dark liver-colored animal with
pure white forefeet, in sharp contrast to the color of the surrounding
])arts, and has the longest claws of any of the bisulcate species.

Ocomys lutescens is a pallid form of the bursarius type, inhabiting the
arid sand hills of western Nebraska and extreme eastern Wyoming,
and ranging thence southerly into northwestern Texas.

Geomys brerieeps inhabits the alluvial lands of Louisiana, Arkansas,
and eastern Texas, the typical form coming from Prairie Mer Rouge,
in Morehouse Parish. It extends thence northwesterly up the valley
of the Arkansas Kiver nearly to the Kansas border. It is a rather small
dark species. On the south, along the coast region of Texas, it splits
up into the two following subspecies :

Geomys breviceps sagittalis inhabits the gulf coast of Texas about
Galveston Bay. It is smaller than true breviceps.

Geomys breviceps attwateri inhabits the coastal plain and islands of
Texas, from Nueces Bay northward to Matagorda Bay, and ranges into
the interior nearly to San Antonio. . It is considerably larger than
typical breviceps.

Geomys texeiisis in its typical form inhabits central Texas. On the
north and northwest it probably passes into lutescens, while on the east
it may intergrade with breviceps. It is much smaller than bursarius or
hitescens and has a pure white belly. Its upper parts are reddish-
brown, paler than bnrsariuSj but darker and brighter than lutescens.

Geomys arenarius inhabits a very restricted area in the upper Rio
Grande Valley in extreme northern Chihuahua, western Texas, and
southern New Mexico. So far as known it is completely isolated, not
coming in contact with any other bisulcate species. It is of medium
size, has a relatively long tail, and the upper parts are drab.

Geomys personatus mhaibitaVadre Island and the adjacent coavSt of
Texas from Santa Rosa southward, extending inland as far as Carrizo,
on the Rio Grande; its range, together with that of its subspecies /rtZ/fw,
thus coincides with the northern arm of the arid tropical belt along the

30 NORTH AMERICAN FAUNA. [no. 8.

Gnlf coast. lu external appearance persoiiatus muck resembles G.
lutescem of the Great Plains, from which it may be distinguished at
once by its larger size, larger and more naked tail, and by important
cranial characters.

Geomys persofiatus faJlax inhabits a small area on the Gulf coast of
Texas, immediately south of Xueces Bay. It is smaller and darker than
tvne personatus.

Geomys tuza, a rather large cinnamon-brown species, inhabits the pine
barrens of eastern Georgia, where it is locally known as tlie 'Sala-
mander.' The same name is applied to the following subspecies:

Geomys tuza Jioridamis is a Florida form of tnza^ as its name indi-
cates, and does not differ materially in external appearance.

Geomys tuza mohllensis inhabits southern Alabama and northwestern
Florida and is a strongly marked form. It is very much darker than
tuza. ( For distribution of United States species see map 4).

DISTRIBUTION OF THE MEXK^AN SPEriES.

At my request Mr. Nelson has prepared the following note, embody-
ing his personal knowledge of the geographical and vertical distribu-
tion of the species obtained by him in Mexico, exclusive of the genus
Thorn omys :

'•One of the most remarkable and interesting features connected with
the Mexican Pocket Gophers is the small area within which most of the
laiown species occur. This area is a belt about 400 miles in length by
()0 in breadth, stretching from the Pacific coast to the Gulf of Mexico,
between the nineteenth and twentieth parallels of north latitude. It
contains tlie thirteen highest peaks of Mexico,* all of which attain an
altitude of lL*,Ono feet or upward. The most notable of these are Iztac-
cihuatl (17,000 feet), Popocatapetl (17,523 feet), and Orizaba (18,314
feet).t

*Tlie only peak in Mexico attaining an altitnde exceeding 12,000 feet, in addition
to tliose here cnnmerated, all of which lie in the Geomys belt, is Mount Zenipoal tepee,
in the State of Oaxaca. This peak is said to reach 12,000 feet, and is inhabited bj' a
new species of gopher here named Orthogeomys nelsoni.

IThe complete list with approximate altitudes, beginning at the westernmost, is as
follows : Yeei.

Sierra Nevada de Colinia 14, 000, State of .Jalisco.

Volcano de Colima 12,000, Do.

Pico de Tancitaro 12, 653, State of Michoacan

Pico de Patamban 12, 200, Do.

Volcano de Toluca 15, 000, State of Mexico.

Cerro de Ajusco 12, 000, Do.

Popocatapetl 17, 523, State of Puebla.

Iztaccihuatl 17, 000, Do.

Cerro de Telapon 13, 575, Do.

Cerro de Maliuche 13, 462, State of Tlaxcala.

Orizaba 18, 314, State of Puebla.

Sierra Negra , 15, 000, Do.

Cofre do Perote 14, 000, State of Vera Cruz.

JAN. 1885.

MEXICAN SPECIES. 31

"The maiu cliaiu of the Cordillera or Sierra Madre extends along this
line and forms here the southern limit of the plateau or table-land region.
The mountains throughout this district are of volcanic origin. They
inclose numerous high valleys, such as that of Toluca (8,000 feet) and
the valley of Mexico (7,400 feet). The main body of the range takes
the form of high rouuded ridges between 7,000 and 9,000 feet in altitude.
On the north the ridges slope down to the adjacent tablelands; on the
south a longer slope carries their bases into the low hot valleys of the
streams that lead out to the sea. The average elevation of the belt
under discussion is far greater than that of any other equal area in
Mexico or Central America; this belt also contains the only peaks of
the region that are permanently capped with snow,

"The characteristic trees of all these mountains are pines, firs, and
alders. In descending toward the hot coast country, below 7,000 feet,
oaks come in, and as the descent is continued they in turn give way
before the subtropical and tropical species. Although most of the area
within the limits given is high and cool, yet at each end a sharp
descent leads to the low, hot coast country.

"Gophers occur throughout this area, from the hot coast districts up
to the scattered vegetation about timber line. Geomys fumosus, the
extreme westernmost species, burrows in the damp clayey soil among
the cocoanut palms about the city of Colima, at an altitude of from
1,000 to 2,500 feet. Geomys M.spidus, the easternmost representative of
the groui), inhabits the coffee and sugar-cane fields of V^era Cruz. In
the intervening district the other species range from 4,000 feet up to
timber line. Although several reach as high as 12,500 or even 13,000
feet, the great majority of individuals of all species occur below 9,000
feet, and a vertical section of the country from 4,000 to 9,000 feet would
include all of the species and nearly all of the individuals of the
interior forms. By far the greatest development of the group is
reached between the altitudes of 0,000 and 8,500 feet. This area is
along the lower border of the pine and oak forest and reaches out along
the adjacent treeless plains for a short distance. Considered faunally,
this area is Upper Sonoran and Transition. The northern base of this
part of the Cordillera forms the southern limit of many species of
birds and mammals belonging to the great interior deserts of the
United States and the x^lateau of Mexico, while their southern base
and adjacent slopes form the northern limit of various tropical species.

"It was observed also that whenever the route led to the north or
south of this belt the pocket gophers became rapidly less numerous,
and ceased entirely except in a few i)laces,

"By far the greater number of species now known from Mexico are
absolutely restricted to limited areas within this district, while others
push out only a little beyond.

" The animals, as a group, are generally found in rather loose soil and
avoid stony areas. In some cases, as with G. funiostis, the soil may be

32 NORTH AMERICAN FAUNA. [no. 8.

a tough clay, but this is exceptional. Wherever fouud in cultivated
districts they invade fields, and frequently commit serious damage to
crops of both grains and tubers. It is a common practice for the land-
owners to pay a fixed bounty to their field hands for them. The owner
of a hacienda near Atlisco, Puebla, told me he had thus iiaid for sev-
enty dozen on his hacienda in a single year, at the rate of (3 cents a
head."

The most interesting and unexpected result of Mr. Nelson's explora-
tions is the knowledge that the family Geomyidw attains its highest
development in a belt about 400 miles in length by GO in breadth
which crosses Mexico from west to east along the southern edge of
the tableland. Within this belt Mr. Nelson collected 175 specimens,
not counting this genus Thomomys. These specimens belong to six
different genera and represent 15 species, no less than 12 of which
were previously unknown.*

WEIGHT OF CHARACTERS.

Nothing is more difficult, in entering upon the study of a new group,
than to determine the relative weight of characters. Structures of known
stability in one group may be highly variable in another, so that char-
acters that are of generic value in the one may be of only specific value
in the other. In framing genera and higher groups therefore it is
desirable to select deep-seated structures and those that are not easily
affected by external influences. In the case of the skull, it is conven-
ient to divide the characters into two categories, fundamental or pri-
mary, and superficial or secondary. Fundamental characters are based
on structures and relations that enter into the ground plan of the skull,
and are of high morphologic weight; super fjciaJ characters are the result
of special adaptations and particular muscular strains, and are of little
value except as aftbrdiug recognition marks for species, and in some
instances for genera also. The fundamental structures are mostly
hidden, comprising the floor of the brain case, the craniofacial axis, and
the turbinated bones. They are seen to best advantage in vertical longi-
tudinal sections and in skulls from whic.li the vault of the cranium has
been removed. On the outside of the skull the palatopterygoid plates,
and perhaps the frontals also, may be regarded as belonging to the
same category. The superficial structures are those that appear on the
outer side of the cranium and are most easily modified by muscular
strain, or are the secondary result of dental peculiarities. They com-
prise the zygomatic arches, muzzle, nasals, occiput, and such parts of

* Since the above note was written — in fact just as tbis paper is going to press —
Mr. Nelson bas sent me 15 si>ecimens of large gopbers from tbe f-'tato of Oaxaca, in
extreme southern Mexico. Ton of these, from Cerro San Felipe, are the species
recently described by Mr. Oldiield Thomas as Geomys scalops; the remaining .5 are a
new &\iL'.c\es, Orlhofjeomys nelsoiii. They were collected at three localities: Mount
Zempoaltepec, Totontepec, and C'omaltepec. All of tbe specimens from the State of
Oaxaca belong to a genus (here named Orthogeomyti) quite distinct from nny of the
genera inhabiting Mr. Nelson's Geomys belt.

JAN., 1895.]

MORPHOLOGY OF THE SKULL.

33

tlie outside of the vaultof the craiiiuni as are niateiially altered in form
and extent (as tlie squamosals) without sensibly changing their relations
on the inner side of the brain case.

LIST OF SPECIMENS EXAMINED.

Geomys tuza (Ord) 32

4*/2'a//o/-irf«H«s( And. and Bach.) 25

titza mohilennis subsp. nov ... 23

bumarhts (Sliaw) 116

lute-scens Merriam 136

hrexnceps Baird 195

hreviceps Hagittalis siibsp. nov. 26

brevieeps (iftwaferisnhs'p. nov. 53

texensis sp. nov 31

arenarius sii.nov 43

personatus Trne 33

])ersonatHS faUaxm\\)&\^. nov.. 22

Pappogeomys bulleri (Thomas) 6

albinasiis sp. nov 1

Cratoyeomys merriami (Thomas) 31

perotensis sp. nov 13

estor sp. nov 10

pcreyrin us sp. nov 1

oreocetes sp. nov 1

Cratoyeomys castanops (Baird)

43

casfanops yoldmani snbsp.

nov

5

fiilvcscens sp. nov

11

Platyycomyfi yymiiiirus Merriam

10

tylorkhi MS sp. nov

9

phin'ireps sp. nov

3

f 11)11 osus Merriam

11

Orthoyeomyfi scdJops (Thomas)

13

iielsoui sp. nov

0

latlfrons sp. nov

1

Heieruyeomys hispidiis (Lc Coute)

9

torridus sp. nov

27

Alacroyeomys heterodus (Peters)

1

dolicliocephahis sp. nov..

2

eostaricensis sp. nov

1

chcrriei (Allen)

1

Zyyoyeomys trichopas sp. nov

12

CHAPTEK II.
MOKPHOLOGY OF THE SKULL.

1. THE CRANIUM AS A WHOLE.

While diversity prevails in the form of the cranium as a whole and in
a nniltitude of minor details, all the members of the family GeomyuJm
agree in the following important characters: The top of the skull is
llattened, the nasals, frontals, and parietals usually forming nearly a
straight line (though the line is decidedly convex in Gratogeomys cas-
tanops ?ind fulvescens). The tympanic or audital bulhe are rather large,
and the external meatus is a long tube directed forward as well as out
ward, and opening externally immediately behind the posterior angle
of the zygoma. There is a well-developed mastoid bulla which is wholly
on the occipital plane, never reaching the top of the skull. The squa-
mosals are largely developed, always overlapping the lower part of the
parietals and hinder part of the frontals, and sending out posteriorly
a lateral arm which enters into the occipital plane and overreaches the
mastoid process of the mastoid bulla. They articulate broadly with
the alisphenoid, but leave a long slit-like vacuity between the postero-
inferior margin and the audital bulla. The hasisphenoid and presplienoid
are higher than broad. The former develops air cells in its body; the
latter is a thin vertical plate always j)erforate anteriorly opposite the
7433— :^o, 8^ 3

34 NORTH AMERICAN FAUNA. [no. 8.

splieiioidal fissure, so tliat in viewiug the skull from tlie side oue sees
completely tlirougli it below tbe orbitospheuoids. The alisplienoids
are larger and reach, or nearLy reach, the upper surface of the cranium;
they are inseparably ankylosed to the basisphenoid before birth. The
orhitosphenoids are small and horizontal and are not united to the ali-
sphenoids except in Zygogeomys and Thomomy,s. The turbinated hones,
while presenting important differences in the several genera, agree in
the following particulars: Anteriorly there is a Hmg\e7naxillo-turbinal,
always attached to the premaxilla; above and i)arallel to it is a large
fiaso-tnrhinal, always attached to the nasal; posteriorly, and attached
to the cribriform plate and os planum are the endoturhinals (of Harri-
son Allen), always four in number and always decreasing in size from
above downward; the uppermost is expanded anteriorly.

The hony palate is long and narrow, broader posteriorly than anteriorly,
and composed chiefly of the maxilld, the body of the |>a/a/m£; being rel-
atively small and situated far back. There is a deep pit on each side
of the palate between the hindermost molars. Posterior to this pit the
palatines usually bifurcate and unite, with the pterygoids to form a
Ungulate or strap-shaped palatopterygoid plate on each side of the poste-
rior nares. On the outside of the skull the palatines are restricted to
the posterior eiul of the bony palate, but on the inside they reach
forward along the crano facial axis all the way to the nasal chamber —
a wholly unnecessary condition so far as the i)resent structure and needs
of the animal are concerned, but a highly interesting and significant
relic of the primitive relations.of these bones. The case is an excellent
illustration of the persistence-of. useless parts.

T\\Q premaxilla is large and heavy, subquadrate in section, and artic-
ulates rather broadly with the frontal. It completely incloses the small
incisive foi-amina except in Zygogeomys.

The jugal is a highly variable bone (as will be seen hereafter), but
it is always restricted to the horizontal part of the zygoma, never
creeping upward anteriorly toward the lachrymal, or inward i)Osteriorly
toward the glenoid fossa.

The vomer bifurcates and sends backward two long vertical wings,
which articulate with the sides of the presphenoid, never with its
inferior surface.

The zygomatic arch varies exceedingly in size and form in the differ-
ent subgenera, but its horizontal part in transverse section is always
distinctly triangular anteriorly, while posteriorly it is fiat or rounded.
Posteriorly it presents two faces, inner and outer; anteriorly a third is
added — a supero-external face. The latter rarely reaches further back-
ward than the middle of the arch and is usually set oft" from the outer
lace by a well-defined ridge, which passes obliquely backward and
upward from the antero-exteriial angle to the tip of the squamosal
arm. This ridge marks the upper limit of attachment of the zygomatic
part of the masseter muscle.

JAN., 1895] MORPHOLOGY OF THE SKULL. 35

There is no true 2)ostorbital process of the frontal except iu Macrogeo-
mys, but the apex of the alisphenoid and adjoiuiug anterior "border of
the squamosal commonly unite to form a decided postorbital ridge, which
slopes obliquely downward and backward from the i)oint where the
frontal, alisphenoid, and squamosal meet, just behind the orbit. This
ridge is made up of the edges of the alisphenoid and squamosal, and
serves to sharply separate the orbit from the adjoining outer side of the
brain case. In Macrogeomys there is a strongly developed circumscribed
postorbital i)rocess, which, with the help of a corresponding eminence
on the middle of the horizontal part of the zygoma, serves to sharply
distinguish the orbital from the temporal fossa. In its component
elements it is peculiar. Its base consists of the frontal, which bone is
notched immediately in front of it, thus emphasizing the appai'ent size
of the process. The summit of the process is made up of the apex of
the alisphenoid, which hero reaches the plane of the upper part of the
skull and is slightly overlapped posteriorly by the autero-external angle
of the squamosal.

T\\Q paroccipital processes stand out sideways above the condyles and
are more or less expanded and flattened — never cylindrical or conical
(figs. 4 and oo pp. and x)l. 15, tigs. 0 and 7).

The^oor of the brain case, as exposed by sawing oft' the vault of the
cranium, affords characters of the utmost value in subdividing the
group into genera (figs. 9, 5G, and 08^, and i)l. 17). As will be seen
on consulting fig. 9, the tympano-periotic capsules, with the inclosed
basioccipital and posterior part of the basisphenoid, form about half of
the floor of the brain case. The alisphenoids (fig. 9, as) are next in
importance, the horizontal part forming abridge across the floor of the
skull above the pterygoid foss;e and immediately iu front of the tym-
panic bulhe, while the ascending wings push forward on each side,
reaching or nearly reaching the orbitosphenoids (os), and forming the
posterior and outer boundaries (»f the large sphenoid fossa. Anteriorly
the orbitosphenoids fill or nearly fill the front part of the floor of the
brain case, on the plane of the orbital constriction. In front of this
constriction, and behind the cribriform plate, the orbital or descending
iflates of the frontal commonly meet in the median line, forming the
floor of the olfactory fossa. In young skulls, as in fig. 9, and iu adults
of the genera Pappogeomys (fig. 56), Orthogeomys, and Thomomys {&g. 08^),
the frontals do not meet below, but the orbitosphenoids reach forward
and articulate directly with the cribriform plate.

A conspicuous and highly important pair of fossa; occupy the ante-
rior part of the floor of the brain case on each side of the median line,
where they are completely surrounded by the several sphenoid bones.
They may be termed the sphenoid fossw. They are directly continuous
and inseparably connected posteriorly with the pterygoid fossce proper,
which latter are widely oi)en in front and are roofed over bv the trans-

36 NORTH AMERICAN FAUNA [no. 8.

verse part of the alisplienoid only. The resulting elongated fossa as a
whole may be named the spheno-pterygoid fossa (Hg. ^yptf). The shape
aud extent of the sphenoid fossa varies materially in the different
genera, as shown in pi. 17: in Geomys (tig. 3) and Heterogeomys (tig. 1)
it is much elongated, reaching anteriorly to the descending plate of the
frontal. In Cratogcomys (fig. 0, pi. 17, and fig. 5), and also in Fappo-
geomys (fig. 56) and Orthogeoinys, it is cut oft" anteriorly by the orbito-
sphenoids. In Zygogeomys (pi. 17, fig. 2) it is still further shortened by
the posterior enlargement of the orbitosphenoids, which are broadly
ankylosed with the alisphenoids. ^

The anterior end of the alisplienoid canal (fig. 9, ac) always opens into
the outer side of the posterior x^art of the sphenoid Ibssa, and its posi-
tion is essentially the same throughout the family (see pi. 17, and text
figs. 9 ae, 52 aud 54 ale, 56, and 68).

The pterygoid fossw are large and widely open (fig. VI, ptf). Poste-
riorly they are bridged by the narrow horizontal arm of the alisphenoid
(tig. 9, as); anteriorly they are not closed or roofed over, but are broadly
continuous with the large and deep sphenoid fossae (fig. 9, pif), which
open into the orbit by means of the broadly expanded lower part of
the sphenoidal fissure. Their floor consists posteriorly of palatine and
anteriorly of maxillary. On the inner side they are bounded by the
pterygoid, the vertical plate of the palatine, the basisi)henoid, and the
presphenoid. On the outer side they are bounded inferiorly by the
external pterygoid plate of the palatine (fig. 12, epl), and superiorly
by the descending wing of the alisphenoid. The outer wall of the j)os-
terior part of the pterygoid fossa thus proves to be double, and the
inner bone — ilie external pterygoid plate — belongs to the palatine and is
overlapped by the descending wing of the alisphenoid, as shown in
figs. 4 and 12.

Thetsphenoidal fissure is a large and nearly vertical pyriform vacuity
at the bottom of the orbit, separating the anterior border of the ali-
sphenoid from the descending or orbital plate of the frontal (fig. 55").
It separates also, to a varying degree, tlie alisphenoid from the orbi-
tosphenoid (fig. 9, sf). Superiorly (above the horizontal jilane of the
orbitosphenoids) it is a narrow slit sloping oblicpiely upward and for-
ward between the brain case proper and the olfactory fossa, and ending
at the base of the thickened interorbital constriction of the frontal
(which continues the line of separation between the olfactory fossa and
cerebral chamber). This slit is permanently open except in Zygogeomys
(in which it is closed by the orbltosphenoid), looking completely through
the skull from side to side. Inferiorly (below the horizontal plane of
the orbitos])lienoids) the fissure is suddenly dilated, forming a broad
and widely open door between tlie deep lateral fossa of the floor of the
brain case and the bottom of the orbit. The corresponding basal parts
of t^e ^s,mv^ Qi] t/)J<? two sides of the skull are incompletely sep^vr^ted

JAN., 1895.]

THE SKUl

37

by a i^erforate septum consisting- of the vertical plate of the presphe-
noid, and in some cases of an ascending wing of the palatine also. The
sphenoidal Assure is bounded by tliree bones : posteriorly by the ali-

Cv I 01^ ^ 5)(\ sc^i

Fig. 4.— Side view of skull of Cratoijeomys merriami from the outside. Zygomatic arch sawed off
to show bottom of orbit. Animal not quite adult. Specimen from Amecameca, Valley of Mexico.
(This figure should be compared with the corresponding view of Geomys bursaiius, fig. 55.)

1 Infraorbital foramen.

2 Posterior (orbital) opening of infraorl)ital canal.

3 Foramen rotundum.

4 Foramen ovale.

5 Meatus auditorius externus.

6 Fenestrum in anterior part of presplienoid (the line pointing to it cros.sos the upper part of

the sphenoidal fi.ssure).
apl Ascending wing of vertical plate of palatine.
• as Alisphenoid (the upper line re.st.s on the ascending wing; tlie lower on the descending wing),
c Condyle of exoccipital.
epl External pterygoid plate of palatine bone.
/(■ Frontal.

/( Ilamular process of pterygoid bone.
I Lachrymal.

in Mastoid process of mastoul bulla.
mb Mastoid bulla.
ms Mastoid process of squamosal.
mx Maxilla.

inx2 Zygomatic root of maxilla (.sawed off fo show orbit),
n Nasal.

off Orbital or dcsceuding plate of frontal.
OS Orbitosphenoid.
im Parietal.
pinz Premaxilla.

pp Paroccipital process of exoccipital.
ps Presplienoid.
pt Pterygoid.
sn Supraoccipital.
fq Squamosal.

sqz Squamosal root of zygoma (sawed off).
tb Tympanic or audital bulla.

sphenoid; anteriorly by the frontal and maxilla; and inferiorly by the
maxilla. The longitudinal vertical septum which forms the floor of the
large inferior part of the sphenoidal fissure is likewise made up of three

38

NORTH AMERICAN FAUNA.

[NO. 8.

bones, the orbitosplienoid, prosphenoid, and palatine — tboiigh tbe lat-
ter is usually so reduced that it appears in tbe anteroinferior corner
only, and in some forms can not be seen from tbe outside at all. But
in tbe elongated skulls of Oeomys bursarius and tuza tbe lower part of
tbe fissure is broadened antero-posteriorly, and tbe ascending wing of

Fig. 55. — Side view of skull of Getmyn bursariua from outsitle, zygomatic arch sawed off to sliow
bottom of orbit. Animal fully adult d- From Knoxville, Iowa. (Tliis figure is duplicated for easy
comparison witli tbe corresponding view of Cratogeomys merrinmi, fig. 4).

1. Infraorbital foramen.

2. Posterior (orbital) opening of infraorbital canal.

3. Vacuity in front of pi-esphcnoid and ascending wing of jialatine.

4. Vacuity in prespbenoid, beliind ascending wing of palatine

5. Optic foramen (in orbitosijbeuoid bone).

6. Foramen rotundum and foramen ovale (wbicli liave liere coalesced).

7. External auditory meatus.

8. Sphenoidal fissure (upper part).

a2}l. Ascending wing of vertical plate of palatine.
as. Alisphenoid.

c Condyle ot exoccipital.
epl. External pterygoid jdate of palatine bone,
/»'. Frontal.
h. Hamular jarocess of pterygoid bone.
I. Lachrymal,
m. Mastoid process of mastoid bulla.
mb. Mastoid bulla.
ms. Mastoid process of squamosal.
inx. Maxilla.

n. Nasal.
pa. Parietal. '

pmx. Preniaxilla.
p}). Paroccipital process of exoccipital.
j}s. Presphenoid.
pt. Pterygoid.
so. Supraoccipital.
sq. Squamosal.
th. Tympanic or auditnl bulla.

tbe palatine is enlarged and extended, reacbing upward alongside tbe
prespbenoid (in front of tbe usual fenestruin) to articulate broadly witb
tbe frontal and orbitospbenoid, on or near tbe plane of tbe top of tbe
prespbenoid (fig. .55). In front of tbe palatine (and also in front of tbe
prespbenoid, wbicb is bere clasped between tbe ascending wings of tbe

JAN., 1895.1 THE SKULL. 89

palatine on the two sides of the skull) is a second feiiestruiii (fig. 55'*)
anterior to the usual one (fig. o")^, which is in the presphenoid), and
likewise looking completely through the skull. This latter opening is
bounded iu front by the maxilla and behind by the i»alatine. It is sit-
uated midway between the sphenoid fenestrum and the orbital end of
the infraorbital canal.

The infraorhital canal is small and does not pierce the root of the
zygoma, but is deeply buried in the maxillary bone, passing backward
and inward from the infraorbital foramen (fig. 4') (on the lower part of
the side of the muzzle Just beliiud the premaxillary suture) to the deep-
est part of the orbit (flg. 4-), its course being wholly internal to the zygo-
matic root of the nuixillary. It curves around the inner side of the
base of the socket of the long upper incisor, and is separated from the
nasal chamber by only a thin lamella of bone rising from the maxillary
floor of the nasal passa^;e and articulating above with the inferior boi'-
der of that j^art of the os i)lannm which supports the endoturbinals.

The foramen rotundum (fig.-4'') is always situated above the foramen
ovale (fig. 4^), and both open into the large longitudinal alisphenoid
canal. In rare instances they coalesce (fig. 55'^).

The narial passage is a narrow vertical ellipse, about twice as high as
broad (fig. 7, nj)).

While most species of the genera under consideration develop a
prominent sagittal crest in adult life, some do not, the temporal imj)res-
sions remaining permanently distant, defining a well-marked sagittal
area. The members of the latter category may be divided into two
sets, (1) those in which the temporal impressions are actual ridges ris-
ing above the level of the surrounding bone on both sides, as in Hetero-
geomijs hispidus (pi. 4), Gcomys tuza (pi. 7, fig. 1), and G. arenarius
(pi. 9, fig. 1) ; and (2) those in which the space between the temi>oral
impressions (the sagittal area) is thickened and as high as the impres-
sions, which thus appear as ridges only when looked at from the outer
side, as in Geomys breviceps (pi. 9, fig. (») and Cratogeomys orcocetes and
peregrinns (pi. 8, figs. 2 and 3).

The lamhdoid crest is broadly and gently convex posteriorly through-
out the group (pis. 1, 2, 5-9, etc.), except in Platygeomys, in which genus
(pi. 3 and pi. 11, fig. 4) it is strongly sinuous — forming a deep and broad
reentrant angle on the median "line, beyond which, on each side, it is
first strongly convex backward and then slightly convex forward — the
extreme mastoid ends curving backward as well as outward. The
bones that take part iu the formation of the lambdoid crest are the
supraoccipital, squamosals, parietals, and interiiarietal.

There is no ossified tentorium in the Geomyidiv.

40

NORTH AMERICAN FAUNA.

[no. 8.

2. TllK INDIVIIHIAL BONES.

Ill the Geomyidw there are normally thirty-three distinct bones in the
skull, not counting" the separate parts of the tympano-periotic capsule,
the turbinated bones of the nasal chamber (which are reckoned with
the bones to which they are attached) or the paired bones that coalesce
before birth. The latter are the preiuaxillje, maxillae, palatines, and
frontals.

The thirty-three bones that go to make up the skull (exclusive of the
paired bones that are fused in the embryo) are:

Basioccipital 1

Exoccipital 2

Supraoccipitai 1

Interparietal ' : 1

Basisplieiioid . . . .* 1

Alisplienoid 2

Squamosal 2

Parietal 2

Prespbeuoid 1

Orbitosphenoid 2

Frontal 1

Ethmoid 1

Vomer : 1

Pterygoid ^ . 2

Palatine 1

Maxilla 1

Premaxilla 1

Lachrymal 2

Jugal 2

Nasal 2

Periotic 2

Mandible 2

33

The hasioccipital in commonly truncate-wedge-shaped, with the pos-
terior edge {basion) rather deeply notched. Its posterior corners enter

Fig. 5. — Basiocciiiital of Gratogeomys merriami, showing diffeiTnce iu form of upper and lower sur-
faces (ankylosert ex()ccii)itaI.s's]iown also): a, inferior surface; h, superior surface; pp, paroccipital
process.

very slightly into the formation of the occipital condyles. The inferior
surface of the body of the basioccipital. is normally broader posteriorly
than anteriorly and the decrease in bread.th from behind, forward is
gradual (pi. 12, fig. 2, a\', but in one species, Cratogcomys casta7iops,tlie
body of the bone is rectangular, its sides being parallel (pi, 12, fig.l, a).
In another, Orihogeomys sealojjs, they may be nearly parallel or even
slightly divergent anteriorly (pi, 19, fig. 2). The basioccipital varies
in breadth according to the development of the audital bullae, by which
its sides are always more or less excavated. Its outer borders are
usually grooved to receive a i)rojectiou from the bulla. The superior
surface (on floor of brain case) is always narrower than tlie inferior
surface. The difference is very marked in some species (see fig. 5, a and
6). The basioccipital early ankyloses with the exoccipitals,* but usually

* The exoccipitals coiissify with the basioccipital very early in Zygogeomys and
Geo/Hj/s proper ; somewhat later in Crdfof/eomi/s, Plati/f/eumys, and Ileteroyeomtjs.

•IAN., 1895.] THE SKULL. 41

remains distinct from tlie basisphenoid, with which it nnites by syn-
chondrosis.

The ejfoccipitah form the whole of the condyles except the extreme
lower ends, into which the outer corners of the basioccipital enter.
They early ankylose with the basioccipital, forming a single bone long
before the animal becomes adult. No part of the exoccipital ever pro-
jects downward below the plane of the condyles. The paroccipital
processes stand out sideways and impinge upon the base of the mas-
toid bulla immediately behind the audital bulla; they are commonly
more or less flattened and expanded, and their distal ends often pro-
ject backward (tig. 12,|>p). In Platygeomys they attain their maximum
development and form the lateral parieties of a deep basin-shaped
depression, the upper boundary of which is formed by the backward
projecting lambdoid crest (pi. 15, fig. 7). The exoccipitals are in contact
anteriorly with the mastoid bulla' and i^eriotic capsules, which they
partly overlap. Viewed from behind, they form the inner boundary
of the exposed i^art of the mastoid bulhie. Vertically they reach the
upper edge of the foramen magnum, and their upper border forms
nearly a straight line across the plane of the occiput.

The supraoccipital forms a small part of the roof of the brain case
and the greater part of the occipital plane, comprising all of the occi-
pital element above the foramen magnum. On the top of the skull it
reaches much farther forward in Plati/fjeomi/s than in the oth'er genera,
(fig. 53, .yo), but is usually nearly concealed in adult life by being over-
lapped by the parietal and squamosal. On the occipital plane its
inferior border forms the superior boundary of the foramen magnum ;
its outer sides curve around the basal part of the exposed mastoid
bullae, though rarely reaching laterally as far as the free ends of the
mastoids. Anteriorly the supraoccipital articulates with the squamo-
sals and parietals, and with the interparietal also in those cases in
which the latter bone has an independent existence. [As a rule the
interparietal is not separate from the supraoccipital.]

The interparietal, which has proved of considerable importance in
furnishing specific characters in the Heteromyidce, is small and of little
consequence in most species of Geomyidw, except in the single genus
Thomomys. Even in very early life it forms an inseparable jjart of the
supraocciptial in the castanops series of Cratogeomyf!, in Platygeomys
gymmirus, in the hursarius series of Geomys proper, and in Pappogeomys,
Heterogeomys, and Zygogeomy-s. It is distinct all around in early life in
most s]>ecies of Thomomys, in the merriami series of Cratogeomys, iu the
tuza series of Geomys proper, in Geomys texensis and hreviceps, in Platy-
geomys tylorhinus and planlceps, but not in P. gymnurus. Fromits varia-
bility in closely related species it is evidently of little importance for
purposes of classification, though its value in Tliomomys is much greater
than in any of the other genera; and it is of some value iu the restricted
genus Geomys also. In the young it is commonly sub(piadrate or
broadly oval and of relatively large size, but with advancing age it

42

NORTH AMERICAN FAUNA.

[NO. 8.

becomes smaller and uanowly triaiii;ulai' or wedge-shaped, its outer
borders being resorbed from pressure of the parietals, which are con-
stantly crowding toward the mediau line. Thus in PlatygeomyH ti/Io-
rhinus several sivulls from the same locality (Tula, Hidalgo, Mexico)
present the following variations in the interparietal:

O Q

A

A

Tig. G.— Forms of interparietal. «, h, c, d. Platiigeom'is tylorhinus showing changes with age.
e. Gcomys tuza iS art- Aiigiist.i, Ga.
/and g G. mohilenais: ci f yg- .irt-: <J / ad. Milton, Fla. All natural size.

A very young male (fig, 6, «, Xo, 51882) has it roughly subquadrate
and broader than long; an immature but older female (fig. C&, No.
51884) has it of the same shape, but narrower and longer than broad;
a still older specimen (fig. Xo. fi, c) has it broadly triangular; while an
adult (fig. G, r7, No. 51883, $ ) has it reduced to a small wedge-shaped
piece squeezed in between the hinder edges of the parietals.

In the young of Zygogeomys irichopus the interparietal is even larger
than in Platygcomys tylorhinus, and is about twice as broad as long
(measuring 8 mm. in breadth in Xo. 50104: juv. fig. 15, a). In shape it is
broadly convex anteriorily and slightly (flatly) convex posteriorly.
The progressive development of the powerful temporal muscles with
consequent enlargement of the parietals posteriorly encroach upon its
size and change its shape, pressing it into an equilateral triangle (as in
No. 4718G S im., fig. 15, h). Its size now decreases rapidly, and as the
temporal impressions meet in a well-developed sagittal crest in the
adult skull it nearly or quite disappears from the upper surface of the
cranium (as in No. 50100 S ad., fig. 15, c).

The interparietal ?s more stable in form in several of the species of
the restricted genus Geomy.s than in any of the other genera under con-
sideration. This is due chiefly to the circumstance that in this genus
several species have permanently distant temporal impressions — for
nothing is so destructive to an interi)arietal as the development of a
sagittal crest. In the species possessing a crest {hnrsarhis, Infescens,
per son atUH, fall a.r, and mohilensis) the interparietal is normally reduced

IAN 1895] THE SKULL. 43

in adult life to an inconspicuous subtri angular wedge. In the species
having a permanent sagittal area it remains of considerable size and
its form is reasonably constant. In G. (ircnarius it is normally sub-
quadrate, though the anterior border may become convex from rounding
oft' of the corners, and it is always truncate behind and persists in old
age (pi. 9, fig. I). In G. texensis it is normally elliptical or oval (broader
than long) and convex posteriorly as well as anteriorly, projecting
nearly as far behind as in front of the lambdoid suture (pi. 9, fig. 2).
In G. hreviceps it is usually reduced to a highly irregular ' wormian'
bone, raucli cut up by contortions of the sutures (])1. 9, fig. 0). In G. tuza
it is very large, occupying nearly half of the broad sagittal area, and is
convex in front, truncate behind (fig. i^e). Inthe closely related G.
mohilensis it is deeply notched behind and is encroached upon and
finally nearly obliterated by the union of the temporal ridges (fig. 6,/
and a).

The hasisphenoid is invariably aukylosed with the alisphenoids and
pterygoids, even in early life, and sooner or later usually coossifies
with the presphenoid; it commonly, though not always, remains dis-
tinct from the basioccipital. Its vertical height is generally greater
than its breadth, and air cells commonly develop in its substance
(fig. 7, bs). Its chief peculiarity is the slight development of the pitu-
itary fossa, which ordinarily is so shallow as to escape notice. But in
Heterogeomys it is a real depression, and in H. hispiiJus it is normally a
pit and completely perforates the bone. In the related species, H. tor-
ridus, it is much less conspicuous and never perforates (so far as the
series of 26 skulls goes).

Tlie basisphenoid articulates with the basioccipital, presphenoid,
alisphenoids (by ankylosis), pterygoids (by ankylosis), and vertical
plates of the palatines (by contact antero-inferiorly — see fig. 7).

The alisjyJienoid is a very important bone, serving to bind firmly
together the middle segment of the vault of the cranium with the pos-
terior part of the upper jaw, and to anchor both securely to the basi-
cranial axis. It maybe described as consisting of three parts, (1) a
horizontal or transverse part, (2) an ascending icing, and (3) a descending
icing.

(1) The transverse or horizontal part is little more than a narrow bar,
inseparably connected with the middle of the outer side of the bas-
isphenoid (figs. 9, as and 54, alh) ; it forms the floor of the brain case imme-
diately in front of the periotic, and the roof of the posterior part of the
pterygoid fossa, the anterior part being uncovered. In passing outward
it bifurcates to inclose the large longitudinal alispheuoid canal, above
which it becomes continuous with the ascending wing, and below with
the descending wing. Posteriorly, the base of the horizontal part of
the alisphenoid is excavated, and usually presents a cup-shaped enlarge-
ment to receive the apex of the audital bulla. It also descends alongside
the basioccipital to unite with the pterygoid posteriorly.

44

NORTH AMERICAN FAUNA.

[NO. 8.

(2) The ascending icing of the alisphenoid differs widely iu form as
viewed from the inside or outside of the braiu case. On the outer side
of the skull (tig. 4, as) it is a long rectangular blade ascending
obliquely in front of the squamosal, with the anterior border of which it
articulates. It also overlaps the posterior i)art of the orbital face of
the frontal, rising nearly to the upper surface of the skull, which it some-
times reaches. The upper part is always roughened, and, with the over-
la]>ping edge of the squamosal, forms an oblique postorbital ridge or
prominence. Sometimes the apex pushes up to the top of the skull,
where it is thickened and forms the major part of a distinct postorbital
process, resting on the frontal, and overlapped posteriorly by the autero-
exterual corner of the squamosal. This process attains its highest
development in Macrogeomijs (see pi. 11, fig. 2, and text fig. 17^). Pos-
teriorly the ascending wing is extensively overlapped by the squamosal,

"F ''^ T3 'ti ^ u a h

ffia, 7.— Longitudinal vertical mediau section of skull of Cratogeoini/i mernami, showinc
of brain caso and nasal chamber. Vomer aud mesethmoid in place.

interior

1 Anterior palatine foramen.

2 Incisive foramen.

3 Meatus auditorius iuternns.

4 Floccular fossa.

5 Upiier part of sphenoidal fis.sure.
as Alisphenoid.

ho Basioccipital.

bs Basiaplienoid.

c Condyle of exoccipital.

fr Frontal.

h. Hamular process of i>terygoid.

ip Interparietal.

tne Mesethmoid plate.

mt Maxillo-turbinal.

mx MaxiUa.

n Nasal.

nt Naso-turbinal.

op Lower border of os planum.

pa Parietal.

pet Petrous part of periotic capsule.
pi Palatine.
pmx Premaxilla.
ps Presphenoid.
Pterygoid.
Supraoccipital.
Squamosal.

Tympanic bulla (antero-superior part,
which alone appears within the brain
case) .
Vomer.

Vomerine sheath of maxilla
First endoturbinal (below and somewhat
behind it the anterior ends of the sec-
ond, third, and fourth endoturbinals
may be seen).

pt

as appears when examined from the inner side of the brain case (fig. 7,
as). Therefore, while the outer face is an obi iquely- vertical i>late, with
essentially parallel sides, the inner face is elongated horizon tallj^, with
an irregularly convex upper border — the difference being due to the fact
that the outer side overlaps the frontal anteriorly and is overlapped b/

JAN., 1895.] THE SKULL. 45

the squamosal posteriorly. The alispheuoid may be separated from
the orbitospheuoid as in Heterogeomys and Geomys (pi. 17, tigs. 1 and 3),
or the two bones may be in contact anteriorly as in Cratogeomys (pi. 17,
tig-. 5, and text tig. 9), or they may be firmly and broadly ankylosed
together as in Zygogeomys (pi. 17, tig. 2).

(3) Tlie flesccnding wing of the ciMsphenoid, on the outer side of the
skull, is a tlatteued plate continuous in breadth, plane, and direction
with the ascending wing, and passing obliquely downward and back-
ward between the posterior border of the maxilla and the antero-
inferior edge of the squamosal (tig. 1, as, lower pointer). Ante-
riorly it forms the outer wall of the pterygoid fossa; posteriorly it
overlaps the external pterygoid plate of the i)alatine. It articulates
Avith the maxilla, x^alatine, and squamosal; and is pierced by two fora-
mina, the foramen rotundum and the foramen ovale, which, in rare cases,
merge into one. The forame)i rotundum (tig; 1^) is.very inucli larger
than the foramen ovale, and is situated immediately below the anterior
end of the squamosal root of the zygoma. It opens into the anterior
part of the large alispheuoid canal, and sometimes also directly into the
deep sphenoid fossa of the floor of the brain case. In Geomys proper it
is higher up than usual and consequently opens downward into the
alis])henoid canal. The foramen ovale (fig. 4*) is a small slit-like
opening beneath the foramen rotundum; it opens obliquely upward
(and usually backward) into the lower part of the alispheuoid canal.
The foramen ovale presents considerable variation in its position and
relations, affording characters of some value in separating the genera.
In Cratogeomys it is near the anterior- border of the lower part of the
alispheuoid, directly beneath the foramen rotundum and far*below the
alispheuoid canal, which it reaches posteriorly by an obliquely ujjward
and backward course. In Platygeomys and Heterogeomys it is simi-
larly situated, except that it is nearer the middle than the anterior bor-
der of the descending wing of the alisphenoid, and is decidedly nearer
the alisphenoid canal i\w([ foramen rotundum. In Heterogeomys it is
not infrequently confluent on one side with the foramen rotundum. In
Platygeomys it is somewhat jjosterior to the foramen rotundum and
nearer it than in Heterogeomys. In Zygogeomys it is immediately below
and close to the foramen rotundum and sometimes confluent with it;
it is high up and opens directly into the alisphenoid canal. In Geomys
proper it is high up also, and often becomes confluent with the fora-
men rotundum (as in fig. 55''"). In the tuza series its size is unusually
small.

The alisphenoid as a whole articulates with the frontal, squamosal,
maxilla, palatine, basisphenoid, pterygoid, tympanic capsule, and in
some genera with the orbitospheuoid also.

The squamosal is a large and highly important bone in the GeomyidK
(figs. 4, 7, 8, and 0, sq). It overlaps to fi cousiderable extent the other
Ijpnes of tUe padetiea of tUe br^iu CfVse, impartial' gre^^t power of resist-

46

NORTH AMERICAN FAUNA..

[NO. 8.

ance to the vault of the cranium. Autero-iuferiorly it articulates with
the alisphenoid for its entire leuj^th. Postero-inferiorly a long slit-like
vacuity seijarates it from the audital bulla, though iu some cases it is
in contact with parts of the bulla. Posteriorly it overspreads the
sni)erior face of the outer part of the supraoccipital and the mastoid
bulla and sends a lateral arm out sideways (the mastoid arm), which
overreaches and articulates with the end of the mastoid process of the
mastoid bulla. Superiorly it covers the posterior i)art of the frontals
and broadly overlaps the i^arietals for their entire length — actually con-
cealing them iu one species, Cratogeomys- merriami. The squamosal
gives off the posterior root ot^ the zygoma, and articulates with the
jugal. In Zij<jogconiy.s trichopHs and Macrogeomys costaricensis, owing
to the much-reduced size of* the jugal, tliC' squamosal" arm reaches
far forward and articulates directly with the maxilla — a most excep-
tional condition among mammals. Below the squamosal' root of the
zygoma is the elongated and ill-detined glenoid fossa, which is com-
pleted posteriorly and on the inner side by the tympanic bulla. The

Fig. 8. — Skull of very young Geuniyi bursaiius from Elk River, Minnesota. Upper .surface, showing
frontals ankj'losed together, and interparietal inseparable from supraoccipital at birth.

/(•, frontal; ip, interparietal; j, .jugal; n, nasal; ^^a, parietal; pmx, ascending branch of jjreniaxilla;
sq, squamosal; zmx, maxillary root of zygoma.

form of the postglenoid notch varies from broadly (J-s^i^PGd in Platy-
geoniys and some others to narrowly V-shaped uiGeomys hursarius. In
Platygeomys and Cratogeomys the glenoid fossa is i^roduced anteriorly
a long distance in front of the squamosal root of the zygoma.

The mastoid arm of the squamosal enters the outer part of the occip-
ital plane above the mastoid bulla and external to the supraoccipital,
where it forms the whole thickness of the lambdoid crest (see pi. 15,
tigs. 3, 4, 6, and 7). In Heterogeomys it is vertically expanded, taking
a more prominent part than usual in the occiput. The variations in
the squamosal are described later (pp. 66-67).

The parietals complete the roof of the brain case posteriorly (fig. 8,^a).
They do not in-esent any unusual variations in the Geomyidw; they over-
lap the frontal anteriorly and the supraoccipital and interparietal poste-
liorly, and are overlapx)ed for their full length inferiorly by the squa-
mosals, which in Cratogeomys merriami gradually overspreadandconceal
them. The parietals are always separate iu early life, but usually coa-

JAN., 1895.

THE SKULL.

47

lesce in the adult. The temporal impressions may remain permanently
distant, defining- a sagittal area, or they may unite in a prominent
sagittal crest.

The pre.sphenoid is a thin vertical plate of bone bridging the gap
between the basisphenoid and mesethmoid cartilage and supporting,
from its superior surface, the horizontally flattened orbitosphenoids
(figs. 4, 7, audi), 29S*), It is perforated anteriorly by a rather large
opening, which, being opposite the sphenoidal fissure, enables one to see
completely through the skull at this point (figs. 46, H)\ and 55^). A
second fenestrum often exists behind the first, and in Orthof/eoiiiys one
or two small perforations usually occur in front of it. 8ui)eri()rly the
prespheuoid supports the orbitosphenoids (tig. 9, oi), with which it is

t\r Kna

^ ^^

Fin. 9.— Young skull of Cratogeomys merriaiai
removed to sliow floor of brain case.

ac Anterior opeiiing of alisiilienoid canal.

as Alisphenoid bone.

bo Basioccijjital.

bs Basisphenoid.

Condyle of exoccipltal.

cr Cribriform plate of etlimoid.

em External auditory meatus.

ex Exoccipital.

ff Floccular fossa.

fr Frontal.

fro Descending or orbital plate of frontal (tlie
animal is so young tliat the plates of the
, two sides have not yet united below).

' Jugal.

Lachrymal.

inseparably ankylosed; anteriorly it abuts against the mesethmoid car-
tilage and is in contact with the ethmoid and usually the vomer; pos-
teriorly it abuts against the basisphenoid, with which it commonly
becomes ankylosed before the animal is fully adult. The ascending

*Iu fig. 9, which is a young skull, the presphenoid is covered by the orbitosphe
noids, making it appear very much broader than it really is.

Irom Amecameca, Mexico, with vault of cranium

ma Meatus auditorius internus.

7nb Mastoid bulla.

n Nasal.

of Optic foramen.

OS Orbitosphenoid.

pet Petrous part of periotic.

pmx Ascending arm of premaxilla.

ps Presphenoid.

ptf Spheno-pterygoid fossa.

s/ Apex of .sphenoidal fissure.

so Supraoccipital.

iv/ Squamosal.

tb Superior face of tympanic or audital bulla.

zmx Zygomatic root of maxilla.

48 NORTH AMERICAN FAUNA. [no. 8.

wings (vertical plates) of the palatines clasi) the sides of the prespheiioid
iiiferioiiy, rising anteriorly. The ends of the vomer reach it also, clasp-
ing it laterally, bnt never underlying it as in many mammals. The pre-
sphenoid ends anteriorly in a somewhat thickened head, with a disk-
shaped cavity iu front, which receives the hinder end of the mesethmoid
cartilage.

The orbitosphe)ioids are a pair of thin horizontal shelves resting upon
and invariably ankylosed to the upper border of the pres])henoid, and
articulating anteriorly with the orbital plate of the frontal (fig. 9, o.«,
and pi. 17). They are normally perforated near the anterior border by
the optic foramen (tig. 9, of), but in Heterogeomys this foramen is incom-
plete superiorly (pi. 17, fig. 1) except in the young. The antero-external
corner sometimes jirotrudes through the sphenoidal fissure, bends up-
ward, and slightly overlaps the j)osterior border of the descending wing
of the frontal, appearing as a small scale in the bottom of the orbit. This
is most often observed in young skulls. In Zygogeomys, Pappogeomys,
and some forms of Thomomys the ascending tongue of the orbitosphe-
noid completely closes the upper part of the sphenoidal fissure, excejit
a small point at its apex, which is left as a permanent foramen (pi. 18,
fig. 2), and becomes ankylosed to the frontal anteriorly and the alisphe-
noid posteriorly (pi. 17, fig. 2). With these exceptions it does not
appear in the parieties of the cranium, though it may always be seen
crossing the sphenoidal fissure, which it divides into two parts. Ante
riorly the orbitosphenoid invariably articulates with the upper surface
of the presphenoid and the descending wings of the frontal, as already
stated, and sometimes also with the palatine, maxilla, and posterior
edge of the cribriform plate; posteriorly it often touches the edge of
the alisphenoid, to which it becomes fixed in Cratogeomys, Orthogeomys,
Pappogeomys, Zygogeomys, and some forms of Thomomys. .

The relations of the orbitosphenoids anteriorly vary in the several
groups and in some cases are exceedingly difficult to ascertain, owing
to early ankylosis with the presphenoid. In Geomys hursariu.s the
ascending wings of the palatine rise high on the sides of the presphe-
noid and articulate broadly with the orbitospheuoids, but iu most
forms it is uncertain whether or not the palatine is reached. The uncer-
tainty is due to the impossibility of determining how far the orbito-
sphenoid descends anteriorly below the top of the presi)henoid, with
which it is inseparably fused. For the same reason it is uncertain
whether or not the orbitosphenoids always reach the cribriform plate of
the ethmoid. They seem to do so in all cases along the median line, but
I have been unable, even in very young skulls, to find the place of sepa.
ration anteriorly between the orbitosphenoids and presphenoid. In
those genera in which the descending or orbital plates of the frontal do
not meet interiorly behind the cribriform plate, tlio orbitosphenoids
articulate broadly with the cribriform (as in Fappogeomys, Orthogeomys^
aucl Thomomys),

JAN., 1895.] thp: skull. 49

In Geoniys proper the orbitosphenoids are narrower than in any of
the other genera, and do not reach the alisphenoids. In Heterogeomys
and Platygeomys also they usnally fall short of the alisphenoid, though
in extreme cases they sometimes cross the anterior edge of the
alisphenoid. In Cratogeomys and Orfhogeomys they articulate with the
alisphenoid anteriorly for a short distance, but do not follow the upper
part of the sphenoidal fissure, though in Orthogeomys they sometimes
send a tongue upward covering part of the fissure. In Pappogeomys
and some species of Thomomys they go a step further, articulating
firmly and broadly with the alisphenoid and normally closing the
greater x)art of the sphenoidal fissure above the plane of the presphe-
noid. Zygogeomys presents a still more extreme phase, the orbito-
sphenoid almost completely closing the upi)er part of the sphenoidal
fissure and ankylosing broadly with the alis])henoids. From what has
been said it must be clear that the orbitosphenoids play a more
important part than any other bones in determining the form of the
floor of the brain case, for the reason that by their expansion or con-
traction anteriorly they completely change the size and shape of the
sphenoid fossa, which is the most conspicuous of the variable land-
marks of the floor of the brain case, as may be §een on consulting i^l. 17.

T\ie> frontals coalesce very early (i)robably before birth), forming a
single large bone (fig. 8,/r) which constitutes the middle third of the
upper surface of the skull and dips deeply into the orbits, where it
makes important connections with the maxilla and other bones. It
forms the roof of the olfactory chamber of the nasal cavity, and the
roof and part of the side walls of the anterior segment of the brain case.
The main body of the frontal articulates anteriorly with the ethmoid,
nasals, premaxilla, maxilla, and lachrymals, and posteriorly with the
parietals, squamosals, and alisphenoids. It is so extensively over-
lapped by the alisphenoids and squamosals that when viewed from the
outside it appears much smaller than it really is.

The descending or orbital processes of the frontal (figs. 4, o/V, and
9,/ro) reach far downward, burying themselves deeply among the bones
of the base of the cranium and face. They articulate with the anterior
border of the orbitosphenoids, clasp the sides of the presphenoid and
palatines anteriorly, and articulate firmly with the maxillaries. Ante
riorly, except in Thomomys^ Pappogeomys (fig. 56), and Orthogeomys, they
completely encircle the cribriform plate of the ethmoid (with which
they early unite by ankylosis) and meet in the median line below it,
thus reaching around the olfactory lobes of the brain case and forming
the floor as well as the roof and sides of the olfactory fossa. At the ijoint
where the two arms come together in the median line, at the posterior
base of the cribriform plate, a small opening is commonly left which
remains as a perforating foramen passing obliquely forward and down-
ward between the presphenoid and mesethmoid plate, and oijening
anteriorly into the olfactory chamber of the nasal cavity immediately
7433— No. 8 4

50 NORTH AMERICAN FAUNA. [no. 8.

behind the lower part of the fourth eudoturbiuals. In Thomomys (fig.
61), and in the young of most of the other genera (as in Cratogeomys, fig.
9, fro), the orbital plates of the frontal are separated interiorly by the
orbitosphenoids. The variations in the form of the frontal are described
further on (p. 05 and fig. 17).

The ethmoid is a highly complicated bone occupying the posterior
part of the olfactory chamber of the nasal cavity, which it completely
separates from the brain case. No part of it appears on the outside of
the skull. It maybe described under five heads : (1) the cribriform
plate; (2) the mesethmoid; (3) the os planum; (4) the ectoturHnals, and
(5) the endoturhinals. There is no apparent 'crista galli' in the Geoni-
yidw. [The naso- and maxillo-turbinals are completely detached, and
are described under the bones to which they are respectively ankylosed,
namely, the nasal and jnemaxilla.J

(1) The cribriform plate is a transverse i)erforated partition, separat-
ing the olfactory fossa of the brain case from the olfactory chamber
of the nasal cavity (fig. 9, cr). It is nearly circular in outline and
slopes or curves forward from the base upward. Posteriorly, in most
of the genera, its entire circumference articulates (and early anky-
loses) with the frontals, which usually separate it interiorly from the
orbitosphenoids, though the latter may always reach it near the me-
dian line by pushing forward beneath the frontals. To its anterior
face are attached the ectoturbinals, endoturbinals, and mesethmoid.

(2) The mesethmoid bone, or perpendicular plate of the ethmoid, is a
longitudinal median partition incompletely dividing the olfactory cham-
ber into two parts ( fig. 7, me). Its superior border is firmly and insep-
arably ankylosed to the frontal; its posterior to the cribriform plate.
Antero-inferiorly it abuts against the cartilaginous mesethmoid, which
latter reaches forward from the presphenoid and is embraced between
the lateral wings of the vomer, completing the partition between the
two sides of the olfactory chamber. The shape of the bony lamella
varies in the different groups and seems to be quite constant in mem-
bers of the same genus. In Cratogeomys (pi. 18, fig. 4), Orthogeomys
(fig. 60), and Geomys proper (pi. 18, fig. 1), it is somewhat like a half
crescent, with the base above, and the apex pointing to the end of the
presj^henoid, the anterior border being convex downward, tn Platy-
geomys it is similar, except that the upper part is strongly rounded
anteriorly, the upper edge being shorter than that part of the lamella
immediately below it (pi. 18, fig. 5). In Heterogeomys it is relatively
small and strongly convex anteriorly (pi. 18, fig. 3). In Zygogeomys it is
nearly rectangular and the front edge is nearly straight (pi. 18, fig. 2).
In Pappageomys (fig. 57) it is higher than long, and its inferior border
dips down between tlie wings of the vomer — a unique condition.

(3) The OS planum is a thin sheet of bone wliich lines the posterior
part of the olfactory chamber (fig. 10, op). It supports the endoturbi-
nals and binds them together (as may be seen by consulting fig. 10 and

JAN., 1895.]

THE SKULL.

51

pi. 10, figs. 3, 4, and 5 of Geomi/s hursarius, Heterogeomys^ and Zycjogeo-
mys). Inferiorly it articulates with the vertical lamella of the maxillary
which lines the nasal passage, and witli the anterior ends of the ascend-
ing wings of the palatines. Near its lower border (just below the fourth
turbinal), it gives off a lateral shelf, which is firmly ankylosed to the
outer side of the posterior third of vomer. In Cratogcomys its antero-
inferior border is cut off" close to the turbinal folds, giving the latter a

Fig 10.— LoDgitudal vertical median section of trout part of skull of Geomijs hursarius. Meseth-
moid and vomer removed to show turbinated bones.

1 Anterior i>alatine foramen.

2 Inci.sive foramen.

3 Vacuity in front of presphenoid (present in Geomys hursarius and tuza only. It is partly over-

lapped posteriorly by the ascending wing of the vertical plate of the palatine, ap.).

4 Presphenoid fcnestrum. Present in all species.

5 Upper part of sjihenoidal fissure.
H First or superior endoturbinal.
2t Second endoturbinal.

3< Third endoturbinal.

it Fourtli endoturbinal.

ap Ascending wing of vertical plate of palatine.

/*• Frontal.

mt Maxilio-turbinal.

nix Maxilla (the upper pointer rests on the maxillary surface of the narial passage, the lower on the

sawed body of the bone).
n Nasal.
nt Naso-turbinal.
op Os planum.
pi Palatine (the upper pointer rests on the palatine face of the narial passage, the lower on the sawed

horizontal body of the bone).
pmx Prcmaxilla.
ps Presphenoid.
vr Vomerine ridge of os planum (unites with the lateral wing of the vomer) .

particularly neat and finished appearance (pi. 19, fig. 6). In Geomys
hursarius, on the other hand, it falls directly downward from the first
turbmal, projecting as a thin sheet considerably in front of the others
(fig. 10 and pi. 19, fig. 3).

52 NORTH AMERICAN FAUNA. fNo.g.

(4:) The ectoturbinals * arise from the upijer and outer corners of the
cribriform plate and occupy a small chamber at the maxillary root of
the zygoma, incased chiefly by the frontal and maxillarybones. When
the lachrymal is removed, they may be seen from the orbital side.

(5) The endoturbinals * arise from the outer sides of the anterior face
of the cribriform plate (on the inner side of the ectoturbinals) and ])ro-
ject into the nasal chamber (lig. 10). They are four in number through-
out the family. Their outer sides are continuous with and form a part
of the OS planum. The first or upi^ermost is always the largest, longest,
and most broadly expanded anteriorly. The others decrease in length
from above downward, and are broadest in the middle or posteriorly.
The fourth or lowermost is broader and shorter than the two middle ones.
The first or uppermost is the only one that need be considered from the
standpoint of variation of form in the several gi'oups. Its front border
usually slopes strongly backward (from above downward), as in Flatygeo-
mys, Cratogeomys, and Zygogeomys; but in Heterogeomys it is straight or
slightly emarginate, vertical, and very broad, and carries with it the
second fold (see i^l. 19, fig. 5). In Platygeoniys it is long and relatively
slender, and its apex projects anteriorly behind the posterior border of
the nasoturbinal (pi. 10, fig. 7). In Zygogeomys also it is pointed and
projects far forward (pi. 19, fig. 1). In Geomys hursarius it is rather
bluntly rounded (fig. 10, and pi. 19, fig. 3).

The vomer is a long and narrow plate of bone, cleft above and bifur-
cate posteriorly, which forms the lower part of the longitudinal verti-
cal septum between the lateral chambers of the nasal cavity (fig. 7, v).
It consists of a median plate and two wings. The median plate is
embraced inferiorly between the wings of the vomerine sheath (which
rises from the floor of the premaxilla and extreme anterior part of the
maxilla). Superiorly it is split lengthwise from above, forming the two
wings, between which the mesethmoid cartilage is received. These
wings are narrowed j)osteriorly and reach the front end of the pre-
sphenoid, which they clasp laterally, but they do not appear on the
inferior surface of the presphenoid, as they do in most mammals. Pos-
teriorly the wings of the vomer separate slightly and are not united
inferiorly. On the outer side they are inseparably united with the
OS planum just below the fourth endoturbinal, thus continuing ante-
riorly the roof of the narial i)assage, which is here sharply separated
from the olfactory chamber above. The vomer articulates with the
premaxilla, maxilla, ethmoid, presphenoid, and palatines.

ThOi pterygoids are more or less quadrangular vertical plates, forming
the lateral walls of the posterior nares (figs. 4 and 7, pt). Anteriorly
they articulate Avith the vertical plates of the palatines ; superiorly they
are firmly ankylosed to the basisphenoid, and usually also with the
posterior downward extension of the transverse arm of the alisphenoid.

* These terms are adopted from Dr. Harrison Allen's admirable paper on the Eth.-
moid.— (Bull. Mus. Comp. Zool., Cambridge, X, No. 3, 1882, 136.)

JAN., 1895.] THE SKULL. 53

They commonly develop a hamtilar process (figs. 4 aud 7, A), which
curves upward and reaches or nearly reaches the audital bulla (except
in Heterof/eomys). The inferior surface of the pterygoid is usually flat-
tened, either horizontally or obliquely ; it may be of uniform breadth
(fig. 11^), or much broader anteriorly than posteriorly (figs. 11^ and IV).
It reaches its maximum length aud slenderness in Zygogeomiis (fig. 11');
its maximum breadth and shortness in Macrogeomys (fig. IP), The
two arms may be divergent posteriorly, convergent posteriorly, or par-
allel.

Fig. 11.— Principal tjpes of palatopterygoids.

1. Zygogeoinys trichopus. 2. Oeomi/s lutescens. 3. Geomys bursarius.

4. Heterogeomya hispidus. 5. Alcfbrogeomys heterodus.

In the share they take in the formation of the palato-pterygoid plates
on the roof of the niouth, and the manner of articulation with the pal-
atine bones, the pterygoids present five types, as follows :

(1) They completely surround the postpalatal notch like a horseshoe,
meeting or so nearly meeting anteriorly that at most a narrow spicule
of the palatine reaches the notch in the median line. This type occurs
in the genus Zygogeomys only (fig. 11').

(2) They form the whole or practically the whole of the sides of the
postpalatal notch, but are separated anteriorly by the full breadth of
the notch itself. This is the commonest type and prevails in the genera
Geomys nudCratogeomys (fig. 11').

(3) They are lingulate in shape and do not reach the base of the jiost-
palatal notch, the palatine bones extending out a considerable distance
to meet them. This is the ordinary condition in Geomys bursarius (fig.
IP).

(4) They are very much reduced, forming only the terminal part of
the palato-pterygoid plates, the palatine part of which is greatly elon-
gated. This condition obtains in Heterogeomys (fig. 11*).

(5) They are short, broad, and abruptly upturned, capping the ends
of the very broad palatines. This type is restricted to Macrogeomys
(fig. IP).

The palatine bones contribute an insignificant part to the external
surface of the skull (fig. 12, pi), but internally their connections are
extensive and important (fig. 7, pi, and fig. 10, pi and aj)). They early
unite (probably before birth) in the median line, forming a single bone,
which may be described as consisting of a body, two vertical plates, and
two lateral wings or external ptergoid plates. The body or horizontal

54 NORTH AMERICAN FAUNA. [no. 8.

part enters the roof of the mouth posteriorly, forming a wedge between
the hinder part of the maxillaries, and never reaching further forward
than the middle molars (fig 12, pi). This part is cut away posteriorly, so
that its inferior surface is on two i)lanes. Anteriorly it is continuous
with the plane of the bony pala'.e; posteriorly with the pterygoitls.
The break in the palatines between these two i)lanes occurs suddeidy
between the posterior molars, forming a step or pit on each side between
the last molar and a median azygos iDrojection of the palate, which con-
nects the two more gradually. Posteriorly the palatals may terminate
oijposite the anterior end of the postpalatal notch (as usual m Crato-
geomys), or they may extend out a short distance beyond the apex of
the notch (as m Oeomys proper), or they may i^ush back still farther,
forming more than half of the side walls of the notch (as in Hetero-
geomys), or they may fail to reach the notch at all, the pterygoids com-
ing forward to the median line (as in Zygogeomys). [See fig. 11 supra.]

The vertical jilcites are tliin lamellae, which reach upward on each side
from the body of the bone to the i)resphenoid, surrounding the middle
section of the narial passage between the maxilla and pterygoid (tig.
7,2)1)- Their upper borders reach the basisphenoid anteriorly and are in
contact with the i^resphenoid for its entire length ; anteriorly they clasp
the sides of the presphenoid and articulate with tlie ethmoid and
frontal — the descending processes of the latter overlapping their ante-
rior prolongations. The front border of the vertical plate of the pala-
tine, on the side of the narial passage, articulates with the correspond-
ing part of the maxilla; the hinder border with the pterygoid. In
Gcomys bursarius the vertical plate rises anteriorly in an ascending iving
which hugs the prespenoid anteriorly and articulates broadly with the
orbitosphenoid, frontal, and maxilla (fig. 10, a.p).

Posteriorly the body of the palatine sends oft", on each side, a lateral
wing — the external pterygoid plate — which pushes its wjiy around behind
the maxilla and along the inner side of the descending wing of the
alisphenoid as ftir as the i^oint where the Litter is joined by the trans-
verse arm of the same bone (immediately below the alisphenoid canal),
and sometimes sends a spicule backward to the audital bulla (fig 12, epl).
The external pterygoid plate of the palatine thus forms the outer wall
of the pterygoid fossa inferiorl}^ It is completely overlapped exter-
nally by the descending wing of the alisphenoid, except along its infe-
rior margin, which projects slightly below the alisi)henoid, thus apx)ear-
ing on the outer side of the skull (fig. 4, epl).

The palatines articulate with the maxilla, pterygoids, alisphenoids,
basisjihenoul, presphenoid, frontals, vomer, and ethmoids and some-
times also within the orbitospheuoids and the tympanic bulla*.

The maxilla is the largest, and after the ethmoid the most compli-
cated bone of the skull, and comprises, roughly speaking, about one-
third of the entire cranium (fig. 12, mx). It i)rimarily consists of two
parts, which are firmly united by ankylosis in very early life (probably

i

JAN., 1895.]

THE SKULL.

55

before birth), forming a single strong bone for the support of the grind-
ing teeth. It articnhites with nearly all the bones of the face and with
those of the anterior segment of the brain case, as follows: Anteriorly
with the premaxilla, ethmoid and lachrymals; superiorly with the pre-
sphenoid and frontal; posteriorly with the palatines and alisphenoid,
and externally with the jugals. The maxilla forms nearly the whole
of the roof of the mouth, the palatines entering it merely as a wedge
from behind. The densest and hardest part of the skull, after the
floor of the premaxilla, is the median part of the maxilla between the

FiQ. 12.— Under side of young .skull of Cratogeomijg merriami. (Specimen from Amecameca, Valley
of Mexico.)

als

Alisphenoid.

VIS Mastoid process of squamosal.

amx Alveolar border of maxilla.

mx Maxilla.

bo

I?asioccii)ital.

n Nasal.

hs

Basiisphenoid.

pi Palatine.

c

Condyle of exoccipital.

pmx Premaxilla.

epl

External pterygoid plate of

palatine.

pp Paroccipital process of exoccipital

fr

Foramen rotundum.

pt Pterygoid.

9/

Glenoid fos.sa.

2itf Pterygoid fossa.

'/

Incisive foramen.

smf Stylo-mastoid foramen.

Jugal.

tb Tympanic or audita! bulla.

ma

"External auditory meatus.

zmx Zygomatic process of maxilla.

mb

Mastoid bulla.

molariform teeth. The infraorbital canal is deeply imbedded in the
maxilla and is very long, reaching back from near the premaxillary
suture on the side of the muzzle to the bottom of the orbit. In the
Geomyidcv it never perforates the zygomatic root of the maxilla, but
passes deeply behind it.

The maxilla gives off anteriorly a vertical lamella, which rises from
the median line of the tloor of the nasal chamber and projects forward
a short distance into the posterior part of the vomerine sheath of the
premaxilla (fig. 13, ms). It is split lengthwise to receive the posterior

56 NORTH AMERICAN FAUNA. [no. 8.

part of tlie median plate of tlie vomer, but the resulting wings do not
spread apart as in the premaxillary part of the vomerine sheath.

On each side of the nasal passage the body of the maxilla gives off
a thin vertical plate or lamella, which may be termed the internal ver-
tical plate of the maxilla. It forms a lining for the narial passage and
articulates above with the lower edge of the os j^lanum of the endo-
turbinal. The infraorbital canal passes for nearly its entire length
between this thin plate and the main part of the maxilla.

The premaxilla is a single bone in the Geomyidce (its two halves unit-
ing before birth, fig. 12, pmx). It constitutes the greater part of the
rostrum and forms the floor and lateral walls of the anterior half of the
nasal chamber. Superiorly it embraces the nasals and articulates with
the frontal and the maxillary root of the zygoma ; laterally it articu-
lates with the outer side of the maxilla a little anterior to the plane of
the infraorbital foramen; inferiorly it articulates with the maxilla
posterior to the middle of the rostrum, and reaches far enough back-
ward to inclose the incisive foramina (fig. 12, if) in all except Zygo-
geomys trichopus. Anteriorly it is perforated on the median line by the

Fig. 13.— Longitudinal vertical section of nasal chamber of Gratogeomys merriami. The vomer
has been removed to show the vomerine sheath and anterior turbinated bones.

mx Maxillary.
n Nasal.

1 Anterior palatine foramen.

2 Incisive foramen.
mt Maxilloturbinal.

WIS Maxillary part of vomerine sheath (which
passes anteriorly into the premaxillary
part of the sheath).

nt Na.so-turbinal.
pnix Premaxilla.

vs Vomerine sheath of premaxilla.

anterior palatine foramen, which descends from the floor of the nasal
chamber to the roof of the mouth, immediately behind the incisors
(figs. 7, 10 and 13 '). On the inner side it supports the maxillo-turbinals
and the vomerine sheath, which latter structure attains a high devel-
opment in this group, particularly in Flatygeomys and Gratogeomys.

The vomerine sheath (fig. 13, vs) is a double lamella rising from the
floor of the premaxilla on the median line and projecting into the nasal
cavity. It is elongated antero-posteriorly, reaching from the hinder
end of the premaxilla forward over half or two-thirds the floor of
the bone. Posteriorly it receives the anterior end of the corresponding
(but very much smaller and narrower) part of the maxilla; superiorly
it receives the median vertical iilate of the vomer.

JAN., 1895.]

THE SKULL.

57

The maxillo-turbinal, or inferior turbinated bone (figs. 7, 10, and 13, mt),
is the lower of the two turbinated bones of the anterior half of the
nasal cavity (the upper being attached to the nasal). It is nearly hori-
zontal, though usually sloping downward posteriorly, and is attached
to the middle part of the inner side of the premaxilla; its free posterior
end projects slightly over the front of the maxilla.

The premaxilla articulates with the nasals, frontal, maxilla, vomer,
and ethmoid.

The jugal completes the zygomatic arch, and is always restricted to
the horizontal ])art, never reaching down posteriorly into the glenoid
fossa, and never creei)ing up anteriorly toward the lachrymal (figs.
9 and 12, j). But its variations in size and form are remarkable (fig. 14
and pi. 13). In some species it is very large and broadly expanded
anteriorly (fig. 11'); in others it is reduced to an insignificant splint,
and the zygomatic arch is complete without it (fig. 14*'). It is com-
monly larger and broader in the male than the female, and sometimes

Fig. 14. — Left zygoma, showing several types of jugal.
\. Platygeomystylorhinus. I 4. Geomysbtirsarius.

2. Heterogeomys hispidus. 5. Cratogeomys perotensis.

3. Macrogeomys hetc.rodus. I 6. Zygogeomys trichopus.

varies greatly in species of the same genus and even in the same sec-
tion. Thus, in Platygeomys it is greatly expanded in gifmnvrus and
tylorhinus, and is slender throughout in planiceps. Similarly, in Crato-
geomys it IS broad anteriorly in merriami, ful,vesce7is, and castanops,
while in perotensis it is slender and small in every way.

The lachrymal is a small L-shaped bone, consisting of a vertical scale-
like part, which closes the vacuity between the frontal and maxillary
root of the zygoma at the inner corner of the orbit; and a thickened
horizontal part which projects outward from the frontal on the upper
surface of the skull and articulates also with the maxillary root of the
zygoma. Its distal end is sometimes elongated and slightly recurved,
and projects freely over the corner of the orbit. The principal or ver-
tical part of the lachrymal is grooved vertically on its outer side, just
anterior to the orbital face, for the lachrymal duct which passes down
into tlie nasal chamber.

The nasal bones fill the interspace between the ascending arms of
the premaxilla on top of the rostrum, thus completing the roof of the
nasal cavity, which they slightly overhang anteriorly (figs. 8 and 9,m).

58 NORTH AMERICAN FAUNA. [no. 8.

They are commonly ankylosed together in middle life, and not infre-
qnently become ankylosed to the frontal.s also. Their actual length
varies greatly in the different species. They are shortest in Gratogeo-
7nysestor and longest in Zygor/eomys triehopiis and Geomys tuza. They
are commonly truncate wedge-shaped; the increase in breadth from
behind forward may be gradual or abrupt. In the latter case the expan-
sion is usually near the middle. In the Geomys tuza group the shai)e
of the nasals is peculiar. They are very long and are constricted near
the middle, giving them an hour-glass shape. In most of the genera
{Geomys^ Gratogeomys, Platygeomys, Zygogeomyn) the nasals are nearly
flat, though they are always more or less decurved anteriorly and
rounded off laterally in front. But in some groups (notably in Hetero-
geomys) they are broadly and highly arched anteriorly, giving them
an inflated appearance. This elevated part of the nasal suijports the
naked nasal pad or callosity. Inferiorly the nasals give oft" a descend-
ing lamella, the nasoturhinal bone, which is elongated antero-posteriorly
and is broadest behind.

The nasals articulate with the premaxilla, frontal, and ethmoid.

The tympa7io-2>eriotio capsule incompletely fills a broad gap in the
posterior segment of the skull, between the basioccipital and squamosal
(ligs. 4, 7, and 9). It is held in place by several bones with which its
connection is more or less intimate, but is never ankylosed to any of
them except in extreme age, when the mastoid process of the mastoid
bulla sometimes unites with the mastoid process of the squamosal. Its
principal stays are the exoccipital and the mastoid iDrocess of the squa-
mosal, between which the mastoid bulla is firmly grasped posteriorly.
In addition to these supports, the inner border of the auditalbnlla com-
monly fits into a groove on the outer edge of the basioccipital, and the
apex of the bulla rests against the base of the horizontal arm of the
alisphenoid near its junction with the basisphenoid. The tympano-
periotic mass as a whole thus has four normal attachments, two of whicli
hold it firmly in place, while the others simply steady it in its position.
In old age the lower edge of the squamosal sometimes reaches the upper
side of the bulla and presses firmly against it.

The tympano-periotic capsule consists of three parts, firmly ankylosed
together: (1) the tympanic, or audital bulla; (2) the petrous, or periotic
proper; (3) and the mastoid bulla. Of these, the mastoid is posterior
to the others, both of which are inseparably ankylosed to its anterior
face. The tympanic protrudes from the base of the skull, forming the
audital hulUv. The petrous projects into the brain case and contains the
organ of hearing. Xo suture or other line of demarcation indicates
the exact i)lace of meeting of the mastoid with either the petrous or
tympanic, but anteriorly the line of nnion between the two latter is
always distinct. The three elements may be described as follows:

(1) The tympanic or audital bulla is almost wliolly inferior, projecting
from the under surface of the outer segment of the cranium between the

.iAN.,1895.] THE SKULL. 59

basioccipital aud squamosal (figs. 4 and 12, tb). Anteriorly it is bounded
by the fonimen lacerum medium basis cranii, in front of which is the
transverse bar of the alisplienoid. Superiorly it is separated from the
squamosal by a long, irregular vacuity reaching upward and backward
from the foramen lacerum medium to the tube of the external meatus,
which latter articulates with the squamosal. Posteriorly it abuts against
the mastoid process of the squamosal above, and the exoccipital below,
and is continuous with the mastoid bulla. Externally it sends off at
right angles a long tube which partly tills the postglenoid notch aud
opens just behind the posterior angle of the zygoma (fig. 12, ma). This
is the external auditory meatus (fig. 4^). The tube of the meatus
curves forward and somewhat upward as well as outward, and forms
the posterior boundary of the glenoid fossa, against which the condyle
of the jaw strikes tliiring the to and fro movement of mastication. The
adjoining upper part of the outer side of the bulla forms the inner side
of the glenoid fossa. It is thus ai)j)arent that this fossa, while mainly
in the squamosal, is completed posteriorly by the tympanic bulla. The
inner side of the bulla fits into a longitudinal groove on the outer edge
of the body of the basioccipital, and the extreme anterior end just above
the entrance of the Eustachian canal rests against the horizontal arm
of the alisphenoid, which sometimes, as in Cratogeomys, sends back a
small tongue of bone to cover its apex. The canal for the internal
carotid artery is absent. On the inferior surface, between the mastoid
and tynq^anic bulhe, is a small oiiening, the stylomastoid foramen (fig.
12, S7nf). The tympanic bulla arches over and protects the tympanum
and the openings leading into the internal ear.

(2). The petrous, or periotic i)roper, in which is lodged the organ of
hearing, is not visible from the outer side of the skull, but is conspicu-
ous on the inner side (figs. 7 and 9, pet)^ where it is saddled upon the
tympanic capsule, which it does not completely cover, a considerable
l)art of the bulla protruding anteriorly (figs. 7 and 9, tb). The line of
demarcation between the two is always evident. The anterior border
of the petrous begins near the middle of the inferior margin of the
inner surface of the bulla and curves upward and forward to the front
end of the ridge that separates the inner from the superior surface of
the bone. On the outer side of this ridge it turns back, forming a deep
reentrant angle, at the apex of which is a small foramen. The petrous
is commonly described as a very hard bone. It is not so in the Geomyidw,
but is soft and spongy, being made up of cancellous tissue like the rest
of the tym})ano periotic capsule. It contains the cochlea (coiled in a
compact cone of 4i turns), the semicircular canals, and the three small
bones of the internal ear — the malleus, incus, and stapes. Tlie jjetrous
may be described as presenting two surfaces, a superior and an inner.
The superior surface is narrow, slopes downward from behind forward,
and is scooped out lengthwise. It is more or less completely separated
from the inner surface by a ridge, which in some forms is sharply

60 NORTH AMERICAN FAUNA. [no. 8.

marked ; in others is incouspicuous. This ridge i)resents various degrees
of developmeut iu the different groups. It is rounded off in Platy-
geoniys, hnt is elevated into a distinct crest in Crafof/eomys, Zygogeomys,
Heterogeomys, and Geoinys proper (pis. 17 and 18), It usually reaches
upward and backward to the upper part of the audital mass, but in
Heterogeomys it fails posteriorly, but forms a sharply elevated ridge
from the plane of the flocculus downward (pi. 18, fig. 3). The inner face
of i\iQ, petrous is always perforated by the internal auditory meatus (fig.
7^ and tig. i), ma)^ above which is a depression called the flocctdar
fossa {Hg. 7^ and lig. 9, ,^). The _^occM/flr/0''>'S«' varies in size and form
in the several genera. Its position is always above and posterior to
the internal meatus, from which it is separated by an elevation which
sometimes amounts to a strongly developed ridge (see pis. 17 and 18).
The ridge is marked in Cratogeomys, but not iu Platygeomys, Heterogeo-
mys, or Geomys proper. In Zygogeomys it is not only present, but a
suiiplemeutary ridge bounds the floccular fossa posteriorly, leaving
another depression behind it, so that the bone presents the appearance
of having two floccular fossa? (pi. 17, fig. 2, and pi. 18, fig. 2).

(3) The mastoid bulla forms the hindermost part of the auditory
apparatus (fig. 4, mh). It appears on the outer side of the occipital plane
as a more or less rounded subtriangular mass, convex posteriorly, with
the base toward the median line and the blunt ^.i^ex {mastoid process
proper, fig. 4, m) directed outward. It is grasped and held in place by
the paroccipital process of the exoccipital below (figs. 4 and 12, pp), and
the long mastoid process of the squamosal above (fig. 4, ms). The for-
mer fits into a notch on the under side between the mastoid and audital
bull re. The latter reaches far outward and curves down upon the head
of the mastoid process, which it overreaches enough to effectually oppose
the action of the exoccipital. The mastoid bulla, viewed from behind,
differs considerably in form in the several genera, and presents specific
differences also (pi. 15, figs. 3-7). It is short and rounded in Zygo-
geomys and Geomys (particularly in the tuza series). It is strongly
triangular in Macrogeomys doUchoeephalus ; triangular with a con-
stricted and elongated neck in M. keterodus, and much produced
laterally with the inferior border concave in Platygeomys. Internally
the mastoid bulla is made up of fine cancellous tissue.

The mandible is usually a large and heavy bone, strongly marked by
processes and ridges for the attachment of the powerful muscles that*
move it. To be understood, it should be studied as a part of the cut-
ting and slicing machine, for it consists, on each side, of a curved
beam or plate built expressly to carry the ponderous chisel-edged inci-
sors and the series of parallel cutting blades of the lower molariform
teeth. The two halves are joined together by an elongated symph3^sis
which admits of a certain amount of movement, and the adjustment is
aided by a transverse muscle which helps bind the jaws together above
the posterior half of the symphysis. Each half of the mandible is

JAN., 1895.]

thp: skull. 61

strongly and rather shortly curved upward longitudinally, and is
broader behind than in front; it also curves outward. There is no
separation into horizontal and ascending rami, although when viewed
from the inner side the condylar and coronoidpart might be regarded as
forming an ascending ramus. The outer side gives off" posteriorly, at
right angles to its axis, a strongly defined angular process which is
alwaysimportant and in some forms, particularly in Flatyf/eomys, attains
enormous i^roportions (pi. 10, fig. 8). Between the angular process and
condyle is a subglobular prominence which covers the root of the long
incisor. The coronoid process is broad at the base anteroposteriorly;
its apex is hamular and rises above the plane of the condyle. In some
forms (notably in Platygeomys) a strong shelf-like ridge runs from the
anterior base of the coronoid to the angular process. The masseteric
fossa is always well defined and reaches anteriorly to the plane of the
front of the i)remolar. On the outer side of the last two molars is a
large and deep pit for the insertion of the principal part of the temporal
muscle (pis. 1-7). The dental foramen enters the, ramus just behind
this pit and just below the condylar process. Behind the symphysis,
inferiorly, is a flange-like prominence for the insertion of the digastric
muscle. The principal differences in the form of the mandible as a
whole result from the amount of spreading posteriorly and the degree
of development of the angular processes. The various types, as seen
from below, are shown on PI. 10. In some cases the base of the angular
process is notched anteriorly, as in Geomys mohilensis (pi. 10, fig. 2.)

3. CHANGES WITH AGE.

Throughout the Geomyidce, except in Pappogeomys, and some species
of Thomomys, the form of the cranium as a whole, and the pattern of
the sutures on the upper surface change greatly with age. The change
marks the transition from immaturity to maturity — from the generalized
type that stands for the group to tlie specialized type that bears the
impress of the species. When the skull of a species fails to show
marked differences with age, that species may be set down as a gener-
alized type — one that is probably but little removed from the ancestral
line. For this reason Pappogeomys hulleri is looked upon as very near
the trunk line of the group.

The principal changes in the form of the skull as a whole resulting
from age are : The broadening out of the zygomatic arches, elongation
of the rostrum, expansion of the squamosal, and development of the
crests and ridges that come with maturity. The anterior or maxillary
root of the zygoma at first slopes strongly backward in all species, and
the arches themselves are narrower anteriorly than posteriorly (as is
the rule in adults of Thomomys). With advancing age they spread apart
anteriorly until in most species they are much broader anteriorly than
posteriorly. At the same time the maxillary root stands out more and
more squarely until it sometimes forms almost a right angle to the axis

62

NORTH AMERICAN FAUNA.

of the skull. Tlie remarkable growth of the squamosal has beeu already
described. Before birth the ascending branches of the premaxilla
end about on a plane with the nasals (sometimes anterior to it), but
they soon push back over the frontals, attaining their permanent
relations at an early age. The muzzle increases in length from birth to
maturity. This may be roughly expressed in the growth of the nasals
as shown in the accompanying figure (fig. 15). In a young skull of
Zygof/eomys trichopus the nasals form 37 percent of the total length of
the u])per surfiice of the skull, wiiile in an adult skull of the same spe-
cies they form 44 i)ercent of the total. The frontal, like the inter-
parietal, though to a less degree, suffers from the encroachment of the
parietals, and in some species from the inordinate growth of the squa-
mosals also. In young skulls the frontal is broad posteriorly and

Fig. 15. — Zyijogeoinys trichojjus, showing changes with age. a, Yonng; b. yovmg ailult; c, adult.

forms an important part of the roof of the brain case, as seen from above
(figs.8, loff, and 16b). In old skulls it is reduced posteriorly, in most
species, to a small wedge between the greatly expanded anterior extrem-
ities of the parietals and squamosals (see pi. 1; pi. 15, fig. 2; and text
tig. 15, c, for adults of same species figured in figs. 8, 15, a and 10, h).

The changes in the suture pattern result mainly from the growth of
the parietals both anteriorly and iiosteriorly, with consequent shrink-
age of the interparietal, and the pi'ogressive development of the squa-
mosal. The decrease in the size of the interparietal corresponds with
the movement of the temporal impressions, whicli approximate with
age, and in many species finally meet in a sagittal crest. The parietals
not only tend to cover the interparietal by meeting posteriorly above
it, but anteriorly they overlaji the sides of the frontal, altering its shape
entirely. The progressive development of the squamosals in some

JAN., 1895.]

THE SKULL.

63

species, as elsewhere shown, is evou more remarkable than that of the
parietals.

Fig. 16.— Skull of very J'oiing Heterogeomys torridus Irom Motzorongo, Mexico (So. 63643).
a, lower surface; &, upper surface. For key to bones see figs. 8 and 12.

4. COOSSIFICATION OF THE PAIRED BONES.

Nearly all the paired bones that meet in the median line become firmly
ankylosed together before birth or in very early life. Those that are
thus coossifled are the preniaxillaries, masillaries, palatines, parietals,
frontals, and frequently the nasals also. Of these, all except the parie-
tals and nasals are ankylosed before birth (see figs. 8 and 16).

The single bones forming the basicranial axis are early ankylosed
with the adjoining paired bones of the same segments. Thus the pre-
sphenoid is inseparably united with the orbitosphenoids; the basisphe-
noid with the alisphenoids and pterygoids; the basioccipital with the
exoccipitals. The union of the lateral with the median elements of
the sphenoidal segments occurs before birth; that of the occipital seg-
ment later. The exoccipitals are always distinct in early life (figs. 12
and 16), but soon become ankylosed with the basioccipital below and
the supraoccipital above. The latter, except in a few species, is insep-
arable from the interparietal. The parietals in adult life are commonly
ankylosed with the squamosals.

5. CRANIAL VARIATIONS— DEPARTURES FROM THE TRUNK LINE.

In external appearance the members of the family Geomyidw are very
much alike, but in cranial characters they present several marked gen-
eric; types. The skulls of these types ditler in size, massiveness, and
degree of development of the crests, ridges, and processes from the
small, thin, and smoothly rounded skulls of Geomys fexensis and bulleri

64 NORTH AMERICAN FAUNA. [no. 8.

to the huge angular crauiums of Platygeomys ffymnurus and Cratogeomys
merriami; and the large, massive skulls differ in the breadth of the
cranium and lateral production of the angle of the mandible from the
extraordinarily broad and flat Platygeomys gymnunis to the long and
narrow Orthogeomys scalops and Macrogeomys doUchocepliahis. The skulls
differ further — and this is much more important — in the relative devel-
opment and relations of certain bones which here assume proportions
and conditions previously unknown. Most if not all of these remark-
able extremes of form are clearly secondary modili cations resulting
from the highly specialized types of dental armature possessed by the
animals, as shown later.

The parts of the skull that exhibit the widest A'ariation and play the
most important part in giving to each type its peculiar impress or physi-
ognomy are the zygomatic arches, the roof of the brain case, and the occiput.
The individual bones that present the greatest range in size and form
are the frontal, squamosal, jugal, pterygoid, and mandible.

The zygomatic arch varies exceedingly in size, form, and the relative
development of its comi)onent elements, according to its importance
as a support for the jugal part of the masseter muscle. It may be small
and slender, with the horizontal part reduced to a mere rod, as in Pappo-
geomys buUeri (pi. 13, tig. 15) and Orthogeomys latifrons (t)1. 13, fig. 16),
or it may be large and massive, with the angle and horizontal arm broadly
expanded, as in Platygeomys (pi. 13, figs. 1 and 2), Cratogeomys (pi. 13,
fig. 4), and Heterogeomys (pi. 13, fig. 20). The area for the attachment
of the jugal part of the masseter muscle may be small and posterior
(fig. 49, jo), or large and extending the full length of the outer side of
the zygoma (fig. 50, jo). The arches may be small and narrow with their
outer sides nearly parallel, as in Macrogeomys dolichocephalus (pi. 5) and
Orthogeomys scalops (pi. 19, fig. 1), or they maybe massive, widely spread-
ing, and broadly divergent anteriorly, as in Platygeomys (pi. 3) and
Cratogeomys (pi. 2). The ratio of their breadth to the basal length of
the skull varies from 54 j^ercent in Macrogeomys dolichocephalus to
upward of 88 percent in Platygeomys tylorhinus, a difference of 34
percent. They may be slightly or strongly decurved; the horizontal
part may be lowest anteriorly as in Platygeomys gymnurus (pi. 13, fig. 2),
or highest anteriorly, as in Macrogeofnys dolichocephalus (pi. 13, fig. 19),
and the angle may be small (pi. 13, figs. 15, 16, and 24) or broadly
expanded (pi. 13, figs. 1, 2, 4, 17, and 18). The expansion, which
normally covers the antero- external angle, as in Platygeomys, Crato-
geomys, and Heterogeomys (pi. 13, figs. 1, 2, 4, etc.) may be drawn
backward so as to occupy the middle i^art of the horizontal arm,
as in Macrogeomys costaricensis and dolichocephalus (pi. 13, figs. 19 and
23). In the latter the zygomatic arch presents a i^eculiarity not
observed in any other member of the grouj). It is narrow, broadly
rounded antero externally, without the expansion of the angle common
to Cratogeomys, Platygeomys, and Heterogeomys, but with a moderate

JAN., 1895.] THE SKULL. 65

expansion near the middle of the liorizontal arm. This expansion is
wholly on the upper or orbital side, and is restricted to the maxillary
part of the arch, wliicli here reaches innch farther back than usual.
On comi)aring the arch caretully with that of 3facro(jeomys heterodus a
curious explanation is suggested, namely, that in the extreme elonga-
tion of the skull of M. dollchocephalus the anterior root of the zygoma
has been moved forward (the post<;rior root being fixed), increasing the
length of the maxillary arm, decreasing the breadth of the arch, oblit-
erating the antero-external angle, elongating the laminar expansion on
the orbital side, and carrying its highest point backward to or behind the
middle of the orbito-temporal fossa (pi. 13, fig. 19, and text fig. 49). At
the same time the upper anterior angle of the jugal has been rounded off,
and the maxillary and squamosal arms of the zygoma have nearly clasped
hands above it. Furthermore, the zygomatic arch as a whole has been
lifted up by the main body of the masseter muscle and as a consequence
the anterior end has been raised higher than the posterior (fig. 49, which
should be contrasted with the corresponding view of Platygeomys gym-
nurus, in which the front of the arch is drawn down, fig. 50).

The form of the occiput as a whole varies considerably in the several
groups. In the less specialized forms, such as Geomy.s texcnsis, arenarins,
and breviceps, and Fappogeomys bullcri {pi. 15, fig. 5), it is rounded and
bulges i)osteriorly to such a degree that the lambdoid suture is left a
considerable distance in front of it. In Zygogeomys, Crafogeomys, and
Geomyn hursarius and lutesceiis, the occiput is squarely truncated. In
Heterogeomy.s (pi. 15, fig. 4), Macrogeomys (pi. 15, fig. o),and Orthogeomys
it is rather high and slopes strongly forward; and in Hetc yog corny nit is
particularly high above the mastoid bulhe. In Platygeomys it is de-
pressed and elongated transversely and presents a unique appearance,
the broad flange-like paroccipital processes curving strongly backward,
defiiung laterally a deep basin-shaped cavity which is completed above
by the overhanging lambdoid crest (pi. 15, fig. 7).

The form of the frontal as seen from above varies greatly in the dif-
ferent groups. lu Geoniys, CratogcomySj Platygeomys, and Zygogeomys it
is narrow and is strongly biconcave between the orbits, with the orbital
margins more or less thickened and raised, leaving a longitudinal
depression or groove between them (fig. 17^). In Heterogeomys it is
broad, flat on top, moderately biconcave between the orbits, and shield-
shaped posteriorly, owing to the elevated temporal ridges ; but the
orbital margins are not rounded, thickened, or raised (fig. 17^). In
Macrogeomys it is moderately broad and deeply constricted between
the orbits posteriorly. Immediately behind the constriction it exi)ands
abruptly at right angles to its axis, forming well-marked postorbital
processes which are capped by the apex of the alisphenoid and partly
overlapped posteriorly by the squamosal (fig. 17^). In Orthogeomys it
is remarkably broad throughout and is not constricted between the
orbits (fig. 17''), though the peculiar inflations at the anterior corners
7433— No. 8^—5

66

NORTH AMERICAN FAUNA.

[NO. 8.

of the orbits in 0. grandis produce the appearance of a constriction
behind them.

The juf/al varies in size and shape from the large and greatly
expanded i^late that forms the major part of the outer side of the zygo-
matic arch in Flalygeomys iylorldnus (pi. 13, tig. 1), to the rudimentary
splint or scale that adheres to the inferior side of the zygoma in Zygo-
geomys trichopiis, the arch being complete above without it (pi. 13,
fig. 24).

Fig. 17. -Types oi frontal.

1. Cratogeomys merriami. 3. Macrogeomys heterodus.

2. Heterogeomys torridus. 4. Orthogeomys scalops.

a?«, apex of alisphenoiil ; I, lachrymal; wixz, luasillary root of zygoma; n, nasal; iimx, ascending or
nasal branch of premaxilla; sq, squamosal.

The variation in the squamosal is hardly less extreme. Throughout the
genus, except in the most generalized forms, this bone exhibits a singu-
lar tendency toward expansion. In Geoniys proper the tendency is
restricted to a slight overlai)ping of the postero-lateral moiety of the
frontal and lower edge of the parietals. But in the genus Crato-
geomys its ambition in this direction is not satislied until the whole of
the posterior half of the cranium is covered. In Cratogeomys merriami
as the animal grows old the upper edges of the squamosals gradually
creep up over the parietals until the latter are comjiletely arched over
and concealed, the squamosals actually meeting above them along the
median line. lu doing this the squamosals cover the posterior part of
the frontal as well as the whole of the parietals and most of the inter-
parietal, and curve up posteriorly to take part in the formation of the
lambdoid crest for its entire length, thus roofing the brain with two

JAN., 1895.] THE SKULL. 67

distinct layers of bone, the npper of which on each side, consisting of
a single bone, overlaps in whole or in i^art five bones of the lower layer
(frontal, parietal, interparietal, supraoccipital, and alisphenoid). The
object of this unique arrangement is not only to furnish a brace to the
zygoma, to which the powerful masseter muscles are in large part
attached, but also to strengthen the vault of the cranium where the
huge temporal muscles take origin. The various steps in the develop-
ment of this extraordinary condition can be distinctly traced in the
series of skulls of dift'erent ages of Cratogeomys merriami collected by
Mr. ISelson in the Valley of Mexico. In Platygeomys another condition
prevails, the squamosal expansion being chiefly away from the median
line. On the inner side it overlaps the lower part of the parietals as
usual; it then extends outward in a broad shelf, carrying the squa-
mosal root of the zygoma far beyond its normal position, and spread-
ing outward and backward so as to completely roof over the post-
glenoid space, behind which .it pushes still further outward and over-
reaches the extreme end of the transversely elongated mastoid. In
Platygeomys gymnurus, tylorhinns, and planiceps the lateral expansion is
so excessive that the breadth of the cranium across the squamosals
posteriorly is actually greater than the breadth across the widely
sjjreading zygomatic arches (pi. 3).

The pterygoids vary surprisingly in size, form, and the extent to
which the inferior surface enters into the lateral walls of the post-
palatal notch, as already shown (pp. 52-53, and fig. 11). In Zygogeomys
they are long and slender and encircle the notch like a horseshoe, meet-
ing or nearly meeting in the median line behind the palate (pi. 14, fig. 1).
In most species of Geoniys, Cratogeomys, Pappogeomys, and Orthogeomys
they are more or less parallel plates forming the greater part of the walls
of the notch but not approximating anteriorly (pi. 14, figs. 7, 11, 13,
15). In Gconiys hursarins they are more posterior, and taper to nearly
a point behind, being Ungulate in shape (pi. 14, fig. 2). In Macrogeomys
they are short and broad and bend abruptly ujjward, capping the ends
of the short and broad palatines (pi. 14, fig. 3). In Eeterogeomys they
are small, and simply form the narrow ends of the elongated posterior
arms of the palatines (pi. 14, fig. 12).

The mandible is relatively small and light in Geomys. It is large
and massive in Cratogeomys, Platygeomys, and the remaining groups.
It is long and narrow, with short truncate angular processes, in Macro-
geomys dolichocephalus (pi. 10, fig. 7). It is broadly spreading, with
greatly elongated angular processes, in Platygeomys gymnurus (pi. 10,

tig. S).

The degree of development of the angular processes is correlated
with definite types of molariform teeth, and afi'ords a key to the domi-
nant movement of the jaw in mastication, the so-called 'grinding move-
ment' being very different in the species with and those without the
greatly elongated processes. Where these processes reach their highest

68 NORTH AMERICAN FAUNA. [no. 8.

development, as in Platygeomys (jymnurus (pi. 3 and pi. 12, fig. 8, and
text figs. 53 and 54) the posterior part of the masseter muscle, arising
from the jugal and squamosal arm of the zygoma, is correspondingly
large and effective; and since the direction of its fibers is nearly trans-
verse to the axis of the skull, it is evident that the resulting movement
of the jaw must be largely lateral. If the two parts of the masseter
contract simultaneously, the resulting motion of the jaw would be
oblique; if they contract independently, a to-and-fro movement would
alternate with a side wise movement.

In the species in which the lateral production of the angle of the
jaw is reduced to a minimum, as in Macrogeomys dolichocephalus (pi. 5
and pi. 12, tig. 7; and text figs. 51 and 52) the i^osterior part of the
masseter nmst be relatively unimportant, and the principal movement
must be to and fro. That this is really the case is shown by the greatly
restricted area of attachment for the jugal end of this part of the muscle
(fig. 49 Jo), and also by the character of the teeth. As would be
expected, the crowns of the molars are broader antero-posteriorly than
in the gymnurus group, and the tooth row on each side is bowed down-
ward—the crowns of the upper series as a whole being convex, the
lower concave, antero-posteriorly (fig. 46). Moreover, the obliquity of
the plane of contact of the upper and lower series is less in dolicho-
cephalus than in gymnurus (see figs. 52 and 54,/).*

* The types of molariform teeth coordinated with the two principal types of jaw
movement, and hence secondarily with the development of the angular processes,
are discussed at greater length under the head 'Mechanism and Dynamics of the cut-
tiiuj machine' (pp. 93-97).

CHAPTER Iir.

THE DEI^TAL ARMATURE.

THE TEETH.

The dental formula of the Geomyid(e is the same throughout the
family, as follows : i ., , c tj? pm j, m o X 2 = 20

All of the teeth of the Pocket Gophers are simple rootless * tubular
prisms, closed at the top aud open at the base. In life the lower part
is filled with a soft, pulp-like substance, supplied with blood vessels
which replenish the tooth from below, enabling it to grow as long as
the animal lives. The hardening of the pulp within the tooth forms

Fig. 18. — Outline of skull of Plalygeomys gynuivrns, showing teeth in situ.

the dentine and osteodentine; the enamel and cement are deposited on
the outside. In the adultt the crowns of the teeth are never compli-
cated by infoldings of the enamel ; the enamel never envelops the prism
continuously and never dips into the interior, but is always attached
to the outside in the form of vertical bands or plates like the staves on

** Althougli the teeth have no true roots, it is convenient to speak of the basal or
growing end as the root. The term is u.sed in this sense in the present paper.

tThe enamel caps of the young teeth, and changes in the enamel pattern due to
immaturity, are fully described under a se2)arate heading (p]). 83-8G).

69

70

NORTH AMERICAN FAUNA.

[NO. 8.

a barrel (pi. 10, fig, 12). The number ofeuamel plates on eacli tooth
varies from one to four. When the tooth is looked at from the side,
the alternating bands of enamel and cement are found to extend ver-
tically from base to crown ; and since the tooth is constantly worn
down from above and as constantly replenished by growth from below,
its original form is preserved and no sensible change in the enamel pat-
tern takes place.

THE INCISORS.

The incisors are long and heavy, with trenchant, chisel-like edges
(figs. 18 and 19). Their massiveness varies greatly in the different genera.
The upper incisor is shortly curved in a single plane, forming a little
more than a complete semicircle, and its root rests either in the ui)per
part of the interspace between tlie divaricating roots
of the premolar and first molar, as in Plafygeomys
(fig. 18), or directly above the root of the first molar,
as in some of the other genera. The lower incisor is
much longer, less shortly curved, and does not form
a complete semicircle. It passes backward beneath
and on the inner side of the molars, its own root rotat-
ing outward in a partial spiral like the beginning of
the twist in a ram's horn, and terminates in a thin
capsule of bone on the outer side of the condylar proc-
ess. The lower incisor is thus considerably longer
than the greatest length of the jaw, from which it
projects at both ends.

Both upper and lower incisors have their anterior
faces covered with a plate of enamel, the edges of
which are bent back over the sides of the tooth far
enough to hold it securely (fig. 20, a, h, and fig. 24)
so that it can withstand, without danger of loosening,
the great strain to wiiich it is subjected in cutting
hard roots.

On the inner side of the tooth the inflexed border of the enamel is
beveled (fig. 20, a) ; on the outer side it retains its normal thickness
(fig. 20, b). The inner edge of the tooth is squarely angular or nearly
so, while the outer edge is always broadly rounded (figs. 20, 21, 22). In
the lower incisor the front face of the tooth is always flat or nearly so
(fig. 24) ', in the up])er incisor it is flat in Macrogeomys and Jleferogeomys
(fig. 20), nearly flat or twice convex in Cratogeomys (fig, 21^ and ^),
Platygeomys (fig. 21^), and Fappogeomys (fig. 21,*); and thrice convex in
Geomys proper (fig. 22^ and ^) and Zygoyeomys (fig. 22^),

The enamel face of the upper incisor is invariably marked (except
in some species of Thomomys) by a conspicuous longitudinal groove or
furrow, resulting from an infolding of the enamel, A second and much
smaller groove is sometimes present also, always near the inner edge
of the tooth. The form and position of the grooves vary in the difl'er-

Fio. 19. — Incisora of
Platygeomys gymnurus
seen from behind, a up-
per; 6 lower.

JAN., 1895.]

THE INCISOR TEETH.

71

pf-^

ent species; there is also considerable range of individual variation.*
Five types of siilcation prevail, as follows :

Bisnlcafe se?(e.s;

Principal sulcus on oM/er side of median line Geomys

Principal sulcus on inner side of median line Zygogeomys

Unisulcate scries:

Sulcus median or slightly on inner side of median line; rather broadly

open Craiogeomys, I'laty geomys, Fappogeomys, Orthogeomys

Sulcus at junction of inner and middle thirds; usually rather narrow and

deep Heterogeomys, Macrogeomys

Sulcus close to inner side or absent Thomomys

In Gcomi/s proper the principal sulcus is decidedlj^ on the outer side,
and the small inner groove is about one-fourth or
one-tifth the distance from the inner edge to the prin-
cipal sulcus; it is nearer the inner border in the tuza
series (flg. 22^) than in tlie hnrsarius series (fig. 22^).
In Pappogeomys there is only a single groove (fig.
21''), and it is median or nearly so, as in Cratogeomys,
and very deep, Avitli the convexities on both sides
strongly rounded.

In Zygogeomys (flg. 22") the principal sulcus is
median or slightly on the inner side, and the fine
inner sulcus is ow the convexity of the enamel about
one-third the distance from the inner side to the me-
dian sulcus. It is not so near the inner side as in
Geomys proper. In tbe latter the inner convexity is
flatter and the small sulcus is on its inner side instead
of on the convexity itself.

In Heterogeomys and Macrogeomys (flg. 20) the
groove is always far on the inner side and some-
times wholly within tbe inner third. As a rule it is
deeper and more abrupt than in the other genera,
and the face of the tooth is flatter.

In Cratogeomys and Platygeomys (flg. 21) the groove, as seen by the

" The exact position of the principal sulcus varies not only in individuals of the
same sjiecies from the same place, but even on the two sides in the same skull. Thus
in Cratogeomys merriami and Platygeomys gymnurus of the unisulcate series it is
usually on the inner side of the median line, but several skulls of each species are
at hand in which it is median on one or both sides. Similarly, in Geomys bnrsarins
and tiiza of the bisulcate series, its distance from the outer side of the tooth is some-
times noticeably different on the two teeth. Its exact position therefore can not be
relied upon as a character in distinguishing species, though its approximate position
is important

Many of the unisulcate teeth show, when examined closely, a faint inner groove
in addition to the deep median furrow. The presence of this indistinct sulcus seems
to be purely fortuitous, occurring here and there irrespective of sex, age, or species,
sometimes on one side, sometimes on both, and is of no value whatever as a char-
acter. Another fortuitous variation is the occasional presence of a line bead in the
median sulcus. When present at all it is rarely symmetrical on the two teeth.

Fig. 20. — Transverse
section of upper in-
cisor in the unisulcate
species iii which the
sulcus is strongly on
the inner side. (1)
Macrogeomys dolicho-
cejjhftlus; (2) Hetero-
geomys hispidus; (3)
M. costaricensis; (4)
M. cherriei (showing
enamel face and single
sulcus), a inner end of
enamel plate; h outer
end of enamel plate.

72

NORTPI AMERICAN FAUNA.

unaided eye, ordinarily appears to be median j but when the tootli is
magnified it is nearly always found to lie sliglitly on the inner side.

Pig. 21. — Tran.sverse section of upper incisor 'Fig. 22. — Transverse section of upper incisor

in the unisulcate .species in wliicli the sul. in bisulcate series —

CU8 is median or nearly median — (1) Zygogeomystrichopus.

(1) Cratogeomys merriami. (2) Oeomys burcariug.

(2) Platygeomys gymnurus. ■ (3) Geomys tuza.

(3) Cratogeomys pcrotensU.

(4) Fap2>ogeomys bulleri.

It sometimes difters noticeably in position in the two incisors, and in
some specimens of C. merriami is further away from the middle than
usual.

In Orthogcomys the groove is on the inner side, but is usually so
widely open that its outer side reaches the median line.
In Thomomys the groove is close

to the inner edge of the tooth (fig.

23) or absent. It is usually j)res

ent, though sometimes very small

and shallow. In a few sijecies it is

deep and strongly marked, as in

T. monticola Allen.
The outline of the incisor in

cross section varies in the differ-
ent species. In some forms the anteroposterior diameter exceeds the
transverse; in others the transverse equals or exceeds the antero-
posterior. Usually the outer side of the tooth is an even curve from
the point where the infiexed border of the enamel stoj^s, to the posterior
convexity of the tooth, but this is not always the case. In the upper
incisor of Cratogeomys oreocctes, and the lower of C. merriami, the
outer side is emarginate, forming a distinct bevel immediately behind
thereflexed enamel edge (fig. 21, h).

Fig. 23. — Transierse
section of upper incisor
of Thomomys douglasi
showing shallow sulcus
close to inner side of
tooth.

Fig. 24.— Transverse
section of lower incisor
of Cratogeomys jnerri-
ami: byhexe] on outer
side.

THE PREMOLARS.

The premolars are double prisms, like a figure 8 in transverse section
(fig. 25 and pi. 16, figs. 8, 12, and 13). Their crowns are worn obliquely
to the axis of the tooth, hence the prisms are of unequal length ; the

JAN, 1895.] THE PREMOLARS. 73

posterior prism is longest in tlie upper premolar and the anterior in the
lower. In size the two prisms of the upper premolar are subequal or
the anterior is only slightly smaller than the posteriory in tbe lower, the
anterior is commonly considerably narrower and more elongated antero-
posteriorly. In form both prisms of the upper premolar and the pos-
terior of the lower are transversely elliptical like the molars; but the
anterior prism of the lower premolar is cylindrical or subcylindrical.
Its transverse section is more nearly circular in Zygogeomys trichopiis
and the Geomys bursarius series than in the others. In Macrogeomys
cherriei it is more elongated transversely than usual in the group.
The neck connecting the anterior and posterior i^risms is usually on or
near the median line of the tooth, but in the upj^er premolar of Hetero-
geomys hispidiis it is decidedly on the inner side.

The premolars are larger than the molars, and the lower premolar is
the largest of the molariform series (fig. 26). The upper premolar is
implanted very obliquely and invariably s?02)es strongly backward from
root to crown, the vertical plane of the root being far anterior to that
of the crown. The lower premolar is strongly curved ; it is always con-
cave anteriorly and convex posteriorly. It is implanted vertically or
nearly so, thougli its root curves forward. The upper premolar is decid-
edly longer than the lower in the genus Geomys (both in Geomys proper,
comprising the bursar ius-tuza series, and in
the Papiyogeomys biilleri series); the two are
subeqnal in all the other genera. The shaft
of the upper premolar may be either straight or
curved. When curved it may be convex
forward or concave forward. It is straight in
Geomys lutescens, but decidedly concave an- , ^'«- ^^-Crowns of upper and

, ' "^ lower premolars of Macrogeomys

teriorly in all the other species of Geomys doUehocephaius : a upper, 6
proper and in Fapimgeomys and Orfhogeomys ; ^*^''^®'"'
it is strongly or moderately convex anteriorly in Cratogeomys and
Macrogeomys^ and faintly convex or nearly straight in Reterogeomys,
Zygogeomys, and Flatygeomys. In the latter genera it is commonly
straight in the young and slightly curved in the adult.

The length of prism of the upper premolar in G. bursarius, turn, and
mobilensis is at least one-third greater than the total length of the tooth
row on the crowns (fig. 26^) ; in G. texensis it about equals the length
of the tooth row. Various intermediate conditions occur in the other
species. The length of the upper premolar with reference to the
molars affords two series: (1) in which the premolar and m' and m^
are of about the same length (comprising G. bursarius and most of the
species in the other genera, fig. 26 1 and '^); and (2) those in which the
premolar is decidedly longer than m' and ni^ {G. tiiza and mobilensis
and Pappogeomys bulleri, fig. 26^). The length of the upper and lower
premolars with reference to each other also affords two series : In the

74

NORTH AMERICAN FAUNA.

[NO. 8.

genus Geomi/s the lower is much shorter tliau the upper (fig. 26^); in
the otlier geuera {Crafogeomys, Heterogeomys^ and Zyyogeomys) the two
are subequal or the lower is slightly the longer (fig. 26^ and ^).

THE MOLARS.

The true molars, except the last upper one (m"), are simple single
tubular prisms, elliptical in transverse section. The last upper molar
is a single prism in some forms; a double prism in others. In both
upper and lower series the posterior molar is the shortest tooth (fig. 20).
In the lower series the teeth are successively shorter from jireinolar to
last molar. In the upper series the premolar may or may not be longer
than the first molar; the first and second molars may be subequal or
either may be. slightly longer than the other. As a rule throughout

Fig. 26. — Type.s of molariform teetli (seen in profile) ;

1. Heterogeomys hispidus.

2. Cratogeomys merriarni.

a upper series; b lower series.
3. Geonigs tuza.

the group, the first and second upf»er molars are as long or nearly as
long as the premolar. This is the case in Geomys bursarius; but in
other species of Geomys proper {fuza, breinceps, and texensh) and in the
genus Pappogeomys they are very much shorter. In Pappogeomys
bulleri and the Geomys tuza series the longest upper molar is only about
two-thirds the length of the premolar, and nr'' is only half as long as the
premolar.

In the lower jaw the molariform teeth are successively shorter from
before backwards, but diversity prevails in the relative lengths of the
several teeth comprising the series. Thus in Heterogeomys Jtisjndns m.^
is but little more than half the length oi pm; while in other species
it is more than three-fourths. The relative length of the individual
molars vaiies in the different species and is subject to considerable
individual variation also.

The last upper molar is always the largest of the true molars. Its
prism may be either single or double, or incompletely double; when
double it nearly equals the premolar in size of crown, but never in
length of shaft. It is invariably the shortest tooth of the upper series,

JAN., 1895.1

THE MOLAR TEETH. 75

aud ill some species is as sliort as the last lower molar. It always
curves backwards and the curvature is sometimes so great as to form
the arc of a small circle. When a double prism, the posterior prism is
always much narrower than the anterior. For purposes of classifica-
tion m' is by far the most important tooth in the skull, its size, shape,
form of crown, and enamel pattern furnishing characters of much value,
as will be seen later.

The last lower molar is ordinarily the shortest tooth in the skull, aud
is always curved— the concavity posterior. In addition to the curvature,
it is implanted obliquely, sloping- strongly backward from crown to
root, the vertical plane of the root being far behind that of the crown.
Its root is also rotated backward and inward, enabling it to lie flat
against the inner side of the incisor, which passes between the roots of
m2 aud m.T (fig. 41). Owing to the strong slope of the shaft of m:,, the
crown is always truncated very obliquely to the axis of the tooth (fig. 18).

The prisms of the intermediary molars in both jaws invariably curve
outward, so that their outer borders are concave and inner borders
convex. The curvature is stronger in the lower than in the upper
series, and strongest in m,, whose root stands further outward (away
from the median line) than any other in the series. The outer borders of
the prisms are shorter than the inner borders, hence the open root-ends
of the teeth always face obliquely outward. The anteroposterior
curvatures of the prisms of the intermediary molars above and below
take the same direction in each jaw, but vary in degree in the different
genera and sometimes in species of the same genus. All of the superior
molars curve backward from crown to root; the inferior intermediary
molars curve forward from crown to root. In the genus Geomys the
aiitero-posterior curvature of m' and m2 is so slight that their prisms may
be described as essentially flat (fig. 2G^). If any curvature is apparent,
it is backward in m' and forward in m2, in accordance with the rule. In
Zyyof/eomys and Heterogeomys the curvatures are slight; in Orthogeomys
they are marked, and in Macrogeomys, Cratogeomys, and Flafygeomys
they iire very strong, m' and m'^ curving strongly backward and mj and
m2 strongly forward (fig. 26^ and^).

In addition to the curves described, the molar prisms are always
more or less twisted on their axes. If the teeth were long enough these
twists would result in spiral curves.

The axes of the elliptical crowns of the intermediary molars are in a
general way transverse to the axis of the skull; but they rarely stand
out at right angles. As a rule they slope obliquely forward or obliquely
backward. When the crowns of the upper molars slope backward from
the median line the crowns of the lower molars are transverse or slope
forward, and rice versa. The axis of the crowns of m' and m^ normally
slopes backward in Geomys, Pappogeomys, and Cratogeomys; it is nor-
mally transverse or slopes forward in Platygeomys, Orthogeomys, Macro-
geomys, Heterogeomys, and Zygogeomys.

76

NORTH AMERICAN FAUNA.

[NO. 8.

Tig. 27.

-Types of ibrm of crown of last
ujiper molar (m^).

VARIATION IN FORM OF LAST UPPER MOLAR.

The form of the last upper molar affords excellent characters. In its
simplest type, as in the genus Geomys (comprising both the tuza series

and the texensk-hur sarins series) it is
a single prism and the shape of the
crown varies from suborbicular to sub-
triangular (figs. 27 ^ and 33). In Pap-
pogeomys (fig. 27 -) the form of the tooth
is similar except that there is a decided
emargination on the outer side, ante-
rior to the middle, behind which the
prism is abruptly narrower. This is
the first step in the formation of the
'heel' or posterior lobe, which is so
conspicuous in Orthogeomys^ Heteroge-
omys, and Macrogeomys (fig. 27^ and '^).
In the genus Cratogeomys the tooth
is partly converted into a double prism
by a vertical groove on the outer side
(fig. 27"*). This genus presents the widest latitude of individual varia-
tion known in the family, indicating that the tooth is in a transition
state and has not yet attained a condition of stable equilibrium.

It is much more variable in Cratogeomys than in Platygcomys. Taking
both genera together the crown presents all sorts of intermediate pat-
terns, from a form in which the posterior prism is hardly more differen-
tiated than in Pappogeomys hullerl, to forms having this prism produced
to such a degree that the superficial resemblance to Hetcrogeomys is
marked (tig. 35). But it lacks the stability of form and fixity of enamel
pattern characteristic of the members of the latter genus.

The variation is greater in the adult than the young, as would be
expected from the increased obliquity of the crown with reference to
the axis of tlie tooth in advanced age, and naturally is most marked in
the length and form of the heel. Sometimes in old age the crown is
worn so obliquely that the heel actually overhangs, acquiring an exag-
gerated length very different from its transverse section (as in fig. 28, d).

1. Geomys breviceps.

2. Pappogeomys hulleri.

3. Platygeomys gymnurus.

4. Cratogeomys estor.

5. Zygogeourys trichopiis.

6. Macrogeomys dolichocephalus.

7. Macrogeomys heterodus.

^Cs^

Fk!. 28. — Varlatious in crown pattern of ni'' in Cratogeomys fulvescens.

In Cratogeomys fulvescens (fig. 28) the variations in form and enamel
pattern of crown are pronounced, but most of them are easily reducible
to one or the other of two types: (1) An obcordate crown, deeply
notched l)etween the prisms on the outer side, with the axis of the pos-
terior loop or heel nearly transverse aiul the outer enamel plate reduced

JAN., 1895.]

THE LAST UPPER MOLAR. 77

to a small U-sbaped piece protectiug the sulcus (tig. 2S,a)', aud (2) a
more or less subtriaugiilar or eveu trefoil-shaped crown with the axis
of tiie posterior h)op very oblique (slopiug strongly backward as well
as outward), and the outer enamel plate more or less elongated (fig.
28, c, (1). In form the second is easily derived from the first by a slight
backward rotation of the transverse axis of the posterior loop. Regard-
ing the shape of the crown as more or less subtriangular, the apex of
the triangle is always toward the median line of the skull and the
notch or emarginatiou always on the outer (buccal) side. Cratogeomys
castanops (fig. 20) stands somewhat apart from the other species. The
double character of the prism is not well marked; the posterior part
of the crown is rather broadly rounded, the lateral enamel plates are
rather short, and the inner one is situated far back. Both tend to
disappear in extreme age — doubtless from atrophy of the enamel organ.

Fig. 29.— Variations in crovf n pattern of m^ in Cratorjcomys castanops.

In the genus Flatygeomys the crown is snbtriangular, narrow behind
the anterior prism, and the axis of the heel is normally anteroposterior,
as in Fappogeomys (fig. 27^).

hiMacrogeomys, Hcterogeomys, and Orthogeomys (fig. 34), the tooth is a
double prism, the anterior and posterior moieties of which are separated
by a groove or depression on each side — that on the outer side being
invariably the deeper, that on the inner side being in rare cases obso-
lete. The posterior prism is always narrower than the anterior (the
narrowing is chiefly on the onter side), aud its anteroposterior diameter
is usually greater. The crown as a whole is thus longer than broad,
and is composed of two parts or lobes: an anterior which is broader than
long (being transversely elliptical, like the other molars); and a narrow
posterior lobe or 'heel' which is commonly longer than broad, and
varies in form and proportions in the different species.

In Heterogeomys the grooves on the two sides are nearly oi:)posite, and
the anterior iirism is narrowly elliptical. In Orthogeomys und Macroge-
omys the sulcus on the inner side is commonly decidedly iiosterior to
the plane of the outer sulcus. In Macrogeomys the anterior prism is
broadly elliptical, and the posterior is elongated antero-posteriorly. In
Maci'ogeomys heterodus the posterior lobe or heel is very long and slopes
obliquely outward; the inner face of the tooth as a whole is unnsually
flat (fig. 27^).

In Zygogeomys the last upper molar is an imperfect double prism, the
depression on the inner side being slight, while that on the outer side
is much deeper. The crown as a whole is longer than broad, and the
posterior loop or heel ends in a broad lip-like extension not protected
by enamel and hence subject to change of shape by wear (see fig. 27^),

NORTH AMERICAN FAUNA.

[NO. 8.

Fig. 30.— Types of enamel
pattern of ujiper premolar.

(1) Cratofjeomys inerriami,-

(2) Heterogeomys hispidus; (a)
anterior enamel biind; (6) lat-
eral band ; (c) posterior band.

ARRANGEMENT OF THE ENAMEL.

After the enamel cap of tlie newly born young lias been ground down
far enough to expose the u^jper ends of the cement bands, the arrange-
ment of the enamel remains the same throughout the life of the indi-
vidual and affords excellent generic and in some cases specific char-
acters. The enamel never envelops the prism in a continuous sheet,
but is deposited in the form of vertical plates or bands which always
alternate with bands of cement. These bands are disposed in a definite
manner on each tooth of the series. In the under ja-w the number in
each tooth is the same throughout the group; in the upper jaw the
number varies in the several genera.

Premolars. — The ijermanent upper premolar has three enamel plates
(one anterior and one lateral on each side* ) in the genera Geomys
_ proper, Pappogeomys, Gratogeomys, and Platy-

geoniys — the posterior being altogether absent
(fig. 30'). In Zygogeomys, Heterogeomys, Macro-
geomys, and Orthogeomys the number is in-
creased to four by the addition of a posterior
plate, which, however,
never covers more than
half of the posterior face
of the posterior prism, and
is always restricted to the
inner or lingual side (fig. 30'^ c). In Orthogeomys
the posterior plate is sometimes obsolete. The per-
manent lower x^remolar always has four enamel
j)lates, the posterior being invariably present and
covering the whole hinder face of the tooth (fig. "25,
b, and fig. 32).

First and second upper molars. — In the first and
second upper molars, which are simple elliptical
prisms, the normal number of enamel plates is two,
one covering the anterior, the other the posterior
face of the tooth, with a narrow interval filled
with cement at each end between them (fig. 31^).
In many species, however, the posterior plate is
obsolete (fig, 31'). It is present and covers the
whole hinder side of the tooth in Geomys, Pappo-
geo-inys, Macrogeomys, Heterogeomys, and Orthogeo.
mys. It is present but restricted to the inner or
lingual half of the tooth in Zygogeomys (fig. 31^),
and is altogether absent in Gratogeomys (fig. 31'^) and Platy geomys.

*Iii both upper aud lower premolars the anterior enamel plate is convex forward;
the lateral are strongly bent, couforming to the sulcus between the prisms and
extending from the convexity of one to that of the other. The resulting shape in
transverse section is usually like that of the letter IJ, with the opening du'ected out-
ward aud the base resting on the median line of the tooth.

Pig. 31. — Types of enamel
pattern of upper niolari-
form series in the diti'ereut
groups :

1 . Geomys bursarius.

2. Ciatogeomys caistanoi)>s.

3. Zygogeomys trichojnis.

4. Macrogeomys cherriei.

5. Thomomys bidbivorus.

JAN., 1895.] THE ENAMEL PLATES. 79

Last upper molar. — Throughout the Geomydiw, except in Thomomys,
the last upper molar has three enamel plates — one anterior, one on the
inner side, and one on the outer side, with interspaces (cement bands) of
varyiug- breadth between (fig. 27). In Orthoycomys scalops the outer
plate is normally divided (tig. 62). The
anterior plate always covers the whole
front face of the tooth, and is the same
in all species ; the two others vary in
length and shape, and furnish excel-
lent characters. In Thomomys there
are but two plates, an anterior and a i
posterior (fig. 31^).

Lower molars. — Except in Thomomys,

the lower ]-0larS have each but a sin- ^'«- 32 -Crowns of lower molariform

series : (a) Geomys bur.mrius; (6) Thomomys
gle enamel plate; it completely covers hulUvorus. Except in Thomomys (D the
the posterior ftice of the tooth, the «^°a™«^^ pattern is the same tLroughout the
, ■ n 1 • 1 1, • 1 family (as in a).

anterior face and sides being covered

with cement (tig. 32, a). In Thomomys each lower molar has two enamel

plates, an anterior and a posterior (fig. 32, h).

PRINCIPAL DIVISIONS INDICATED BY THE ENAMEL PLATES.

The foregoing study of the enamel plates shows that all of the 37
species and subspecies herein described, and all the species of Th6md-
mys,may be arranged in five principal groups, according to the i)resence,
absence, or relations of the posterior enamel plate in tlie upper molari-
form series, as follows:

1. Posterior enamel plate absent in pm and present in m^ and m- Geomys,

Pappoocomys, Orthoueomya. *

2. Absent iu both 2)m and m' and m- Crutoyeomys, Flatygeomys.

3. Present on inner (lingual) side in both jj?)t and m' and m- Zyyogeomys.

i. Present on inner (lingual) side in jrm and complete in m' and m^ Hetero-

geomys, Macrogeomys, Orthogeomys. *
5. Present in pm and m', m-, and m* Thomomys.

NORMAL NUMBER OF ENAMEL PLATES — SUMMARY.

The number of enamel plates actually present in the different teeth
has been shown to vary from one to four. The number on each tooth
has been found constant in the lower series; inconstant in the upper
series. The lower premolar (which is a complete double prism) invari-
ably has four, and the lower molars one each, except in Thomomys iu
which they have two (fig. 32). The upper premolar (a complete double
prism) has four in some genera ; three in others. The upper interme-
diary or elliptical molars (m^ and m^) have two in some genera; one in

*Orthogeomy>i is losing the posterior enamel plate of the upper premolar. It is
present in 0, latifrons, but greatly reduced or altogether absent in nelsoni and
scalops,

80 NORTH AMERICAN FAUNA. [no. 8.

others. The last upper molar (an incomplete double prism) invariably
has two in Thomomys and three in all the other genera. These facts
indicate that the normal number of enamel ])lates in simple elliptical
prisms is two, and that one has been suppressed iu all of the elliptical
molars having only one (the lower molars in all except Thomomys aud
the first and second upper in Platyc/eomys aud Gratogeomys)^ and in the
^PP^r premolar when it has only three plates (as iu Platygeomys, Crato-
geomys, Pappogeomys^ and Geomys proper). This view is sujjported by
a study of the mechanics of the grinding process. (See pp. 90-97,
107-108).

VARIATIONS IN ENAMEL PLATES OF LAST UPPER MOLAR (m*).

Throughout the family, except in Thomomys, the last upper molar is
strengthened by three vertical plates or bands of enamel, alternating
with three interspaces filled with cement (figs. 33, 31). The anterior
of the three enamel plates is constant in form and relations; the two
others inconstant. The anterior invariably covers the whole front face
of the tooth and is convex forward (the convexity may be slight or
great). The others vary in position, shaj)e, and relative breadth. In a
single species, Orthogeomys scalops, the outer plate is normally divided
(tig. 62). In the simplest forms, in which the tooth is a subcylindric
or subtriangular prism, as in texensis, breviceps, and allied si)ecies (fig.
33), they are simple vertical bauds of enamel, subequal in size, one on

Fig. 33. — Variations in form of crown and enamel pattern of m^in restricted genus Geomys.

1, 2. Geomys tuza. G. Geomys personatus.

3. tuza /I arid anus. 7 — 10. texensis.

4. mobilensis. 11 — 13. breiriceps.

5. arenarius.

either side of the tooth posteriorly, separated from one another and
from the anterior enamel plate by similar vertical plates or bands of
cement. The genus Geomys proper presents no variations from this
type except in the relative breadth of the inner (lingual) and outer
(buccal) enamel bands. The inner is more constant than the outer and
is commonly somewhat broader.* Sometimes the two tend to define a
lip posteriorly (fig. 33^" and '^). Marked departures from this simple
type occur in those species iu which the last upper molar is a double
instead of a single prism ; and since various intermediate conditions in

*In G. tuza the outer plate is much narrower or shorter than the inner. Since the
teeth are commonly looked at endwise from above, the enamel pattern is ordinarily
seen in transverse section, and the three enamel i)lates ap]iear as narrow bands on
the periphery of the prism. Their breadth on the sides of the tooth i.s shown iu the
lenfjth of the baud as it appears on the crown. In describing the patteru. therefore,
it is convenient to use the term length instead of breadth to designate the rel?itive
width of the vertical enamel plates.

i

JAN., 1895.]

LAST UPPER MOLAR.

81

tlic evolution of the double ])iisiu are preseuted by living- species, so
the several stages in the adaptation of the lateral enamel plates to the
development of a posterior loop or heel are clearly shown. These
changes consist in a lengthening or shortening of the enamel plate (as
it appears on the crown of the tooth) and in the development of a bend
or flexure by virtue of which the enamel conforms to the curvature of
the anterior and posterior loops, resulting from the development of a
dee]) sulcus on one or both sides of the tooth in those species that have
a double prism. And since the sulcus on the outer side appears flrst
and is always deepest, it follows that the outer enamel plate is the one
most affected and shows the greatest range of variation (fig. 34).

Outer {buccal) enamel plate. — The first step in the formation of a dis-
tinct and permanent flexure may be seen in Pappogeomys bulleri (fig.
34,'), in which species the anterior end of the outer enamel plate bends

Fig. 34.— Forms of crowu aud enanifl pattt-rii of m^ iu the genera in whicli this tooth is a double
prism.

1. Pappogeomys bxtUeri. 8, 9. Orthogeomys nelsoni.

2. Platjigeomys gijnimirus. 8. Totontepec ; 9. Comaltepec.

3. Cratogeomyty estor. 10. Heterogeomyshispidus.

4. oreocetes. 11. torridus.

5. peregrinus. 12. Macrogeomys cherriei.

6. Zygogeomys trichopus. 13. contaricensis.

7. Orthogeomys latifrons. 14. dolichocephalus.

15. Alacrogeomys heterodus.

outward in front of the vertical sulcus that marks the outer side of
the tooth. A slightly more accentuated condition is found in Platy-
geomys gymnurus (fig.34,-). The extreme development of this flexure
is attained in the genera Heterogeomys (fig. 34, '" and ''), Macrogeomys
(fig. 34, '^, '^,i'^), and Orthogeomys (fig. 34, " and **), iu all of which the
bend is essentially a right angle — a result of the deepening of the
sulcus between the prisms. At the same time the posterior arm of the
enamel plate is considerably lengthened in order to protect the elon-
gated posterior lobe or heel to which it conforms. In Orthogeomys and
all the known species of Reterogeomys and Macrogeomys the posterior
limb is about double the length of the anterior; and except in M.
heterodus it actually reaches the hinder border of the tooth. In Ortho-
geomys scalops a very remarkable condition i)re vails; the outer enamel
plate is normally divided (fig. 62).
7433— Fo!! 8 0

82

NORTH AMERICAN FAUNA.

[NO. 8.

Ill riaiiigcomyH the outer enamel baiul is iiornially either straight or
bent outward at the extreme anterior end — not U shaped as in Crato-
geomys proper.

In the remaining groups a widely dilt'erent condition obtains: The
outer enamel plate is much reduced, and as a rule the two arms are sub-
equal. This type prevails in Cratogeomys proper and in Zygogeomys —
groups whose interrelations are distant and obscure. In Crato-
geomys the outer plate is normally ( ?) reduced to a mere angle or
U-shai^ed piece at the bottom of the sulcus that gives the outer side

of the tooth the semblance to a dou-
ble prism (tig. 35, •'" and ''), leaving a
wide unprotected interval (cement
band) on each side. It is variable,
however, and in some specimens the
posterior arm reaches nearly to the
end of the heel (fig. 35, ^). The dif-
ference maybe sexual; but owing
to the difficulty in determining the
sex in these animals, which difflculty
is greatly increased in the case of
the young, it is unsafe to place
much reliance ou the sex marks
accompanying the specimens. Still
there is reason for suspecting that
those specimens in which the outer plate is elongated x)osterioiiy are
females. The variation is much greater in some species than in others.
It is most extreme iu C, castanopi^ (fig 29), and least, so far as our
material goes, iu G. perotensis and estor. In advanced age it some-
times happens that the lateral enamel bands become abnormally short
on one or both sides and very rarely become divided in the middle.
Accidents of this sort are probably the result of shrinkage or atrophy
of the enamel organ.

Iu the genus Zygogeomys the outer angle is more open and the enamel
plate covers about half of the outer side of the tooth.

The outer enamel plate is slightly longer than the inner in Platy-
geomys^ and much longer in Heterogeomys, Orthogeomys, and Macro-
geomys (except in M. heterodus); it is subequal or shorter in all the
other known forms.

Inner [Ungual) enamel plate. — The inner plate is much less variable
than the outer, as jn-eviously stated. It is straight or slightly convex,
except in the few species that have a real sulcus on the inner side, con-
verting the tooth into a complete double prism. In these its anterior
part curves or bends outward. This condition is known in the three
genera, Heterogeomys., MaerogeomySj and Orthogeomys. In Heterogeomys
the outward curvature is slight (fig. 34, ^" and i^); in Macrogeomys cJoli-
eliocephalus and Orthogeomys latifrons it is strong (fig. 34, i* and'). In

Pig. 35.— Variations in form of crown and
enamel pattern of ni^ in Platygeomyg and in
Cratogeomys merriami.

1, 2. Flatygeomys gymnurus.

3. Flatygeomys tylorhinus.

4. Flatygeomys ftimosus.

5-8. Cratogeomys merriami (all from Ameca-
meca, Mexico).

JAN., 1895] THE UNWORN TEETH. 83

leiigtli and position the inner plate is much more variable: It reaches
the hinder end of the toDth in Geomys proper, Gratogeomys, Pap^wgeomys,
Flatygcomi/s* Zygogeoinys, and OrfJiogeomys; falls slightly short of the
end in Macrogeomys, and very considerably short in Ileterogeomys. In
Heterogeomys it barely covers half of the inner side of the tocth; in all
the other known species it covers nearly two-thirds or more than two-
thirds of the inner side. The condition in Ileterogeomys therefore is
clearly exceptional.

CHARACTERS OF THE UNWORN TEETH.

Specimens of pocket gophers young enough to show the deciduous
premolars and the unworn crowns of some of the molars are so exceed-
ingly rare that I have seen but four in the entire series of specimens
of this genus examined in the preparation of the present paper. Two
of these are Geomys bursarius from Elk River, Minn., collected by Ver-
non Bailey April 29, 1888, and May 14, 188G (Xos. 4909 and 2927, Mer-
riam collection); f he third is a young Geomys mohilensis from Milton,
Florida. The fourth is a juvenile specimen of Heterogeomys torridus
from Motzorongo, Mexico, collected by E. W. Nelson March 5, 1894
(No. 03043, IJ. S. :N^. M.). The unworn teeth are so much alike in the
two genera that they may be described together.

Incisors. — In both genera the grooves in the front face of the upper
incisors are very much deeper and larger than iu the adult, and the
convexities are much more strongly rounded. In the young of Geomys
hursarius the two grooves do not present the dispro[»ortion character-
istic of the adults, the small inner groove being relatively much deeper
and larger, though by no means so large as the median groove.

Deciduous premolars. — The crown of
the upper deciduous premolar is much
elongated and has an anterior prism in

addition to the double prism of the per- t,T>in--/y/. | |\| l)\\f-\""~^3
manent tooth (pi. TO, figs. 1 and 3). The
double prisms are united on tiie inner '^ r^

(lingual) side, forminga U-shaped grind- fig. se.-Lower moiariform teeth of a

iug surface (with the opening directed "•'^^'S' yo«ng Geomys bunanus, showing

. „ ]\-j? j.j?i-i j_ii ileciduovis and permauent premohir in

outward) m front of which, separated by ,i,„^ ,,,^ nn worn crown of m , which has

sulcus, is the small transversely elongated not yet reached tlie plane of the crowns

summit of the anterior prism. ^ The crown °' *''" "''^''' '"'"'•
of the lower deciduous premolar is likewise much elongated, and it is
irregularly and incompletely divided into three lobes (pi. 10, figs. 2 and
4&). Both upper and lower premolars have the anterior and posterior
roots far apart, and the permanent premolar may be seen between them
(fig. 30, and pi. 10, figs. 1-4, a).

*In Platygeomys fumosus the inner enamel band seems to be normally shorter than
the outer, and only half or less than half the length of the anterior band (fig. 35^).

tKT

84

NORTH AMEKICAN FAUNA.

[NO. 8.

Permanent premolars. — One of the upi^er deciduous premolars (pi.
16, fig. 1 h) has been removed from the baby skull of Heterogeomys tor-
ridiis, exposing the unworn crown of the permanent premolar (pi. 10,
fig. l.r). The permanent j)remolar also has been removed and figured
in several positions to show the form, size, and relations of its primi-
tive enamel cap (pi. 16, figs. 5, 6, and 7). For ready comparison, the
corresponding tooth in an adult of the same species has been figured
also (pi. 16, fig. 12). On reference to pi. 16 it will be seen not only
that the crown of the young premolar is completely enveloped with
enamel, but that the enamel cap reaches down over the shaft of the
double prism, covering nearly half of the tooth (figs. 5, 6, and 7) and
passing continuously into the four enamel bands that alone remain in
the adult (fig. 12*). The fact that the young of the various species as
usually obtained rarely show any trace of the enamel cap indicates
that the growth of the young teeth and grinding down of the crowns
progress with surprising rapidity. A very young Cratogeomys casta
7iops from Las Animas, Colo., collected by Dr. A. K. Fisher, has only a
remnant of the enamel cap left (i)l. 16, fig. 14).

The unworn crown of the upper premolar (pi. 16, figs. 1 a', 5, 6, 7) has
a single transverse crest on the anterior prism, an incompletely double
transverse crest on the posterior prisni, and an oblique ridge connect-
ing the two on the inner side. The single crest of the anterior prism
is notched or bifid at the apex, and has a small upright lobule at the

base of the notch on the inner side.
The double crest of the posterior prism
is open on the outer side, and the crest
as a whole is roughly and narrowly
U-shaped. The summit of the anterior
crest is bilobate; that of each arm of
the posterior crest is irregularly tri-
lobate or trituberculate. . fl
0 ••-! ; I ■" s - >* The enamel cap of the permanent

lower premolar is a complete double
prism, each moiety of which bears an
independent transversely elongated
crest (fig. 37). The summit of the an-
terior crest (fig. 37-), is trituberculate;
that of the posterior is incompletely
double, being split lengthwise into two
unequal parts, the posterior of which is
the shorter and more irregular. The trituberculate crest of the anterior
prism is bilaterally symmetrical. There are two large tubercles or lobes,
one on each side, and a smaller median one, which is much elongated
antero-posteriorly and is continuous ■oith the ridge connecting the
anterior and posterior prisms.

*In figs. 5, 6, 7, and 12 the cement bands are shaded, thus serving to hringout
the enamel more distinctly.

eo--

Fig. 37. — Right lowenin worn permanent
premolar of Heteroijeomi/s torridus: (1)
inner or lingual side ; (2) enamel cap from
above; c, cement bands; e, enamel; eo,
enamel organ.

JAN., 1895.

THE UNWORN MOLARS.

85

e --

c --

Molars. — In all of tlie young skulls under consideration the decidu-
ous premolar and the intermediary molars (m 1 and U) have been
used, and their enamel caps have been partly ground down, while the
permanent premolars and last molars have not yet sutiered attrition.
The premolar has been already described. The enamel cap of the last
hirer molar, which has not yet reached the plane of the crowns of the
other teeth (lig. 38 and pi. 16, tigs. 2, f7, 4, ^Z, and 9, d),
presents two complete transverse crests, each of
which has an undulating summit incompletely
divided into three lobes. The two crests are sepa-
rated by a deep furrow and show no tendency to
come together at any point. The enamel cap covers
a little more than half of the tooth (fig. 38, e). The
last upper molar (pi. 10, figs. 1, c and 3, c) has just
reached the level of the other teeth. Its unworn
crown in both genera presents a well-defined anterior
and a less distinctly defined po.sterior crest, sepa-
rated by an interspace which is bridged over by an
oblique enamel ridge on the inner side of the median
line. The anterior crest is incompletely trilobate.
The posterior crest is thickened and less symmetrical
than the anterior, and in Heterogeomys torridus (pi.
10, fig. 1, c) it is incompletely double, being partly
divided by a transverse excavation.

The crowns of the first and second upper molars present different
degrees of wear in the three young specimens at hand, and none of
them are young enough to show the transverse crests by which they
were undoubtedly crowned before the tops of their enamel caps were
ground down. The wearing, however, has not progressed so far as to
obliterate the double crowns characteristic of immaturity except in the
upper molars of one specimen of G. bursarius (I^o. 4009). In the other
skull of this species (jSTo. 2927) a transversely elongated loop of enamel
incompletely divides the grinding surface of m'\ indicating the former
presence of two transverse loops, as in the lower molars. In the lower
series the double crowns are well shown in both Geomys bursarius (pi.
16, fig. 4) and Heterogeomys torridus (pi. 16, fig. 2). In one skull of
Geomys bursarius (pi. 16, fig. 4) the second lower molar is only slightly
worn, and its crown presents two transverse loops separated by a
decided depression. In the other skull it is more worn, but still is incom-
pletely divided. The crown of the first lower molar in both skulls is
deeply notched on the inner side and slightly on the outer, showing that
when unworn it resembled the others.

i^ummary.— The summits of the unworn molariform teeth in Geomys
and allied genera are not only completely covered with enamel, but the
enamel cap is complicated by crests and tubercles. The permanent
premolar, which is a double i)rism, has a single transverse crest over

- eo

Fig. 38.— Right last
lower molar of very
yonng Heterogeomys tor-
ridus (from game speci,
men as fig. 37) ; inner or
lingual side, showing
unworn enamel cap, and
relations of enamel and
dentine lower down : c-
cement bands ; e, enamel ;
eo, enamel organ.

86 NORTH AMERICAN FAUNA. [no. 8.

the anterior prism and a partly double crest over the posterior. The
true molars are bilophodont, each carrying two transverse crests. In
the case of the last upper molar, the i)osterior crest is thickened and
somewhat irregular and may represent the coalescence of two crests.
It is joined to the anterior by an oblique ridge on the inner side. In
the premolar and last molar, above and below, the summit of each crest
is more or less distinctly divided into two or three lobes or tubercles.
There is every reason to believe that the crowns of the intermediary
molars (m ' and ') are similarly crested-tuberculate when in the unworn
condition, but in the specimens at hand their summits are worn down
too far to show it.

The crowns of the unworn teeth are bilophodont in all the lower
molars and in the first and second upper molars. The premolar and
last upper molar (m^) may be considered as imperfectly trilophodont,
the posterior prism in each instance being incompletely double.

The theory that permanently rootless teeth with flat griiiding crowns
are more primitive and less specialized than rooted teeth with tubercu-
late crowns receives a decided setback in the circumstance that the
young unworn molars in the Geomyiilw are provided with crested-tuber-
culate enamel caps, and that the adult teeth, though simple when con-
sidered singly as individual prisms, constitute, when taken collectively,
one of the most highly specialized grinding and cutting machines thus
fardiscovered.

CHANGES IN FORM AND ENAMEL PATTERN OF YOUNG TEETH
RESULTING FROM WEAR.

As already stated, the bilophodont crowns of the embryo and very
young molars are hardly ever seen, the wearing down of the primitive
enamel cap proceeding so rapidly that the youngest specimens ordina-
rily coming under the eye of the naturalist have flat grinding surfaces as
in the mature animal. During the reduction of the young crown four
different types of enamel i)attern, represeutingas many stages of wear,
succeed one another as follows:

First stage (before the crests are completely obliterated) : iico parallel
disconnected transverse loops.

Second stage (when the sulcus between the crests is reached): a
figure 8.

Third stage (after the sulcus is passed and before the tops of the
cement bands are reached) : a continuous ring or circle.

Fourth stage (after the tops of the cement bands are reached): the
pattern of the mature tooth, consisting of from one to three bands of
enamel alternating with the same number of bauds of cement, as already
explained in detail.

The first stage is of brief duration ; the second still more evanescent;
the third decidedly longer than the first and second together; the fourth
continues throughout the life of the animal.

A

JAN., 1895.] ENAMEL ORGAN AND OSTEODENTINE. 87

During the-eaiiy part of the fourth stage the form of the shaft of the
tooth changes, the double prism characteristic of -extreme youth giving
place to the single elliptical prism of the adult (except in the last upper
molar, which in some genera remains permanently double). It seems
remarkable that a tooth having a large double crown like the first and
second lower molars of the very young animal (pi. IG, figs. 2 and 4)
should be capable of changing its form to that of the single transverse
ellipse of the adult (pi. 10, fig. 17) in a very brief period and without
molting the tooth. That it does so is not open to question, and may
be demonstrated by making a section of the lower part of the young
touth. This has been done in the case of the second lower molar, as
shown in pi. 16, fig. 4, where 4c.x is a transverse section of the same
tooth from the lower fourth. 'The antero-posterior diameter of the
tooth decreases from above downward and the vertical groove on each
side becomes shallower and shallower and finally disappears. The
change in the shape of the crown takes place naturally by the rapid
wearing down of the grinding surface, which brings successively lower
parts to the top.

THE ENAMEL ORG^AN.

Throughout the group the enamel organ is situated at the base of the
teeth, as usual in rodents having prismatic molars. In the young tooth
the enamel organ is very much larger than in the adult, owing doubt-
less to the greater rapidity of growth in early life. Thus on referring
to pi. 10 (figs. 5, 0, and 7) it will be seen that the enamel organ occupies
about one- fifth of the length of the upper premolar in a very young
animal, while in the-correspouding tooth of an adult of the same species
(fig. 12) it occupies only about one-fifteenth of the length of the tooth. In
extreme age partial atropliy of the enamel organ sometimes takes place,
causing a shortening of the enamel on that side. In a few instances an
enamel plate has been found divided in the middle, due doubtless to
atrophy or injury of the enamel organ in the same vertical plane.

OSTEODENTINS.

A core of osteodentine traverses the central jiart of each tooth. In
the premolars and all of the molars except m^ it forms a large elliptical
shaft in the middle of each prism. In m^, whether single or double, the
osteodentine is a single core, conforming in shape to the shape of the
tooth. On all sides it passes into the true dentine, by which it is com-
pletely enveloped except at the free ends. At the lower end it passes
insensibly into the growing pulp. In other words, the osteodentine is
a central core consisting of the hardening pulp .and containing the
vessels by means of which the tooth is nourished. In the Geomyidct' it
forms a considerable part of the substance of the tooth, as usual in pris-
matic teeth growing from persistent pulps. In the genera Geomys and
Cratogeomys it is pale bufty or yellowish brown in color, and conse-

88 NORTH AMERICAN FAUNA. [no. 8.

quently not conspicuous. In the genera Heterogeomys and Zygogeomys
it is dark brown, in striking contrast to the white of the rest of the
tooth.

MECHANISM AND DYNAMICS OF THE CUTTING MACHINE AS A

VTHOLE.

The individual teeth have been described. It remains to consider
them as parts of a complex and highly specialized mechanism for cut-
ting and slicing the food, to describe the muscles that operate the
machine, to mention other structures concerned in the act of mastica-
tion, and to show how a bit of root or other hard vegetable tissue is
cut loose, sliced, and reduced to puli> ready to pass into the stomach.

The primary object of the dental armature is twofold: (1) To enable
the animal to bite or chisel off" pieces of the hard vegetable substances
on which it feeds, and (2) to reduce these pieces to a condition of
minute subdivision suitable to be tuined over to the stomach for
digestion. The. incisors serve the additional jHirpose of bars, axes, and
picks in helping the animal overcome the various obstacles encoun-
tered in driving its tunnels through different soils. When the front
teeth are used for this purpose, the resulting dirt and chips are kept
out of the mouth x^roper- by a furry partition, elsewhere described,
which divides the mouth- as a whole into two chambers.

MANNER. OF ATTACHMENT OF THE TEETH.

The- way the teeth are fastened in their sockets is in harmony with
the other remarkable adaptations of the grinding apparatus. The
attachment is effected by means of the periosteum of the alveolus,
which does not invest the teeth, but is firmly adherent to the cement
bands, leaving' the enamel faces free. Thus each tooth is suspended
by one or more vertical cushions, which extend all the way from root
to gum. This method of attachment not only relieves the tender pulp
at the base of the tooth from pressure, but gives to the cutting edge
or edges an elasticity that must be highly effective. In the case of
the incisors, the area of attachment is very extensive, comprising the
whole of the tooth below the gum except the enamel face. The lower
molars throughout the entire group, and the intermediary uj)per
molars in the genus Cratoffcomys, are attached in the same way on one
side only — the side opposite to the enamel ov cutting edge. In the
case of the upper premolars the principal attachment is along the
posterior face of the posterior prism, while a supplementary band on
eacli side of the anterior prism serves to keep the cutting edges always
in place. In those species in which the posterior prism of the upper
premolar develops an enamel band on its inner or lingual side, the
tooth is suspended by four cement bands. The lower premolar is
attached by four narrow lateral bands. The»last upi^er molar is inva-
riably held firmly in place by three cement bands, one on each side
anteriorly and one on or near the median line behind.

.TAN., 1895.1 THE CUTTING MACHINE. 89

DYNAMICS OF THE INCISORS.

The upper incisor has been shown to curve in the arc of a circle, to
cover a little more than a complete semicircle, and to lie in a single
plane (figs. 18 and 19). Its root is very long with relation to the length
of the muzzle, always overreaching the first upper molar. It is
implanted in such manner that its cutting edge is directed downward
and slightly backward. The hiwer incisor has been shown to curve
outward in an incomplete spiral, and to traverse the entire length of
the mandible — its root projecting on the outer side of the condylar proc-
ess, where it is incased in a thin capsule of bone. This small capsule
contains the pulp from which the tooth continually grows to replace
the wear at the other end. The extreme development of these teeth is
proportionate, of course, to the strain put u^jon them in chiseling hard
roots. The upper incisor is subjected to less strain than the lower, and
its principal function seems to be to anchor the cutting machine to the
substance operated on, while the greatly elongated lower incisor does
most of the work. The free end of the lower incisor slopes forward
and upward, its angle of implantation being difierent from that of the
upper. Tims, while the upper incisor remains stationary, its recurved
and usually divided tip enabling it to hold fast to the object to be cut,
the lower incisor plays rapidly back and forth like a steam drill, its
straight enamel edge doing the cutting.

The great length of the incisors within the alveolus is necessary in
order to counterbalance the length of the part that protrudes beyond
the jaws, and also to afford a large surface for attachment within the
alveolus so as to relieve the growing root from pressure. The way the
teeth are attached to the jaw by a long belt or cushion, which envelops
all but the enamel ftice, gives to the cutting edge an elasticity that
must be of great service, not only in increasing the efficiency of the act
of chiseling, but also in relieving the tooth from jar.

It remains to notice the interesting secondary modifications of the
skull and molariform teeth, by means of which the animal is enabled to
open the front part of the mouth wide enough to use the incisors to
advantage. The molariform teeth stand much higher out of the jaw
anteriorly than posteriorly, and their roots increase in length propor-
tionally (fig. 18). The premolars, both above and below, protrude
twice or more than twice as far as the last molars. Thus, when the
mouth is shut and the teeth pressed firmly together, the jaws are at
least twice as far apart at the anterior as at the posterior end of the
molar series. N"ow, the.distance from the crown of the premolar to the
putting edge of the upper incisor is two and one-half to three times the
length of the molariform series on the crowns, and the axis of the skull
is nearly parallel to the plane of the crowns of the molar teeth. Hence,
without any other help and with the mouth shut, the ends of the jaws
(where the incisors cut the gums) would be from five to six times fur-

90

NORTH AMERICAN FAUNA.

[NO. 8.

ther apart than at the plane of the posterior molars.* This arrange-
ment permits the necessary i^rotrusion of the jncisors, the cutting
edges of ^vhich, as a rule, reach the plane of the crowns of the molars
in the upper jaw and slightly pass this plane in the lower jaw. The
great advantage of this arrangement is most apparent during the act
of biting off hard roots, when a very slight opening of the mouth proper,
entailing only a slight separation of the molars, is sufiBcient (multiplied
along the length of the strongly divaricating jaws) to separate the
chisel ends of the incisors widely, enabling them to grasp objects of
comparatively large size.

DYNAMICS OF THE MOLARIFORM TEETH.

(a) Planner of implantation and curvatures.

The angle of implantation of the molar series as a whole in both
upper and lower jaws is peculiar. A transverse section of the skull
(fig. 39) shows that the roots of the upper molars are nearer the median
line than the crowns.! It follows that the upper tooth rows are strongly
divergent from root to crown (fig. 39, e). In the lower series the con-
Terse occurs, the tooth rows converging from root to crown (fig. 39,/).
The upper molars slope strongly and curve moderately outward from
root to crown, while the lower molars both slope and curve strongly
outward from crown to root.

The crowns of the opposing series do not meet in a horizontal plane,
^ but are obliquely truncated: the

upper series face obliquely doivn-
tvard and outward; the lower series
obliquely upward and inward (fig.
39). When the jaws are shut, lateral
movement in a horizontal plane is
impossible. If a circle is drawn
around the upper molars (fig. 40) it
is at once apparent that during the
lateral movement of the mandible
the crowns of the teeth move side-
ways in the arc of a circle, thus giv-
ing the utmost possible mechanical
advantage. The axis of rotation is
in or near the basicranial axis, and
the axis or arc of oscillation is short,
as in a pendulum. To enable the teeth to withstand the great pressure
to which they are thus subjected, they have developed very Ion >■ roots

*The actual condition is not exactly as here described. In the case of the lower
jaw the distance is decreased by the upward curvature of the anterior end of the jaw
and the shortening of the diastema. lu the upper jaw it is increased by the excava-
tion of the under side of the rostrum between the molars and incisors.

t The roots of the upper premolars are even nearer together than those of the molars ;
they are, in fact, almost in contact.

Fig. 39. — Transverse section of skull of
Platygeomys gi/mmirus, showing manner of
implantation and relations of niolariform teeth-
a, Frontal; &, zygoma; c, palate; d, mandible;
e, upper molar ; /, lower molar ; g, incisor.

JAN.. 1895.]

DYNAMICS OF THE MOLAR TEETH.

91

Fig. 40.— Upper and lower
molars of Flatygeomys gym-
7iurus in normal position,
showingangle of truncation
of crowns, necessitating
lateral movement in arc of
circle.

and a system of complex curvatures and oblique implantations, and
uie suspended in their sockets by vertical bands of periosteum, as
already described. When the jaws are shut, the molars on each side
curve outward so strongly that the distance be
tween them below (between roots of lower series)
is several times greater than above (between roots
of upper series). The result of this arrangement
is that the molar teeth, during the lateral move-
ment of the act of grinding the food, press upon
the opposing series not only iu such manner as to
secure the greatest mechanical advantage, but
also so as to produce the least jar, since the press-
ure in both directions is distributed over arcs of
circles. But this is not all, for if the tooth rows
are viewed from the side another remarkable
complex of curvatures appears (figs. 18 and 20).
It is now seen that in addition to the lateral curvatures there are
strongly developed antero-posterior curves and incomplete spiral
curves. In the upper series the premolar always slopes strongly
forward, and the molars curve backward from crown to root. In the
lower jaw the premolar and intermediary molars (mi and m-z) curve for-
ward from crown to root and the posterior molar backward. The
lower premolar is the largest and heaviest tooth of the molariform
series; it is strongly concave forward, convex backward, and is im-
planted nearly vertically. The last molar is the smallest tooth, and
both slopes and curves strongly backward from crown to root. The
end teeth of each series thus act as braces to support the tooth row as
a whole during the antero-posterior movement of the jaws in grinding,
and to keep the molars constantly 'keyed up,' so preventing any tend-
ency to spacing between the crowns.

In addition to the curvatures described,
the molariform teeth are usually more or
less twisted spirally on their vertical axes,
so that the two ends lie in different tan-
gential planes. Furthermore, the outer
(concave) edge is commonly shorter than
the inner (convex) edge.

The molariform teeth are so implanted
that the roots of each lateral series, above
and below, lie iu at least two antero-
posterior planes, the roots of the premolar and last molar in both jaws
l^eing nearer the median line of the skull than those of the intermedi-
ary molars. The discrepancy is most marked in the lower series, where
the posterior lower molars (mj and m.^) actually straddle the root of the
incisor (fig. 41). The roots of nii and m2 curve down outside (on the
buccal side) of the incisor, while that of m3 lies on its inner (lingual)
side. In order to do this the latter tooth (ms) not only curves strongly

-"Vn"!

Fig. 41.— Cross section of mandible
of Plati/geomys (jymnimis, showing how
roots of nij and ms straddle the incisor.

92

NORTH AMERICAN FAUNA.

[no. 8.

backward but is twisted on its own axis sufficiently to enable its root
to lie flatwise against the inner side of the incisor.

(&) Injiuence of the direction of the jmc movement on the molariform

teeth.

The direction of the dominant movement of the jaw exerts a marked
efl'ect upon the size, curvatures, proportions, and number of enamel
plates of the molariform teeth. This is well shown in comparing teeth
from skulls of the same size of Macrogeomys doUchocephalus and
Platygeomys gymnurus.

(1) Effect on the size and curvature of the prisms. — The length of the
molariform series on the crowns is approximately the same in both.
In M. doUchocephalus, in which the principal movement is antero-pos-
terior or nearly so, the premolars and last molars,
which form the end posts of the series, are very
much lengthened and enlarged, while the inter-
mediary molars are essentially the same size as
in P. gymnurus, in which animal the principal
movement is transverse or obliquely transverse.
The lower premolar of doUchocephalus (fig. 42, a)
contrasted with that of gymnurus (fig. 42, b) is
not only larger and longer, but its root curves
forward much more strongly, increasing its resist-
ing power as a brace. Throughout the group
this tooth (the lower premolar) acts as an immov-
able post or buttress against which the molars
l^ress during the to and fro grinding movement;
hence it is naturally largest in those species in
which the principal movement is antero-pos-
terior (see fig, 26).* The intermediary upper molars (m^ and m^) are
longer and less curved in doUchocephalus than in gymnurus; the inter-
mediary lower molars (mj and m2) are essentially equal in length in
the two forms and are equally curved, but the curvatures are differ-
ent: In M. doUchocephalus the upper half of the prism is nearly
straight, particularly in m^; the curvatures are more abrupt; the ante-
rior curve is much greater than in gymnurus, and the spiral twist is
more pronounced, the root end of the teeth rotating more strongly
inward. The posterior molar, both above and below, is much broader
and heavier in doUchocephalus than in gymnurus, and the upper one is
more strongly curved backward. The strong outward inclination of
the roots of the end teeth of the series tends to keep the molars per-
petually keyed up, preventing any spacing between the crowns. The
destructive effects of the to-and-fro movement of the powerful planing
machine are thus successfully offset.

*VVliat the lower premolar accomplishes by its massiveness and fixed position, the
upper premolar accomplishes by its length and angle of implantation.

Fig. 42.— Lower premolar
showing difference in size
and curvature according to
whether the domiuaut jaw
movement is to and fro or
sideways, a Macrogeomys
doUchocephalus; h Platy-
geomys gymnurui.

.iAN.,J895.] DYNAMICS OF THE MOLAR TEETH. 93

(2) Effect on the proportions of the prisms. — The breadth of the molar
prisms with respect to their anteroposterior diameter is materially
affected by the direction of the dominant movement of the jaw. This
is readily seen in the crowns which are much more elongated trans-
versely in those species in which the principal movement is obliquely
transverse (P. gymnurus and others) than in those in which it is chiefly
antero -posterior {M. dolichocephalus and others). In the former series
the transverse diameter of the crown (of upper molars) averages two
and one-half times the antero-posterior; in the latter, only two times.

(3) Eff^ect on the number and size of the enmnel plates. — Perhaps the
most conspicuous and important of the differences in the molariform
teeth, resulting from the direction of the dominant movement of the
jaw, is in the number of the enamel plates on the upper intermediary
molars. Two plates are invariably present in those forms in which
the dominant movement is antero-posterior (genera Geomys, Zyyogeomys,
Orthogeomys, Macrogeomys, and Heterogeomys) ; while only one is present
in those in which the movement is obliquely transverse (genera Platy-
geomys and Gratogeomys). In the latter case the enamel is restricted
to the front face of the tooth, the posterior plate being obsolete, and
the upper premolar resembles the molars in this respect, the pt»sterior
enamel plate being invariably absent.

ARRANGEMENT AND MODE OF OPERATION OF THE CUTTING BLADES.

The arrangement of the enamel plates and the direction of the dom-
inant movement of the jaw in mastication present two widely different
types in the animals under consideration. In one of these types the
principal movement is obliquely transverse; in the other it is antero-
posterior. They may be best considered separately.

{a) Dominant movement of jaw obliquely transverse. — When the upper
tooth row of Platygeomys gymnurus, or any other species in which the
dominant movement is obliiiuely transverse is
examined as a whole, it is found to be made up
of five flattened columns of dentine arranged
seriatim one in front of another, and each
faced in front with a vertical plate of enamel
which projects a short distance beyond the
crown (fig. 43 '). These five enamel plates are
strongly convex forward and their curvatures ^.^.o t ■. a ^

Fig. 43. — Longitudinal section

are essentially parallel (fig. 44 ^). An addi- of molariform teeth of piaty.
tional enamel plate covers the posterior face ^'""''"* i/i/ ''"'«'•«« (cUagram-

n .. . . .,, „ , , matic). (1) Upper; (2) lower.

ot the anterior pillar of the premolar and the

isthmus connecting the two parts of this tooth; and the two lateral
plates of the last upper molar may be considered as together forming
another cutting plate, making seven in all in the upper series. Turning
now to the opposing series — the lower molars — the opposite or
complementary condition prevails, a curved enamel plate covering

94

NORTH AMERICAN FAUNA.

[no. 8.

the posterior face of each of the live flattened columns of dentine
(figs. 43' and 44-). Two additional transverse plates complete the
armament of the lower premolar, making seven in all, as in the upper
series. It should be observed further that the concave sides of the five

regular enamel i)lates face bad-ward in the
upper series and forward in the lower series.
If now the two series are superimposed in
the position they naturally assume in the
mouth (tig. 45), and the lower series is moved
obliquely forward and outward in the direc-
tion it normally takes when drawn by the
masseter, the two sets of curved enamel
blades come together like the opposing
blades of seven pairs of shears working
almost simultaneously, with this difference
in favor of the teeth, that in addition to the anteroposterior closing
movement the curved blades slide over one another laterally, thus giv-
ing the greatest possible advantage in slicing the hard roots and other
unyielding substances on which the animals feed. The length of the
blades gives a long sweep, while the curvature* insures the passage of

Fig. 44.— Crowns of luolariforni
teeth of Flatygeomys gymniirns.
(1) Upper series; (2) lower series.

Fig. 45. — Superimposed molar series of Platygeomys gymnurus showing relations of enamel blades
(light outlines lower .series; dark, upper); a front end.

each particle of food against the cutting edges. The action is still
further favored by the obli(i[ue truncation of the molar crowns and the
peculiar method of suspension already described whereby the unyield-
ing enamel blades gain an elasticity which gives them a shearing motion
of the highest efficiency. The cutting is done during the obliquely for-
ward movement of the mandible; the complementary movement is sim-
l)ly one of recovery and has no effect on the food.

The forward movement is evidently complex and apparently consists
of three independent motions by which the mandible is shifted from
side to side in a zigzag manner, as follows: (1) The mandible is carried
obliquely forward and to one side until each of the enamel blades has
completed a shearing cut against one of the blades of the upper series;
(2) it is then carried obliquely forward in the opposite direction until
each blade completes another cut; (3) it then turns again and the
molar blades accomplish a third cut, leaving the upper and lower series

* The concave sides of the enamel blades move toward aud over one anotlier,
inclosing the food in a rapidly contracting loop, the opposite sides of which meet
and pass, leaving no chance for food to escape.

I

JAN., 1895.] DYNAMICS OF THE MOLAR TEETH. 95

nearly in the same vertical plane. The lower series has been carried
forward so that each tooth stands considerably in advance of the
corresponding- tooth of the upper series. A fourth movement, that of
recovery, brings the mandible back to the starting point. The limit of
the to-and-fro movement is nearly the same throughout the family
Geomyirl(e and is measured by the anteroposterior diameter of the
crown of the premolar, which it slightly exceeds. When the jaws are
at rest the front face of the lower premolar rests on or slightly behind
the corresponding face of the upper j)remolar. When the jaw is drawn
forward until the lower incisor strikes the posterior beveled face of the
upi)er incisor, the lower premolar stands free from and wholly anterior
to the upper. Hence, the thickness of the premolar is slightly less
than the distance covered in the to-aud-fro movement of the jaw. This
being the case, it is easy to ascertain the number of cuts made by the
enamel blades during each stroke of the jaw in mastication. By super-
imposing tracings of tlie upper and lower molar series (fig. 45) and
moving the latter oblicpiely forward and outward under the former
it appears that of the four cutting blades of the lower premolar the
first is unimportant, the secontl glides over two cutting edges of the
upper premolar during each stroke, the third and fourth cut against
three edges each, and the single blade of each of the three true molars
cuts over three enamel plates of the upper series (counting as one the
two lateral plates of the last upper molar against which they cut),
making seventeen cuts for each stroke of the jaw.

In a tame Geomys lufescens it was found (by actually counting the
contractions of the temporal muscle) that the mandible makes 200 com-
plete strokes a minute, which, at the rate of 17 cuts with each stroke,
is equivalent to 3,400 cuts by a single pair of blades. This is the num-
ber of cuts made by the blades of a single ramus; but since the blades
of both sides doubtless act simultaneously the number should be
doubled, making a total of G,800 cuts each minute!

The enamel plates are so spaced, by means of slight differences in
the anteroposterior diameters of the upper and lower molars, that
when the jaws are shut together and the movement of mastication
takes place, only one pair of cutting edges comes into bearing at a time.
The seven sets of blades, therefore, instead of cutting simultaneously,
follow one another in rapid succession, one pair just completing its
stroke as the next begins. By means of this delicate adjustment only
one-seventh the power is required that would be necessary if all oper-
ated together.

If, in the animals having the above described shearing movement of
the molars, a i)osterior enamel plate was present in the upper inter-
mediary molars, or an anterior plate in the lower molars, the possession
of such plates would obviously be a raechauical disadvantage, as they
would not only be of no use but would be actually in the way. Ilence,
in the evolution of this specialized type one plate has been gupx)ressedj

96

NORTH AMERICAN FAUNA.

[NO. 8.

Fig. 46. — Longitudinal section of
molariform teeth of Macrogeomys
dolichocephalus (diagranmiatic) .
(1) Upper series; (2) lower.

and the fact should be emphasized that the loss of a useless enamel
plate is as clearly a sign of specialization as the development of an addi-
tional plate where needed. In the less specialized genus Thomomys
both plates are always present (tig. 32, h).

{h) Dominant movement of jaw anteroposterior.— In the remaining
groups the movement of the jaw is chieiiy anteroposterior, the crowns

of the teeth are more broadly elliptical, and
enamel plates are i^resent on both sides of the
upper molars (figs. 46 and 47). In some genera
Ihe posterior plate, which is always thinner
than the anterior, covers the whole hinder
face of the tooth; in others it is restricted
to the inner side, according to the exact axis
of jaw movement. Whenever the ellipse is
broad, and is so directed with reference to the
enamel plates of the adjacent teeth that it
presents a free edge toward the food that
is being ground, this edge is invariably protected by a plate and cutting
edge of enamel. Conspicuous illustrations of this law may be seen in

the upper premolar of Zygogeomys, Macrogeo-
mys, and Heterogeomys, and in the upper inter-
mediary molars of Zygogeomys, in all of which
the posterior enamel plate is restricted to the
lingual side — the side impinged upon by the
food. On the other hand, non-cutting edges
protected by the enamel plates of adjacent
teeth are better off without enamel of their
own, because such enamel, if present, would
not only be of no use, but would be actually in the way, as already
explained.

By superimposing tracings of the upper and lower molar series of
Macrogeomys dolichocephalus (fig. 48) and moving the lower backward
and forward under the up^jer as nearly as possible in the way they are
moved by the living animal, it is found that the cutting blades make
nineteen cuts during each forward stroke of the jaw, as follows: The

Tig. 47.— Crowns of molariform
teeth of Macrogeomys dolicho-
cephalus. (1) Upper; (2) lower.

Fig. 48.— Superimposed molar series of Macrogeomys dolichocephalus .showing relations of enamel
blades. Light outlines, lower series; dark, upper, a front end.

anterior plate of the lower premolar does not cut at all, or, if it cuts its
action is so limited as to be of no particular consequence; the posterior
plate of the anterior prism makes two cuts ; the anterior plate of the pos-
terior prism, three cuts ; the posterior plate of the premolar and that of

JAN., 1805.] DYNAMICS OF THE MOLAR TEETH. 97

the first molar make four cuts each; the second and third mohirs, three
cuts each (counting the two lateral plates of the, upper molar, against
which ma acts, as if they were a single plate), making nineteen in all.
During the return movement fourteen cuts are made, as follows: The
second and third transverse plates of the i)remolar make two cuts each ;
tbe fourth, three; the first molar, three; the second and third molars, two
each. The backward stroke is evidently less powerful and less effective
than the forward stroke.

Since the teeth on both sides of the mandible cut simultaneously, the
total number of cuts during each complete stroke will be double the
number above mentioned, or 38 for the forward stroke and 28 for the
backward stroke. Assuming that the luimber of complete strokes each
minute is the same as in Geomys luteficens, namely, 200, the total number
of cuts made each minute on the forward stroke would be 7,600, and on
the backward stroke 5,(300, making a grand total of 13,200 cuts each
minute while the jaws are in active operation !

Stroke of the jaw. — There being no postglenoid process, the backwaid
movement of the jaw is not interrupted until the condyle strikes the
auditory bulla at the base of the tube of the meatus. When the con-
dyle rests in this position and the molar series are in api)Osition, the
front faces of the premolars above and below are in line. The forward
movement of the jaw is stopped by the incisors and reaches its limit
when the front face of the low^er incisor strikes against the posterior
face of the beveled edge of the ujiper incisor. When this happens
the upper premolar usually rests on the back part of the first lower
molar.

From the foregoing account it must be clear tliat the molars, which,
considered as individual teeth, are simple elliptical tubes, lacking the
complicated enamel patterns of the beaver, porcupine, and many other
rodents, are so constructed that collectively they form one of the most
powerful and highly specialized cutting and slicing machines known.
The way the narrowly elliptical crowns are placed side by side flatwise,
the hard projecting enamel blades alternating with surfaces of soft den-
tine, results in the production of a cutting and rasping apparai us e(iual
if not superior to that possessed by those rodents and ungulates tliat
have complicated enamel folds within the substance of the teeth. The
obli(puty of the crowns, whereby the upper and lower series are brought
together in the arc of a circle, gives them remarkable power under
the transverse movement of the jaws, while the way the teeth are stis-
pended on vertical cushions, together witii the angle of imi)lantati()n
and the double curvatures of their prisms, enables them to withstand
the great strain to which they are subjected Mithout danger of dis
])laeement and without injury to the tender pulps at their bases.

The secondary modifications of the skull resulting from the action of
the muscles operating this wonderfully effective machinery are dis-
cussed elsewhere (pp. 104-107),
7133— Is^o, 8 7

98 NORTH AMERICAN P^AUNA. [no. 8.

TREATMENT AND COURSE OF THE FOOD.

Tho cireniiistance that all the iiieinbers of the (icomyhlwYwo, mider-
grouiid has an important l)earin.t;- on tliekind of food habitually eaten,
and is thus the remote cause of the special adaptations of the dental
armature, and of the secondary cranial moditications necessitated
thereby. The animals sometimes come to the surface and cut the stems
and leaves of plants, which they draw into their subterrannean tunnels,
but in the main the choice of food is restricted to such parts of plants
as nuiy be found within the ground. The food therefore consists chietiy
of tubers and roots, including the hard roots of trees and shrubs, the
tough rootstalks of the mescal or agave, and tlie like. In dealing with
these unyielding substances the aniaial gains one decided advantage —
the roots on which it feeds are held firmly in place by the earth while
pieces are chiseled off by the broad, trenchant cutting edges of the
powerful incisors. In the case of certain relatively soft substances,
such as j)otatoes, the lower incisors are sometimes used alone, both as
a i)ry to dislodge pieces and as a scraper to scrape off thin slices, but
as a rule both upper and lower incisors operate together. The prin-
cipal function of the upper incisors seems to be to transfix the tuber and
oppose the action of the lower while the latter do most of the work,
moving rapidly backward aiul forward (and at the same time upward),
until a piece of food is cut loose or sufficiently undermined so that it
may be torn loose by a backward movement of the head while the teeth
are held firndy together. The bit of food thus dislodged is eitlier
reduced in size by trimming — during which operation it is held between
the large Ibrefeet, the long claws turned inward toward one another — or
is passed directly into the mouth or cheek pouches. The mouth i)roper,
it should be remembered, is separated from the incisors by a furry parti-
tion which is directly in front of the molars. This diaithragm-like par-
tition is of great service to the animal, keeping dirt and chips out of
the mouth. When the food reaches the mouth ])roper the tongue and
lips keep it between the teeth, where it undergoes the treatment com-
monly described as grinding. But in the highly specialized forms of
the Geonn/idcv no real grinding occurs — the whole process is one of cut-
ting or slicing. The arrangement of the enamt 1 plates that form the
blades of the cutting machine has been already described in detail.
In those species in which the principal movement of the jaw is antero
posterior the mechanism is essentially a phoiiu;/ machine, wliile in
those in wliich the dominant movement is obliquely transverse it is a
shc(tri)u/ or HJicing machine. In either case the tough vegetable fibers
composing the food are quickly reduced to a pulp, which is promptly
l)assed on to the stomach for digestion.

MUSCLES THAT OPERATE THE CUTTING MACHINE.

The principal muscles concerned in the movements of the jaw are (1)
temporal, (2) masseter, (3) internal pterygoid, (4) external pterygoid,

JAN. 1895.1 MUSCLES THAT MOVE THE JAWS. 99

(o) dig'astric, ami (6) trausverse mandibular. Cf these, by far the most
important single muscle is the masseter.

The temporal muscle occupies the whole of the upper surface of the
cranium behind the orbits, covering the parietal, squamosal, and pos-
terior part of the frontal as far forward as the postorbital prominence.
It arises from the flat upper suifacesof these bones and from the larab-
doid and sagittal crests. The muscle is indistinctly divided into two
parts — a superficial and a deep — which are m)t well detiued in their
origin. The fibers of the muscle as a whole converge anteriorly; those
of the superficial part are inserted into the apex, posterior edge, and
inner side of the coronoid process; those of the deep part play over the
trochlear groove and at the margin of the orbit drop vertically down-
ward and are inserted by a dense aponeurosis on the anterior edge of
the basal half of the coronoid ramus from the plane of the molar crowns
upward to a point slightly above the plane of the coronoid notch; pos-
teriorly the muscle remains fleshy and c(3vers the inner side of the cor
onoi<l ramus wliere its insertion extends downward to the bottom of
the deep pit between the ramus and the posterior molar. The function
of the temporal muscle is to shut the mouth, and in some species to
draw the mandible slightly backward. Operating in connection with
the digastric, it performs the backward stroke of the to and fro move-
ment of the jaw in the (Jolichocephidic series, the masseter producing
the forward stroke.

The masseter is a large complex muscle and is by far the most impor-
tant of the muscles concerned in the act of mastication. It is incom-
pletely divided into three parts, which, from their principal sources of
origin, may be described as the rostral or suiierficial, maxillary, and
zygomatic parts.

(1) The rostral or superficial part arises by a long an<I dense aponeuro-
sis from the outer side of the rostrum on tlie line of the preniaxillo-
maxillary suture, its npper border being immediately in front of the
infraorbital foramen. It passes thence obliquely downward and back-
ward, developing muscular fibers and spreading- out posteriorly into a
flat muscular band which is inserted upon the inferior crest of the
masseteric fossa and the inferior surface of the mandible from the
digastric crest posteriorly to the base of the angular process, its inser-
tion being wholly fleshy. It is the most i>owerful muscle in drawing the
jaw straight forward, and is aided in the dolichocephalie species by the
zygomatic branch of the masseter.

(2) Tlie main body of the masseter arises from the side of the anterior
part of the maxilla and adjacent parts of the maxillary root of the
zygoma. Anteriorly it slightly overlaps the posterior part of the pre-
maxilla immediately below the top of the rostrum, where it forms a
distinct crest continuous with the anterior edge of the maxillary root
of the zygoma. The principal origin covers the whole of the anterior
face of the vertically expanded zygomatic process of the maxilla, and

100 NORTH AMERICAN FAUNA. [no. 8.

in additiou a thin supplenieiitary sheet takes origin from the posl-erior
face of the same bony plate (withni the orbital chamber). J'osteriorly
its origin is limited on the outer side by a thick aponeurosis, which is
firmly attached to the inferior surface of the antero external angle of
the zygoma. The part within the orbit follows the inner face of the
korizontal part of the zygoma all the way back to the glenoid ligament,
to which its posterior libers are attached. This part of the muscle is
insei ted on the outer side of the neck of the condylar ramus just above
the incisor capsule.

(3) The ziif/oniatic part of the masseter arises from the outer side of
the horizontal partof the zygoma, its origin embracing the outer sur-
face of the scjuamosal root of the zygoma and the outer side of thejugal
below the oblique crest which marks the limits of its insertion above
and in front. It arises also from the aponeurotic septum which sepa-
rates it from the mam body of the muscle. It is inserted ujion the angu-
lar process of the mandible, its insertion covering the upper surface of
this process irom the incisor capsule outwardlj^ to and over the head of
the process, and also the under surface of the process to its very base,
where its insertion becomes continuous with that of the main body of the
muscle. Its function in Geomi/.s i)ioper and in all the (loUchoccphalic
si)e('ies is to draw the jaw forward. In the phitucepludic species its
insertion is carried so far outward by the great elongation of the angu-
lar process that it serves to move the jaw sideways, in which act it is
aided by the pterygoid muscles.

'J he internal pferyf/oid mvscle arises from the pterygoid fossa of the
skull, which it completely tills. Passing directly outward and slightly
downward, it is inserted into the pterygoid fossa of the jaw, where its
line of attachment has developed a strong crest along the posterior edge
of the angular process. Its function in Geomi/s proper and in all of the
dolichocephalic species seems to be to bring the posterior end of the molar
series firmly together when the jaw is shut. In the platycephalic species
it aids the masseter in moving the jaw sideways.

The external pterygoid arises from the alispheuoid bone on the outer
side of the root o^ the last upper molar and is inserted into tlie inner side
of the neck of the condyle. Its function is evidently mainly the same as
that of the internal pterygoid, though in addition it tends to move the
mandible slightly forward.

The digastric arises from the paroccipital process and adjacent parts
of the mastoid and audita! bulhie, and is inserted on the digastric crest,
Avhich projects backward from the hinder part of the symphysis of the J
mandible. It is largely developed, its function being not merely to
open the mouth, but, operating with the temporal, to draw the jaw
strongly backAvard in the to and fro movement of mastication in the
dolichocephalic series. Its action is very direct and powerful.

The transverse mandibular muscle connects the two halves of the lower
jaw immediately behind the symphysis, where, in many species, there is

JAN., 1895.] MUSCLES OF THE CHEEK POUCHES. 101

;i distiuct fossil for its lodgment. It must fulfill an important function
in regulating tlie adjustment of the tootii rows during mastication.

MUSCLES OF THE CHEEK POUCHES.

1 have not dissected the muscles of the cheek pouches, but they have
been described by Dr. C. E. McChesney* and Prof. H. L. Osborn.t
Dr. McChesney states that the aperture of the pouch is surrounded by
a narrow delicate constrictor muscle, and that the long pouch itself,
which extends back to the shoulder, is enveloped by a contractor mus-
cle which seems to be a modified part of the platysma myoides. This
muscle consists of two parts : (1) a retractor part, reaching from the
extreme posterior end of the pouch backward over the muscles of the
back and ending in a broad thin tendon which blends with the tendons
of the superficial dorsal muscles, to be inserted into the spines of the
three last lumbar vertebne; (2) an anterior part which envelops the
pouch proper. This latter is in turn subdivided into two parts — exter-
nal and internal. The former covers the upper or outer T)ortion of the
[louch and is inserted into the maxillary bone ( probably j^rcmaxillary).
The latter covers the inner and under sides of the pouch and is attached
to the mandible, though the uppermost fibers join those of the former
division, to be inserted on the upper jaw. Dr. McChesney states that
the lower and inner surface of the muscle is thickest, the outer surface
l^eing thin and of little power.

Prof Osborn describes the muscles of the pouch as follows: "There
are three distinct sets of muscles; these are, first, a circular muscle that
runs around the margin of the pocket in its outer bounding fold. This
by its contraction would seem to purse the opening of the pocket. The
second set of muscles are those that I will call the protractors of the
pockets. These are two in number on each side. They are spread out
in the skin of both the inner and outer posterior portions of the pockets,
and their fibers converge forward to finally form somewhat definite
bands. The outer of these is attached in the skin at the origin of tlie
fold on the upper jaw. The other is attached to the lower attachment
of the fold at the lower jaw. These two muscles thus surround the
po(;ket, and their contraction pulls its recess forward to the opening of
the vestibule. The third set of nuiscles are the retractors of the pocket.
These arise funnel-wise from surface of the pocket, both on its inner
and outer aspects, and they run backward and dorsally ])arallel to the
fibers of the latissimus dorsi and totally free from the skin. They form
a band three or four inches long and nearly an inch wide, and are
filially inserted in the tendinous aponeurosis that covers the inser-
tion of the latissimus dorsi and is attached to the neural spines of the
anterior lumbar vertebme. These by their action retract the pockets."

"Bull, U. S. Geol. aud Geog. Survey Terr., iv, No. 1, Feb., 1878,214-215.
t Science, xxiii, Feb. 23, 1894, 102-103.

102 NORTH AMERICAN FAUNA. [no. 8.

PRINCIPAL MUSCLES CONNECTING THE HEAD WITH THE NECK.

The sterno-mastoid muscle arises by a teudiuous aponeurosis from the
manubrium of the sternum and is inserted into the mastoid process of
the squamosal immediately behind tlie auditory meatus.

The clei do-ma staid arises from the middle part of the clavicle and is
inserted on the upper or dorsal aspect of the mastoid process of the
squamosal immediately over or above the insertion of the sterno-mas-
toid. Its libers are but little separated from those of the trapezius.

The trapezius muscle arises from the veutr.il surface of the outer
third of the clavicle and the adjacent acromial process of the scapula
and the spine of the scapula for its entire length : near the median line
its fibers seem to be continuous with those of the median part of the
hitissi)iiiis dorsi. It is inserted on the lambdoid crest for its entire
length, its outer edges being continuous with the insertion of tliecleido-
mastoid.

The rhomboidcus lies immediately below the trapezius. It is much
less extensive than the latter, but considerably thicker. It arises from
the superior face of the spine of the scapula and the adjacent anterior
part of the vertebral border of the sca])nla, and is inserted into the pos-
terior face of the lambdoid crest immediately beneath the insertion of
the trai)ezius.

ANALYSIS OF .JAW MOVEMENTS.

Turning now from the consideration of the individual muscles to the
study of tlte origin of the complex movements of the jaw in chiseling
and slicing the food, even greater dififlculties are encountered. The
following attempt, therefore, is subject to correction. fl

(1) The act of chiseling. — From what has been said it appears that
the act of chiseling" is performed in essentially the same way in both the
platyecphalic and dolichocephalic members of the group, and that it is
due to the joint action of the masseter and temporal muscles, the former
being more effective than the latter.

The thin enamel edge of the upper incisors is used chiefly as an
anchor to fasten the cutting machine firmly to the object operated upon,
while the lower jaw plays back and forth like a drill in accomplishing
the work. The exserted part of the upper incisors, therefore, is curved
downward and inward, and the edge, which is very thin and sharp, is
broken by one or more grooves, which enable it to penetrate hard sub-
stances more easily than if it were straight. The face of the lower
incisor slopes strongly forward as well as upward and the axis of its
movement in cutting must be obliquely forward and upward. The
principal muscle concerned in chiseling is the masseter, which is aided
by the temporal, and in some cases also probably by the pterygoids.
The way the posterior part of the ramus of the mandible curves upward

•JAN., 1895.] ANALYSIS OF JAW MOVEMENTS. 103

ill tlie arc of a circle lias a biglily important bearing- on the efficacy of
the action of the niasseter, and lias doubtless been molded into its
l)iesent sliape by this all-important muscle. The rostral part of the
niasseter is nearly horizontal; from its aponeurotic origin on the sides
of the rostrum it spreads out posteriorly and is inserted broadly over
the posterior curvature of the upturned ramus of the mandible, its
action being- to draw the mandible as a whole directly forward. The
main body of the muscle is uearly vertical, but slopes slightly back-
ward from its maxillary origin to its insertion on the outer side of the
mandible; in contracting draws the jaw slightly forward and power-
fully upward. In those species in which the zygomatic part of the
niasseter is nearly vertical instead of transverse this part of the muscle
aids the rest in moving the jaw forward and upward. The masseter is
aided still further by the temporal muscle, which, using the condyle as
a fulcrum, moves the lower incisors upward.

(-') The act of sUcin;/. — The act of slicing the food is performed in dif-
lercnt waysin the two series of animals, being chietly ato and-fro move-
iiK'iit in the (loHchocephalic species and a transversely oblique rotary
movement in the liJaiyeephalic species. In the dolichocephalic species
botli the forward and backward movements are important, while in the
pi at ycephaltG i^liecies the backward movement is merely one of recovery.

In the (lolichocepJialic series, therefore, the forward movement produced
by the masseter reipiires a powerful counter movement in bringing the
•jaw bick. This is supplied, apparently, by the joint action of the
digastric and the deep part of the temporal. The latter holds the teeth
firmly together and draws the jaw slightly backward, while the digastric,
contracting at the same time, pulls the jaw powerfully backward, the
superficial part of the temporal, which is inserted ou the coronoid proc-
ess, preventing it from opening the mouth.

In t\\& platycephalic series, as already stated, the principal movemeut
is obliquely transverse, the jaw being drawn outward and forward.
The muscles proilucing this action are the zygomatic part of the mas-
seter and the pterygoids. It is probable that they are largely aided by
the deep iiorticm of the temi)oral, which is inserted into the pit on the
outer side of the posterior molars. The fibers of this ]>art of tlie tem-
poral muscle being- vertical, bring the teeth firmly together and draw the
under jaw slightly outward, which movement, in connection with the
angle of truncation of the crowns of the teeth, must result in the trans-
verse rotary motion.

The mouth is opened by means of the digastric muscle, which is
beautifully adapted to this end, its origin taking hold of the posterior
part of the cranium on each side of occipital condyles, while its iuser-
tioii is carried forward all the way to the symphysis of the jaw. The
digastric does not ai)pear to be assisted by any other muscle iu i)erform-
iug its function of opening the mouth.

104

NORTH AMERICAN FAUNA.

[NO. 8.

IKFLVENCE OF THE MA8SETER MUSCLE IN MOLDING THE HKULL
AND MODIFYING THE TEETH. *

Throughout the Geomyidw the masseter muscle has profoundly modi-
fied the form of the skull and the character of the teeth, and is largely
responsible for tlie extraordinary cranial peculiarities that distinguish
the several genera. Perhaps it would be better to say that slight dif-
ferences in the direction of the principal movement of the jaw in grind
ing the food, which have proved an advantage to the animal, have by
natufal selection developed certain fibers or parts of the muscle at the
expense of other parts, and that the differences thus origiaated have
been i)erpetuated and intensified until the muscle has in turn molded
the bones to which it is attached, and also those with which it comes
in contact, thus altering the form and proportions of the cranium as
a whole, and giving rise to extreme variations in the size, shape, and
position of the zygomatic arch and in the development of the angle of
the jaw. At least two very distinct typi^s of skull have been estab-
lished in this way — a broad ot platycephalic type (pi. 3) and a narrow
or dolichocephalic type (pi. 5).t

By contrasting the accompanying figures of representative skulls of
these two types, with respect to the areas of attachment of the princi-

riG.49. — Side view of skull of Macrogeomyn doLichucephalus, showiug relationa of mandiblo, and
fossae for attachment of muscles.

a Anglo of mandible.
ic luci.sor capsule.
jo Jugal origin of masseter.
711 Mastoid process of mastoid bulla.
ms Mastoid process of s(iuaniosal.

mf Masseteric fossa.

mo Maxillary origin of main body of masseter.
m«.? Mandibular slielf (leading to angle in Pia-
tygeomys gymnurus).

pal parts of the masseter, the action of the muscle and its effects on
the skull may be better understood. Without rei)eating the detailed

"P^'or an irnijortaut chapter on the general snbject of the intineuee of the inu.scles
in shaping the skull in the Rodentia, see Herluf Winge, Jordfundne og nnlev.
Gnavere fra Lagoa Santa, Minas Geraes, Brasilieu, 1888, 103-110.

t These extremes in the form of the skull are brought about mainly by alterations
in the superficial or outer parts, the fundamental structures and relations remaining
very much the same in both, as shown by sectionized skulls (pis. 17 and 18).

INFLUENCE OF MASSETER MUSCLE.

105

descriptions already given under the liead of the muscle (p. !M)), it may
be stated that the principal part of the masseter arises from the side of
the maxilla in front of the zygomatic arch, and from the adjacent parts of
the premaxilla and the maxillary root of the zygoma (fig. 49, mo). It is
inserted upon the outer side of the mandible, and the area covered by its
insertion — the masseteric fossa — extends from the angle to the plane of
the front of the premolar (fig. 4!), mf). Its origin, insertion, and relations
are essentially the same throughout the group. The jugal part arises
from tlie horizontal arm of the zygoma and is inserted upon the upper
side and end of the angle of the jaw. Its size, form, area of origin, axis,
and relative imi^ortance differ conspicuously in the various members of
the series. In some forms it arises from the entire length of the hori-
zontal part of the arch (fig. 50, jo); in others from the j^osterior part
only (fig. 49, /o). The upper limit of its origin is marked by an oblique
line and a change of direction in the outer face of the jugal*

Effect on the skull. — In the long and narrow skulls, of which Macro
geomys dolichocephalus may be taken as a type, the great body of the

Fiu.50. — Side view of skull oi Platy geomys fji/umicrui nhowiog relatious of inaudible ami fos.sii^ for
atta('limeut of muscles. Letteriug same as in fig. 49.

masseter is parallel to the side of the face, its function being to close
the jaws firmly and draw the mandible forward. Its princii)al origin
is maxillary, the jugal part being small and posterior to the plane of
the middle of the orbit (fig. 49, ;>>). The resulting i^rincipal movement
of the jaw is antero-i)osterior. The action of the muscle has narrowed
the zygomatic arches, rounded off their anterior angles, and lifted them
out of the way until the horizontal part of the arch is much nearer the

* Owing to the scarcity of material for dissection tlie masseter muscle itself has
been actually examined in two forms only, namely, Geomiisbursarins and Miicrofjcomiia
dolichocephalus. Its relations in these species, studied in connection with the well
defined fossae on the skull marking its origin and insertion, furnish a very good
guide to its modifications and to the jiart it has played in producing the several
types of cranium known in the group.

106

NORTH AMEKICAN FAUNA.

[NO. 8.

top of tbe skull in front tliaii l)eliin(l (tig. 40). The fibers of tliejugal
braneli are nearly vertical, and are of little use except in drawing up
the back part of tbe jaw. This may be seen from fig. .lU: the muscle
passes downward from the zygonni {.zy) to the angle of the Jaw [a).

In the broad and fiat skulls, of which Flatygeomys (jymnnrus may be
taken as a type (fig. 50), the Jugal branch of the masseter is largely
developed, its function being to move the jaw sideways at the same
time that the maxillary i)art brings the teeth firmly together. The
resulting principal movement of the jaw is obliquely transverse. In
producing this lateral movement tlie jugal branch is aided by the
pterygoid muscles, but the latter must have played a very subordinate
part in molding the skull. The jugal part of the masseter in the
j)latycephalic series is not only of relatively large size, but the area of
its origin is greatly extended (fig. 50, jo) and the axis of its fibers has
become more nearly horizontal than vertical (fig. 54, a to zy). Its origin
occupies the outer and inferior surface (and i^robably most of the
inner surface also) of the horizontal part of the zygomatic arch for

ZX-.

a

KlG. 51. — Macfogeomys dulichvcephalus. Via. h'i. — Flatygeomys gyinnurus.

Posterior part of craiiiiiiii from above, showing relations of maudible iu place.

a Angular process of uiaiulible. //• Frontal.

cp Corouoid prooes.s of mandible. pa Parietal.

ic Incisor capsule (covering root of lower so Supraoccipital.

incisor). sq Squamosal.

ip Interparietal. 2y Zygoma.

its entire length, its anterior end being in front of the plane of the
orbit. The action of this part of the masseter has drawn the zygomatic
arch far outward and has pulled the anterior angle downward until
the latter is lurther from the plane of the top of the skull than the pos-
terior end of the arch. The angle is thus drawn down until it reac^hes
four-fifths of the way from the plane of the toj) oi the skull to the plane
of the molar alveolus, overreaching and overarching the maxillary or
principal part of the masseter muscle, which ojx rates beneath it (tig.
50, which should be cdiitrasted with fig. 40 of Macrof/coniys doUchoceph-
aius). The insertion of the muscle has produced an equally extraordi-

I

JAN., 1S95.]

INFLUENCE OF MASSETER MUSCLE.

107

nary effect upou the shape of the under jaw. The sides of the Jaw are
not only spread widely apart in conformity with tlie great breadth of
the skull, but in addition tlie fibers of tlie uiasseter that are inserted
on the angular process have stiniuhited this process to push out side-
ways until it reaches off like a long arm at nearly a right angle to the
axis of the skull (figs. 53 and oi, a)* The lengthening of this proc-
ess was clearly necessitated in order to coutinue the effective action
of the muscle. Furthermore, the segregation and specialization of the

Fig. 52. —Macrogeoniys dolichneephalug. Fig 54. — I'latygeomys gymmuus

Transverse vertical section of skull, with mandible in position, showing relations.

a Angular process of mandible.
ale Alisphenoid canal.
alh Horizontal arm of alisphenoid.

c Con<lyle of mandible.
cp Coronoid process of mandible.

/ Angle of crowns of closed molar.i.

ic Incisor capsule (covering root of lower in-
cisor) .
np Narial passage.
pa Parietal.
pf Pterygoid fossa.
iq Squamosal.
zy Zygoma.

two parts of the masseter in iho, platycephalic series has resulted in the
production of a long and well-defined horizontal shelf extending for-
ward from the angle of the jaw to the base of the ascending ramus
(fig. 50 mss). This shelf is totally wanting in Macrogeomys doliehoceph-
alus and the other doUchocephalic forms in which the jugal part of
the masseter is relatively unimportant and the principal movement of
the jaw is fore and aft instead of transverse. The relations described
may be seen to good advantage in the accompanying drawings (figs.
49-54).

Effect on the teeth. — While from the nature of the case it is ciearly
impossible to observe exactly what happens, either in the muscles or
the teeth, during the act of mastication, it is at the same time i)ermis-
sible to draw certain inferences from the mechanical construction of
the apparatus. In the case of tlie teeth, considered as the focus of the
cutting machine, it has been already shown that two types exist, one

* In M. dolichocephalus the angle projects only 2| mm. beyond the plane of the
zygoma (fig. 52), while in P pymnurus it jtrojects 10+ mm.

108 NORTH AMERICAN FAUNA. [no. 8.

in which the crowns of the upper inteiinediaiy inohuvs are broadly ellip-
tical and bear two enamel i)lates (one on eueh face); the other in which
the crowns are narrowly elliptical and bear only one enamel plate
( which is on the anterior face). It has been shown further that the pres-
ence of two enamel plates is always correlated with an anteroposterior
movement of the jaw, and that tiie presence of a sinj>le plate is always
correlated with an obliquely trausv^erse movement of the Jaw. A care-
ful study of the cutting- blades in each instance shows that an antero-
posterior movement is accompanied by a to-aud-fro i)laning' action in
which two enamel blades are serviceable; and that a. transversely
obli(iue movement is accompanied by a lateral shearing- action in which
only a single blade can be used. In accordance with the well-known
law that useful structures are preserved and useless structures sup-
pressed, it is logical to infer that tlie direi'tion of the dominant move-
ment of the jiiw has determined the presence or absence of the posterior
enamel plate; and since the movement of the Jaw is controlled by the
masseter muscle, it is evident that the number of enamel plates on the
u])per intermediary molars may be traced back to the influence of tiiis
nuisde.

In the course of the evolution of the two types Just described it seems
evident that as soon as the principal movement of the Jaws in the line
leading- to Macrofjeonii/.s doUchocephaluH came to be fore and aft it was
settled that the form of the posterior part of the cranium should be
narrow; that the angle of tlie under Jaw should b(^ shortly truncate;
that tlie grinding- teeth should be broadly elli|)tical, and that the poste-
rioi- enamel plate of the upper series should be retained; and when the
priiKii)al motion of the Jaw in the ancestors of Phityfjeomys gymnurm
v-Mwy to be obliquely transverse, from that moment it was predeter-
mined that the hinder jiart of the skull should be broadly expanded;
that a long- arm like process should spring from the angle of the jaw;
that the grinding teeth should be transversely flattened, and that the
posterior enamel i)late of the upper series should disappear.

CHAPTER IV.
SYSTEMATIC DESCRIPTIONS OF GENERA AND SPECIES.

Genus GEOMVS Rafinesque, 1817.

Pis. 1,7; 9, 12; pi. 1.5, lags. 11 Jind 12; pi. 17, lig. :^: pl.l8,tig.l; pi. 19, fig. :5, ami text

fig. .5.5; maps 1 ;ni(l 4.)

Type Mus iuza Onl. 1815, from Au(JU.si a, (Jkorgia. {z^Geomi/s phietis Raf., 1817).

Geomi/s Raliuesciiic, Am. Monthly Magazine, II, No. I, Nov., 1817,45. Type G. pine t is

Raf. (=.yM.'* titza Ord. 1815), from pine liarrens near Augusta, Ga.
Diplostuma Ratiuesqne, Ildd, 1817, 44-45.
Saccophonis Kuhl, Beitriige zur Zool., 1820, 65-66.
Pseudostomn Say, Long's Exi)d. to Rocky Mts., I, 1823,406.
Asromys Lichtenstein. Abh. Akad. Wiss. Berlin ^822), 1825,20, fig. 2.

Dental characters. — Upper pieinolar with three enamel plates (the
posterior absent). Tapper pni decidedly longer than lower (in the other
genera they are subeqnal); shaft of upper pm decidedly concave for-
ward, except in a single species {G. IvJescens). First and second upper
molars with two enamel plates each, the posterior complete; posterior
curvature of m^ and anterior curvature of iii, hardly ai)parent.

Last upper molar a single subcylindric or subtriangular prism with-
out lateral sulcus on either side (and consecpiently without heel) : outer
enamel plate normally straight; inner and outer plates commonly sub-
equal, or outer somewhat shorter, both reaching posterior face of tooth.
I'pi^er incisor strongly hisulcatc (fig. 1*2^ and 2li '; pi. 15, figs. 11 and 12).

Cranial characters. — Skull simi)le. without any very striking external
characters. Orbitosphenoids small and narrow, not reaching alisplie
noids (pi. 17, fig. 3); sphenoid fossie C()rre^l)ondingly elongated, reach-
ing forward to orbital plates of frontal; alisphenoids short posteriorly,
ending on floor of brain case about on plane of front ends of audital
bulhe; pterygoids large, always forming more than Imlf of the palato-
pterygoid extensions; mesethmoii plate large, some wliat rectangular,
nnich longer than high, and wholly superior to vomer (not dipping-
down between vomerine wings as in Pappof/romys); endoturbinals col-
lectively forming a quadrate plate, the anterior border of which is ])ar
allel to the cribriform plate (pi. 19, fig. 3) ; first endoturbiual rounded
and only vSlightly expanded anteriorly, its inferior border fiiHing (as
the OS planum) in the front of the others and articulating with the
anterior third of the internal vertical plate of the maxilla — the os pla
num thus extending anteriorly in front of the lower endoturbiual nuich

further than the length of the latter.

109

110

NORTH AMERICAN FAUNA.

In the elongated sknlls of Geomys bursarius and tuza the lower part
of the sphenoidal fissure, on the floor of the orbit, differs from its con-
dition in any of the other groups (fig. 55). In all of the others a fenes-

I

Fig. 55. — Side view of r^knll of Geomys hursariiis from outside, zygomatic
sawed off to sliow bottom of orbit. Animal a fully adult S , from Knoxville, I
(This Hgnre should be compared with the corresponding view of Cratoc/eomys
riam'u fig. 4.)

1. Infraorbital foraiiien.

2. Posterior (orbital) opening of infraorbital canal.

3. Vacuity in front of prespbenoid and ascending wing of palatine.

4. Vacuity in prespbenoid, l)ebind ascending wing of palatine.

5. Optic foramen (in orbitoax>benoid bone).

fl. Foramen rotundum and foramen ovale (whicb have hero coale.'iced).

7. External auditory meatus.

8. Spbenoidalfi.ssnre (upper part).
.\scendiiig wing of vertical plate of palatine.
Alisplicnoid.
Condyle of exoccipital.
External pterygoid plate of palatine bone.
Frontal.
Hamular ])rocess of pterygoid bone.

/. Lachrymal.
n. Mastoid proce.s.s of raa.stoid bulla.

6. il.Tstoid bulla.
«. Mastoid process of squamosal.
X. Maxilla.
/(. Nasal.

Parietal.

Premaxilla.

Paroccipital process of exoccipital.

Prespbenoid.

Pterygoid.

Supraocciiiital.
sq. Squamosal.
tb. Tympanic or nudital bulla.

arch
owa.
mer-

ajA.

as.

c.

epl.

St.

h.

pa.
pmx.
pp.
ps.
pt.
so.

I

truin (fig. 4,")* penetrates tlie iiiterorbital septum, which at this point
consists of the presphenoid oul^'. In Geomys bursarius and tuza the basal
part of the sf^henoidal fissure is unusually broad, and the septum at

* In some cases, particularly in Orthoyeomys and Zjigoyeomys, this fenestrnm is sub-
divided into two or even three parts, but they all invariably penetrate the prespben-
oid; they are never in front of it.

JAN., 1895] THE GENUS GEOMYS. Ill

its bottom, which here consists of both palatine and pres]ihenoid, is
l)i'r(orated by two fenestra, whicli hiok completely throngh the sknll
from orbit to orbit. The posterior is the usual opening- in the anterior
part of the prespheiioid (flg. 55,^) ; the other is in front of the pre-
sphenoid and is bounded anteriorly by a process from the maxilla, which
here rises to join the frontal (tig-. 55,'*). Hence in Geomys hursarhis
there are three openings in the bottom of the orbital fossa, arranged
seriatim, one in front of the other. Tlie first is the posterior outlet of the
infraorbital canal (fig-. 55,-); the second is the vacuity here mentioned,
which penetrates the skull in front of the pres])henoid (fig. 55,^); the
third is the usual fcnestrum in the anterior part of the presphenoid
(fig. 55,'*). The ojiening in front of the presplienoid is completely sur-
rounded by the maxilla and ascending wing of the palatine — the former
bounding it in front, the latter behind — for the ascending wing of the
vertical plate of the palatine (fig. 55, ajH) here rises along the front of
the presphenoid between the two fenestra in question and articulates
with the maxilla, the orbital plate of the frontal, and the orbitosphe-
noid. (See also fig. 10.)

Tlie condition here described has not been observed except in the
elongate skulls of Geomys bursarius, tuza, a.i\d per sonat us, and is imper-
fectly developed in the latter. It reaches its highest development in
Geomys bursarius, and does notocccurin the closely related G. lutescens,
which has a short skull. A condition simulating it sometimes exists
in Orthogcomys, in Avhich there are several (usually two or three) small
jierforations in the anterior part of the presphenoid, but the relations
of the ascending wing of the palatine are not the same. Very young
specimens of Cratogeomys resemble the adult of Geomys in the presence
of a fenestrum in front of the presphenoid and ascending wing of the
palatine, but the fenestrum disappears as the animal matures, a vestige
of it remaining as a foramen (on each side), which opens from the floor
of the orbit obliquely forward and downward into the narial passage.

The genus Geomys, even as here restricted, comprises three series or
groups of species: (1) the texcnsis-brericeps series, (2) the tuza series,
and (3) Geomys busarius.

(1) The texcnsis-breviceps series inhabits Texas, Louisiana, Arkansas,
and the Great Plains, and includes eigiit species and subspecies, as
follows: arenarius, tcxensis, lutescens, breviceps,breviceps sagittaHs, brevi-
ceps (ittwateri, personatiis, and personatus fallax. Most of these, par-
ticularly arenarins, texensis, and brericeps, are small generalized forms
suggesting relationship with Ihomomys and Pappogcomys. Indeed,
these animals are very much alike in many ways and the skulls agree
in general form, lightness, in the small rounded brain case, slender
and nearly parallel zygomata, narrow pterygoids, and many other
characters, though dilfcringconspicuoiisly in the teeth. It seems evident
that they are but little removed from the trunk line of the group, and
that both the tuza and the bursarius series are offshoots from the bred-

112 NORTH AMERICAN FAUNA. [no.8.

cejys stem. Geomys hreviceps seems to be the central or parent type
from wWcb three widely different species originated, tuza on the east,
bnrsariiis on the north, and luteseeiis on the west. To tlie eastward
only ai narrow gap separates the range of hreviceps from that of niobi-
lensis of the tuza series, which, though specifically distinct, was evi-
dently derived from the hreviceps stock. Still further east mohileusis
passes in totuza. On the west hreviceps shades toAvard and probably
will be found to intergrade with lutescens. On the north only a nar-
row hiatus separates it from hnrsarins, the most specialized type of
the series. Specimens of hursarins from southern Missouri suggest
that the gap between it and hreviceps is not very wide; if continuity of
range between the two forms is anywhere found this gap may be
bridged even at the present time (see map 4).

(2) The tuza series inhabits the South Atlantic and Gulf States south
of the Savannah Kivcr and east of the Mississippi (mah 4, A), and com-
prises three forms, tuza, tnza mohilensis, and tuza floridanus. They are
locally known by the singularly inappropriate and misleading name
'Salamander.' The members of the tuza series agree among themselves
and differ from the remaining forms of the genus Geomys in having
longer and more naked tails, and in numerous cranial characters. The
shape of the skull in potile is decidly convex, the rostrum long and
decurved, the nasals long and slender and constricted in the middle, giv-
ing them a somewhat hour-glass shape. The interparietal is perma-
nently distinct from the supra-occipital and is normally much larger
than in any of the other groups, though in G. mohilensis it is nearly
obliterated in old age by the encroachment of tlie ridges that unite to
form a sagittal crest.

The tuza group differs not only from hursarius, but from all other
known members of the family, in the disproportionate length of the
upper premolar in relation to the other molarilbrm teeth. It is merely
double the length of m-^. The lower premolar is much shorter, particu-
larly in jioridanus.

(3) Geomys hursarins inhabits the u])per Mississippi Valley (map 4, b)
and stands alone at the end of the northern branch, just as Geomys tuza
occupies the end of the eastern branch of the restricted genus Geomys.
The skull is elongated and angular, the frontal compressed between the
orbits, the palatopterygoids broadly Ungulate, and the sagittal crest
high; but the most important departure fi'om its allies is found in the
anterior part of the craniofacial axis, and consists mainly in the broad
articulation of the ascending wings of the palatine bones with the hori-
zontal shelf of the orbitosphenoids, and in the presence of a fenestrura
looking completely through the skull in front of the presphenoid. G.
hursarius presents the extreme of differentiation occurring in the bisul-
cate series inhabiting the United States.

The following brief tabular statement of some of the cranial char-
acters of the three members of the ^»vrt group may facilitate the identi-
fication of specimens:

JAN., 1895] KEY TO SPECIES OF GEOMYS. 113

IHfferentiftI cr(iiti((l iharaitcm of the ineinhcr.'s nf llie tiiza t/roiqi.

Tcraporal impvossions

Frontal (iiiteroibitally)

Ascending brauclies of premaxilla

Palato]iteryg(>id

Audital bnllii!

Interparietal

Mobilensis.

Tuza.

Floridanus.

United in a sagittal crest . . ! Distant 1 Distant.

Very broad ! Narrow ; Narrow.

Motleratu ; Moderate i Very broad and

j ! blunt.

Narrow, sides i)arallel- ... Lingulate-ctineate Lingnlate eune

i I '^^^'

Small Small i Large.

Deeply notched posteriorly Not notched i Not notched.

KEY TO SPECIES AND SUBSPECIES OF GEOMYS BY CRANIAL AND DEN-
TAL CIIARACTERS.

[Based on skulls of adult males only.)

(1) .TUGAL equal to or shorter thai) basioecipital (measured from condyle),
a' Sagittal erestprenenl.

ft' Zygomata strongly angular (standing out at right angles) ; jugal
broadly rounded anteriorly.

Size large ; audital biilbe nornial persouaiits

Size medium; audital buUie short and swollen (almost 8ubglobular)./rt?/ax

h- Zygomata rounded ; jugal narrow anteriorly ; size small safjittaJis

a- Saijiltal crest absent.

Temporal ridges prominent ; squamosal arm of zygoma ending in

a knob arenarins

Temporal ridges not prominent; squamosal arm of zygoma not

ending in a k nob iexensis

(2) Jugal longer than hasioccipital (measured from condyle).

c' Sagittal crest strongly dcreloped — long and high; size larg<«t hursarius

c- Sagittal crest feebly developed or absent; size medium or small.

(P Nasal bones hour-glass shaped; strongly constricted near middle.

e' Temporal impressions uniting in sagittal crest mobilensis

e'^ Temporal impressions not uniting in sagittal crest.

Audital bulhe small; not swollen; nasals broad posteriorly tnsa

Audital bulhe large, swollen; nasals narrow posteriorly .. .floridanii't
rf^ Nasal bones not hour-glassed shaped; slightly or not constric-
ted near middle.
/' Frontal strongly depressed interorbitally ; zygomata l)r()adly

rounded; nasals very narrow posteriorly, notched behind. ftrtr(cej;s
/^ Frontal slightly or not depressed; zygomata angular,
strongly divergent anteriorly.
Temporal ridges prominent, divergent anteriorly; nasals

abruptly narrow and convex posteriorly attwateri

No temporal ridges; temporal impressions parallel or
meeting in sagittal ridge; nasals truncate or emargi-
uate posteriorly liitescens

GEOMYS TUZA (Ord).
(Frontispiece and pi. 7. fig. 1; pi. 13, tig. 9; i)I. 15, fig. 12.)

Mas tnza Ord, Guthrie's Geog.,2d Am. ed., Ii, 1815, 292 (based on Mitchill's "unde-

scribed little <iuadruped of Georgia'" — atm postea).
Geomys pinetis Rafinesque, Am. Monthly Magazine, vol. ii, No. I, Nov., 1817, 45 (type

of geuus Geomys).
Undescrihed little quadruped of Georgia, Mitchill, New York Medical Repository, V,

1802, 89. (Descr. orig. on which the name .Vns tuza of Ord was based.)

7433— No. 8 8

114 NORTH AMERICAN FAUNA. [no. 8.

Eamster of Gcoryia, Audersou, 2d Am. from 8tli London ed. of Bewick's Hist, of
Quadrupeds, 1848,* 326 (accompanied by figure Avith claetik pouches properly
turned in).

Type locality. — Pine barrens uear Augusta, Georgia, t

Geographic distribution. — Pine barrens of (leorgia ( and probably
northern Florida also), within the Anstroriparian fannal area (map 4).

General characters. — Size medinni or rather large; tail long atid
naked; feet moderately well haired; a small naked pad on end of nose.

Color. — Upper parts cinnamon brown, strongly tinged with fulvons
in fresh pelage; only a faint trace of darker median dorsal stripe;
under parts dull ochraceous buff; hairs of feet whitish.

Cranial characters. — Skull rather large and angular (PI. 7, fig. 1), its
upper surface convex in i^roflle (due in part to the strongly decurved
rostrum and in part to the absence of sagittal crest); zygomata diver-
gent anteriorly, the maxillary root sloping strongly backward; temporal
impressions never uniting in a sagittal ridge, but forming permanent
temporal ribs, which in the males are elevated on both sides and sepa-
rated by an interspace or sagittal area 3 to 4 mm. in width. In the
females the intersj)ace is broader and usually thickened so that it is
flush with the top of the temporal imj^ressions. Interparietal very
large and broad. The frontal is narrow interorbitally; postorbital
prominences marked; palatopterygoids lingulate-cuueate, the base
slightly or not excavated on outer side; audital bulhe small, normal;
basioccipital strongly wedge-shaped, truncate anteriorly.

Skulls of (t^. tuza may be distinguished from those of mohilensis by the
presence of distant temporal ridges instead of a sagittal crest; by the
narrow frontal (interorbitally); by the lingulate-cuneate (instead of
narrow strap-shaped) palatopterygoids, and by the very large inter-
parietal which is not notched behind (fig. G e). Skulls of tuza differ from
those oi Jioridanus in much narrower ascending branches of premaxilla,
broader nasals posteriorly, more strongly wedge-shaped basioccipital,
and much smaller audital bulhe. The relationship with Jloridanns is
much closer than with mohilensis. The protile of the top of the skull
is more convex than in either of the others.

Measurements. — Average of ten males from type locality (Hollywood,
Georgia, 11' miles soutli of Augusta): Total length, 2G9; tail vertebni?,
89,5; hind foot, 34.4.

* The copy cited by Coues (Monograj)hs of N. Am. Rodentia, 1877, 615 footnote) has
the same pagination, but a somewliat different title page (different publisher) and is
not dated. The eighth Loudon edition of Bewick was published in 1824. The
only mammals described in the American reprint not in the original are the grizzly
bear, hamster of Georgia, and mammoth.

iThe tj'ps specimen was sent Dr. Mitchill from Augusta, Ga., in July, 1801, by
.Josiah Meigs, president of the University of Georgia. In the letter that acco.iipa-
nied the spccimeu Mr. Meigs said: "For the space of about 100 miles, between
8avannali and Augusta, the land on each side of the road is almost covered l»y the
heaps of loose earth raised by it." — New York Medical Repository, Y, 1802, 89.

JAN., 1895.]

GEOMYS TUZA. 115

Average of nine females from same j)]ace: Total length, 249; tail
vertebra', 82 ; liiud foot, 32.

For cranial measurements see Table C, p. 208.

Specimem examined. — Total number 32: twenty from type locality,
Hollywood, 12 miles south of Augusta, Georgia; and twelve from
Butler, Georgia, (latter not typical).

General reniarls. — SpeL-imens from Butler, near the western border of
Georgia, are intermediate between fuza and mobilensis. In color they
resemble the latter, while in cranial characters they are nearer the
former.

It is an interesting fact that the first description of this species — and
not a bad description either, considering it was written nearly a century
ago — was from the pen of a member of Congress, the Hon. John Mil-
ledge, Ilepresentativ^e from Georgia. It was published by Dr. Mitchill
in the New York Medical Repository in 1802 (vol. v, p. 89), and runs as
follows: ''One of the little animals that burrows in the pine land, only
known in Georgia, was caught by Mr. Stephen Pierce, living midway
between Savannah and xVugusta. Its body is of the length and thick-
ness of a common-sized rat, amd of the same color: the head between
that of a rat and a mole, with small whiskers and short snout: the tail
without hair, but shorter than that of a rat : the fore feet like those of
a mole, with nails near an inch long: the hind feet like those.of a rat,
but the nails not of the same length, each foot having five claws : very
sparkling small eyes: also short ears: teeth like a squirrel, and full as
long. On both sides of the jaw, externally, are sacks or wallets, where
it deposits its food, and each will contain as much as can be put in a
large tablespoon. Little or no fur, and the hair of the length of a wood
rat. The whole face of the pine country is covered with little mounds
made by this animal, of the circumference of a peck, and from 6 to 8
inches high. It is by no means active, but remarkably fierce. No
common wooden place of confinement can hold it long, as it gaaws its
way out. It lives entirely on roots, and is very fond of the sweet
potato, and often i^roves injurious to the planter by getting under'his
stacks. It ai^pears to move nearer the surface in the spring and fall
than at any other season. It is surprising, that though the work of
this creature is seen throughout the country, in the region of the long-
leaf pine, and in that region only, yet sach is its skill in burrowing, and
acuteness of hearing, that there is no animal in all our State so seldom
caught or seen."

CxEOMYS TUZA FLORIDANUS (Aiul. and Bach.).
(PI. 7, figs. 3 and 4; PI. 10, fig. 1; PI. 14, fig. 16.)

Pseudostomn Jior'ulana And. and Bach., Quadrupeds of Xorth Am., Vol. iii, 1854, 242-

24.5.
Geomtjn tiiza Goode (not Ord), Powell's Report Colorado R?ver, 1875, 281-285 (habits).

Type locality. — St. Augustine, Florida.*

*Auilubon and Bachman did not discriminate between the Georgia and Florida
animals, but all of their Florida specimens came from St. Augustine.

116 NORTH AMERICAN FAUNA. [no. 8.

General characters.^^umhiv to G. tnza, but much (linker in color;
fore feet larger; tail sHghtly more hairy; <lifi(er.s also in cranial
characters.

Color. — Upper parts dull sooty-plumbeous, becoming cinnamou-drab
on the sides; under parts plumbeous, more or less washed Avith buffy;
an irregular white patch under chin and throat.

Cranial characters. — Skull long, with very angular zygomatic arches,
much as in tuza and mobilensis. G. floridanus differs from G. tuza in
broader and blunter ascending branches of premaxilla, narrower nasals
posteriorly, somewhat broader jugals anteriorly, more rectangular (less
strongly wedge-shaped) basioccipital, and much larger audital bulhe;
from mohilensis in much larger audital bulhe, narrower frontal, less
spreading and more depressed arches, much broader ascending branches
of premaxilla, .less flattened brain case, lingulate-cuneate instead of
narrow palatopterygoids, and in the presence of temporal ridges instead
of a sagittal ridge. The angular process of the mandible is nuich less
deeply notched at base anteriorly. In G. floridanus the interspace
between the two grooves of tlie upper incisor is broader than in either
tuza or 'mobilensis, and the head of the jugal is more deeply mortised
into the maxillary arm of the zygoma.

i^pccimens examined. — Total number 25, from the following localities
in Florida: Chattahoochee, 2; Pomona, 4; Gainesville,!; San Mateo, 0;
Tarpon Springs, 12.

Measurements. — Average of three males from San Mateo, Florida
(measured in flesh by Dr. W. L. Ralph) : Total length, 288; tail vertebr;e,
94 ; hind foot, 35.5. Average of three females from same locality : Total
length, 235; tail vertebrie, 77 ; hind foot, 33. For cranial measurements
see Table C, p. 208.

General remarls. — The foregoing description has been drawn up from
specimens from San Mateo, Putnam County, Florida,* only 25 miles
from St. Augustuie, the type locality of the species. Specimens from
further south on the peninsula are somewhat different.

The best and almost the only authentic account of the habits of this
species is from the pen of the eminent director of the TT. S. National
Museum, Dr. C. lirown Goode, by whom it was contributed to Cones'
monographic paper on the group, published in 1875. f Dr. Goode kept
a number in confinement for several weeks and was thus enabled to
make the "following interesting observations on their habits. He says:
" They may easily be coniined in a wo®den box, with sides 8 or 10 inches
high, having dry. sand 2 or 3 inches dee»p on the bottom. No cover is
necessary; I have never seen one look up from the earth, and have

*These specimens Averekindly presented to me by Dr. W. L. Ralph, of Utica, Ne
York, who collected them himself and measured them m the flesh.

t Abstract of results of a study of the genera Geomt/s and Tliomomys. Powell's Expi.
Colorado River, 4^, 187.5, 21.5-28.J. Addendum B.— Notes on the "Salamander" of
Florida, by G. Brown Goode, 281-285.

JAN., 1895] GEOMYS TUZA PLORIDANUS. 117

rarely known them to attempt to escape. They require no water, and
no food except sweet potatoes. A single potato of moderate size will
feed a salamander for three days.

" The senses of sight and hearing seem in them to be very dull. An
object may be held within a short distance of their eyes without attract-
ing their attention; but the moment one is touched, he turns with a
jump, snapping fiercely, much to the detriment of fingers which may be
near. If two are confined in the same cage, the one does not seem
aware of the presence of the other, unless they accidentally come in
contact. Their eyes are small, dull, and without expression. Their
sense of smell I judge to be very delicate, from the manner in which
they approach the hills of potatoes. Their motions are surprisingly
quick and energetic, their activity never ceasing from morning to night.

" They are very pugnacious, and a rough-and-tumble combat between
two vigorous males woald seem terrific, if their size could be magnified
a few diameters in the eye of the spectator. Every muscle of their com-
pact, elastic, stout bodies is brought into action, and they plunge and
bite with wonderful ferocity. A battle is usually followed by the death
of one or both. I have examined them after death and found the
whole anterior part of the body bruised almost to the consistency of
paste, the bones of the legs crushed in four or five places. When two
come together in the cage, their salutation is a plunge and a bite.

" I watched their burrowing with much interest. They dig by grub-
bing with the nose and a rapid shoveling with the long curved fore
paws, assisted by the pushing of the hind feet, which remove the dirt
from beneath the body and propel it back with great power a distance
of 8 or 10 inches. When a small quantity of earth has accumulated in
the rear of the miner, around he whirls with a vigorous flirt of the tail
and joining fore paws before his nose, he transmutes himself into a sort
of wheelbarrow, pushing the dirt before him to a convenient distance,
and repeating the act until the accumulation is remo^'ed, then resuming
his mining. Any root or twig which blocks his way is quickly divided
by his sharp chisel-teeth. * * * Tlie direction of the burrows may
easily be traced by the loose hillocks of white sand which are thrown
up along the line at intervals of 3 or 4 feet. These are the -dumps'
made by the burro wer in throwing out his refuse accumulations. Each
consist of about a peck of loose sand, and, by the casual observer,
might easily be mistaken for an ant-hill. No opening is visible, but by
digging under the hill a hole is found, the mouth of the adit to the
mam tunnel, which may be 3 feet below the surface if made in cold
weather, but perhaps not more than (> inches if in summer. One of the
mounds is thrown up in a very few moments. I have seen 30 raised in a
single night on the line of one tunnel; this would represent nearly 100
feet of tunneling. I have seen 150 in one continuous row raised in
about two days ; this would make between 400 and 500 feet of burrow
completed in that short time, apparently by one little animal, an amount

118 NORTH AMERICAN FAUNA. [no. 8.

of work which may seem incredible to one who lias not watched the
restless movements of these animated plows, which are seemingly as
well adapted for piercing tlie sand as birds are for -cleaving the air.
The bnrrows are about 2i inches in diameter. * * * The nests are
large chambers, 1 or 2 feet from the main tunnel, with which they are
connected by side passages, which leave nearly at right angles. Here
the miners lay up a supply of provisions and the chambers are often
found to contain a half bushel of sweet potatoes cut up into chunks
as large as peach stones, and of convenient size to be carried in the
pockets. * * * In these side chambers the salamanders rear their
young, building a nest of grass, i)ine needles, and live-oak leaves. 1
found them breeding in April."

Dr. Goode remarks that the name ' salamander,' by which the species
is universally known in the South, " luiiy allude to the safety enjoyed
by these little animals in their subterranean abodes at the time of the
devastating fires which sometimes consume the pine forests. After such
a conflagration has passed over their heads, destroying every other kind
of life, they are seen at work among the ashes, very good types of the
salamander of fable."

Mr. Morris M. Green, who obtained specimens for the Division at
Pomona, Putnam County, Florida, in June, 1S89, furnished the follow-
ing notes respecting their habits : " The hills of the ' salamander,' as the
Florida Geomys is called, are abundant in the pine woods and clearings,
on rather low and moist land. Their tunnels were from 4 to 24 inches
below the surface; the hills were thrown up at intervals of from 2 to 6
feet, and contained about a peck of dirt each. The night and early
morning seemed to be their favorite time for working. It is very easy
to trap a 'salamander' when fresh mounds are found. By sweeping to
one side the heaps of dirt, traces of the hole through which the earth
was brought and its direction can be easily found. A minute's work
with the spade will usually expose the tunnel lying to one side of the
hill. Place a steel traj) in the tunnel, and cover up the breach with a
piece of pine bark or some palmetto 'fans.' If the breach is left open,
the animals will carry dirt to shut out the light, and thus clog the trap,
whereas if the opening is closed they will step in the trap and are
caught. A break is often repaired within half an hour, or it may be
left for nearly a day. In mending an opening it is astonishing how
compactly the earth is packed; in one case an animal closed an opening
so securely that the tunnel could not be found at all until another shaft
was sunk in search of it.

"A 'salamander' caught in a trap is a picture of fury and spite, bit-
ing at everything within reach of its jaws, and sometimes breaking its
front teeth in venting its rage on a trap.

" In the cheek pouches of one were some pieces of pine roots, and
some grasses were found in the tunnels. The animals do serious injury
to orange and pear trees by gnawing tlie roots. Sometimes the roots

i

JAN., 1895.]

GEOMYS TUZA MOBILENSIS. 119

are gnawed off so completely that the tree can be pusbed over with one
hand. They also feed on sweet potatoes. But when an animal enters
a garden or an orchard, and betrays itself by throwing up hills, there
is no excuse for not ridding the place of it, as it may be easily caught
in a steel trap. It is claimed that the 'salamander' works near the
surface from September to March, retiring deeper in the ground during
the hot season."

GEOMYS TUZA MOBILENSIS subsii. nov.

(PL 7, figs. 2, 5, aud 6; pi. 10, fig. 2; pi. 14, fig. 15; text fig. 6, f and g.)

Typ6 from Mobilf, Bay, Alabama. No. ^^U i ad. U. S. Nat. Museum, Department
of Agriculture collectiou. Collected April 26, 1892, by Russell J. Thompson.
(Original No. 50.)

Geographic (listribution. — Soutliern Alabama and adjacent part of
northwest Florida, witiiin the Austroriparian zone (map 4).

General characters. — Similar to G. tuza, but somewhat smaller, aud
much darker in color; tail shorter, nearly naked; feet scant haired.

Color. — Upper parts dark, generally sepia or bistre, washed on sides
of face and body with golden brown or ochraceons, intimately mixed
with black-tipped hairs; top of head, between eyes and including ears,
dusky, with an ill-delined dorsal band of the same jolor. Under parts
dark plumbeous, faintly Avashed with dull pale fulvous. Hairs of feet
whitish. More or less white about throat and pouches.

Cranial charactcrs.-^SkuW very long aud angular (pi. 7, lig. 2); fron-
tal broad aud high; top of skull in profile strongly convex; isygomatic
arches broadly spreading, divergent anteriorly, and angular; brain
case broad and tlat; palatopterygoids narrow, their sides parallel;
temporal impressions in adult males meeting in a low but well-developed
sagittal ridge; interparietal deeply excavated posteriorly (trousers-
shaped), reduced in advanced age by meeting of temporal ridges (fig.
6, /and g). G. mobilensis differs from G. tuza in the great breadth of
the frontal interorbitally; the narrow palatopterygoids; the presence
of a sagittal ridge in adult males, and the very different shape of the
interpariteal (fig. G). It differs from florid anus in much smaller audita!
bulkv, broader frontal, lower and more depressed brain case, more
divergent zygomatic arches, narrower ascending branches of premax-
illa and much narrower palatopterygoids. G. mobilensis differs from
G. brcrieejys^ its nearest neighbor on the west, in general form of the
skull and in numerous details : in profile the top of the skull is strongly
convex instead of concave; the zygomatic arches are more angular and
more divergent anteriorly; the frontal is nnich broader interorbit-
allj'-; the brain case flatter; the nasal bones broader and constricted in
front of the middle; the angular process of the mandible deeply notched
anteriorly.

Measurements (taken in flesh). — Ty2)e specimen: Total length, 2G0;
tail vertebrae, 82 ; hind foot, 33.

120 NORTH AMEIRICAN FAUNA. [no. 8.

Aiwrage of four males from type locality: Total length, 250; tail
vertebr;e 81 ; liind foot, 33.5.

Avcrmje of four females from same place: Total length, 229; tail
vertebrae, 7(!; hind foot, 30.5,

For'crauial measurements see Table 0, p. 208.

Specimens examined. — Total number 23 : 9 from Point Clear, Mobile
Bay, Alabama, 2 from Brewton, Alabama, and 12 from Milton, Florida.

General remarks. — Geoniys mobilensis is an inhabitant of the low-
lands bordering the Gulf of Mexico east of Mobile Bay. How far its
range extends to the east and north has not been ascertained. In size
and coloration it seems to bear the same relation to its neighbor {G.
tuza) of the adjacent pine barrens of Georgia that G. hrericeps of the
lowlands of Louisiana and Texas bears to its relative of the higher and
drier ground further west {G. lutescens).

It seems a pity that such a strikingly marked animal as mobilensis
must stand as a subspecies, but there is no reasonable doubt of its
complete iutergradation with tuza in western Georgia.

GEOMYS BURSARIUS (Sliaw).

(Pl.l; pi. 9, figs. 8 and 9; pi. 10, fig. 6; pi. 13, fig. 11; pi. 14, fig. 2; pi. 15, fig. 11; pi.
17, fig. 3; pi. 18, fig. 1; pi. 19, fig. 3; text fig. 55.)

Mhs hursariiis 8haw, Trans. Linnean Soc, v. 1800, 227-228, pi. 8; Genl. Zoology,

Mammalia, Vol. ii, pt. 1., 1801, 100-101, pi. 138.
? Miis hidovicianus Ord. Guthrie's Geography, 2d Am. ed., 1815,292 (Nomen nudiDn).
BipJostomafusva Rafinesqiie, Am. Monthly Magazine, Vol. ii, No. i, Nov. 1817, 45.
Geomys ci neren Rafincsqne, Am. Monthly Magazine, Vol. ii, 1817, 45. {Miis I) ur sarins

renamed.)
Saccophorus hiiysarius Kuhl, Beitriige ziir. ZooL, 1820, 65.
MnssaccalHs Mitchill, New York Medical Repository, Vol. vi, n. s., 1821, 249. (Type

from Lake Superior, probably Minnesota.)
Psendostoma biirsariKS Say, Long's Expd. to Rocky Mts., i, 1823, 406.
Asromi/s caxadensis hicht., Abh. Akad. Wiss. Berlin (1822), 1825, 20, fig. 2.
Geomijs'/ biirsariiis Richardson, Fauna Boreali-Americana, i, 1859, 203.
Geomys canadensis LeConte, Proc. Acad. Nat. Sci., Phila., vi, 1852, 158.
Geomys orcfionensis LeConte, Proc. Acad. Nat. Sci., Phila., vi, 1852, 160. (Locality

erroneous.)

Type locality . — Unknown; somewhere in Upper Mississippi Valley.

Geographic distribution. — Upper Mississippi Valley from a short dis-
tance south of the Canadian boundary, in longitude 97° (Warren, Min-
nesota, and Grand Forks, North Dakota), southward to eastern Kan-
sas (Neosho Falls), southeastern Missouri (Williamsville and Hunter),
and southern Illinois (Belleville); east nearly to Lake Michigan (Win-
nebago and Fond du Lac, Wisconsin, and Cook County, Illinois);
west in the Dakotas and Nebraska to the ninety-eighth or ninety-
ninth meridian (Valley City and Hamlin, North Dakota; Burch,
Mitchel, and Scotland, South Dakota; Niobrara, Ericson, and Kearney,
Nebraska). The species belongs to the Upper Sonoran and Transition
zones. See map 4, b.

JAN., 1895] GEOMYS BURSARIUS. 121

General characters. — Size large; coloration dark; tail raedium or
rather long, scant haired, the terminal half nearly naked.

Color. — Dark liver brown or chestnnt above and beloAv, somewhat
[)aler on the belly (belly very rarely whitish); fore feet white; hind feet
soiled white; hairs of tail usually brown on basal part and white on
terminal part.

Cranial characters. — Skull long, large, and angular (pi. 1); zygomata
spreading, widely divergent anteriorly, angular; a well -developed
sagittal crest; rostrum long and narrow for size of skull; frontal nar-
row and rounded interorbitally; palatoptery golds broadly Ungulate,
tapering posteriorly, not notched at base on outer side (pi. 14, fig. 2).
The skull of G. hursarins does not require close comparison with any
other species, though the young and females are sometimes difticult to
distinguish from lutescens. The skull of the female differs from that
ot the male in much smaller size, shorter rostrum, broader interorbital
region, fuller brain case, in the absence of distinct sagittal and lamb-
doidal crests, and in the less development of processes and ridges for
nuiscular attachment. Skulls of G. bursarius differ from those of lutes-
cens chiefly in greater length and angularity, the ratio of zygomatic
breadth to basilar lentli of Hensel rarelj'' exceeding 73 percent in
adults; while in I ntesceiis this, ratio runs from 75 to 79. The brain
case is higher posteriorly and the sagittal crest is much more highly
developed. G. hnrsarius (in common with lutescens) differs from per-
sonatus notably in the angle of the anterior part of the zygomatic arch
and in the length of the Jugal. In both bursarius and lutescens, even
in old age, the anterior root of the zygoma slopes back at a consider-
able angle; in personatus it stands ont at nearly a right angle. In
bursarius and lutescens the Jugal is much longer than the basioccipital ;
in personatus it only equals the basioccipital.

Dental cliaravters. — Face of upper incisors strongly bisulcate; small
sulcus fine and close to inner edge of tooth ; principal sulcus much
larger and on or slightly external to median line; enamel face rounded
externally and between sulci (fig. 22^, and pi. 15, fig. 11). Molariform
teeth much smaller than in the other sections of the genus; crown of
last upper molar suborbicnlar, without heel.

Upper molariform series. — The upper premolar curves and slopes
strongly forward and is concave anteriorly; the last molar curves
strongly backward and is concave posteriorly. The intermediate molars
curve both backward and outward; the first only slightly backward,
the second strongly; both are concave externally; their roots divari-
cate, the first sloping forward, the second backward. (A second and
greater point of divergence is between the premolar and first molar.)
Tlie ])remolar is nearly one-third longer than the last molar. The inter-
mediate teeth are about as long as the premolar — sometimes longer.

Lower molariform series. — All the teeth are short compared with those
of the upper series; premolar longest, largest, heaviest, and curves

122

NORTH AMERICAN FAUNA.

[no. 8.

strouffly forward; last molar smallest, shortest, and curves strongly
backward; the intermediate teeth intermediate in length* Premolar
strongly concave anteriorly and nearly as concave ontward ; nij slightly
concave anteriorly, strongly concave outward, and somewhat twisted
on its axis; nij strongly concave outward and faintly anteriorly, with
a slight twist; nis strongly concave posteriorly and moderately so out-
wardly.

Average measurements of 26 specimens of both sexes from eastern
North Dakota (measured by J. Alden Loring) : Total, 270; tail vertebriie
80; hind foot, 35. Average of C males from same localities: Total, 21)0;
tail vertebrae, 90; hind foot, 37. Average of 10 females: Total, 2G5;
tail vertebra^ 78; hind foot, 34. Average total length of 20 males and
20 females from Elk Eiver, Minnesota, measured in tlesh by Vernon
Bailey : Males, 284; females, 243. In both cases many of the specimens .
are not full grown, hence the measurements are toe small. Unfortu-
nately no satisfactory series of measurements is available.*

For cranial measurements see Table A, p. 204.

General remarls. — Geomys Jmrsarhis is a well-marked species, easily
distinguishable by color alone from all the, other bisuicate forms. It is
also the largest species inhabiting the United States, although varying
considerably in size in different localities. The largest form inhabits
the region about Knoxville, Iowa, where the males average a foot in
length.

Geomys hursarius is of much greater economic consequence than all
the other species combined, for the reason that its home is in the fertile
prairie region of the Mississippi Valley from central Missouri northward,
covering the whole State of Iowa, nearly the whole of Illinois, and the
richest and most densely populated agricultural lands of eastern Kan-
sas, eastern Nebraska, eastern South and North Dakota, Minnesota,
and southern Wisconsin.

Specimens examined. — Total number 116, from the following localities:

North Dakota: Portland, 18; Erie, 3; Oasselton,2; Buflalo, 2; Valley
City, 3.

*Dr. C. E. McChesney, U. S. Army, iu a paper on the Mammals of Fort Sisseton,
Daliota, has recorded a valuable series of measurements of this species, all taken at
that locality. While his measurements are not strictly commensurate with ours, and
while many of his specimens were not full grown, his means are important, particu-
larly as showing the average sexual diiference. Reduced to millimeters his most
important means are :

Mea,n of—

Tliirty-tliree males, Fort Si.sset.on, South Dakota
TJiirty five female.s, Fort Si.s.setoii, South Dakota
Sixty-oight specimeu.s, both sexe.s

Head and
body

Tail ver-
tebrai .

Hind
foot.

214
182
198

79

72.5

76

35
32
33

(Bull. U. S. Geol. andGeog. Survey, Terr, iv, No. 1, Feb., 1878, p. 213.)

JAN., 1895] HISTORY OF GEOMYS BUESARIUS. 123

Soiitb. Dakota: Flandreau, 1; Fort Sissetou, 1; Travare, 2; Scot-
land, 1.

Minnesota: Ortouville, 2; Browns Valley, 1; Elk River, 39.

Iowa: Council Bluffs, 1; Kuoxville, 16.

J^ebraska: Xiobrara, 3; Verdigris,!; Columbus, 1; Ames,!; Blair,
1; Norfolk, 2.

Kansas: Onaga, 3.

Missouri: Hunter, Carter County, 4; Williamsville, Wayne
County, 8.

EARLY HISTORY OF GEOMYS BURSARIUS.

The early history of this gopher is somewhat obscure. It was origi-
nally described by Shaw m the year 1800 and was named Mm bursa
rins. * The description is very brief and is as follows : " Ash-coloured
rat, with short round nearly naked tail, pouched cheeks, and the claws
of the forefeet very large, formed for burrowing in the ground." Shaw
states further: " This quadruped was taken by some Indian hunters in
the upper parts of interior Canada, and sent down to Quebec. It is
now in the possession of Governor Prescot." The description is accom-
panied by a full-size engraving of the animal, with cheek pouches
turned inside out and distended. The skin evidently was greatly
overstuffed. No grooves are shown on the upper incisors.

The next year (1801) Shaw redescribed the same specimen as follows:
" It is about the size of a brown or Norway rat, and is of a pale greyish-
brown colour, rather lighter beneath; the length to the tail is about 9
inches, and that of the tail, which is but slightly covered with hair,
about 2 inches: the legs are short; the fore feet strong, and well adapted
fov burrowing in the ground, having five claws, of which the three mid-
dle ones are very large and long; the interior much smaller, and the
exterior very small, with a large tubercle or elbow beneath it. Tlie
claws on the hind feet are comparatively very small, but the two mid-
dle are larger than the rest, and the interior one is scarce visible: the
teeth are extremely strong, particularly the lower i)air, which are much
longer than the upper: the ears are very small." (General Zoology,
vol. IT, part 1, Mammalia, 1801, pp. 100-101.) He gave a new engrav-
ing of the animal, stating that in the figure previously published (in
the Transactions of the Linneau Society) "the claws on the fore feet
are represented as only three in number, and are somewhat too long,
weak, and curved. The engraving in the present plate is a more faith-
ful representation, and is accompanied by an outline of the head, in its
natural size, as viewed in front, in order to shew the teeth and cheek-
IKHU'lies." This plate contains three figures: a side view, as in the
earlier engraving; a front view, reduced, and a natural-size front view
in outline. The cheek pouches are everted, as before, jirotruding from

"Transactions of the Linnean Society, London, vol. V, 1800, pp. 227-228; descrip-
tion read before the society June 4, 1799.

124 NORTH AMERICAN FAUNA. [no. 8.

the sides of the face as great bursie. Although the teeth are distinctly
shown in these engravings, no trace of a groove is apparent, unless an
incomplete dotted line near the middle of each upper incisor in the out-
line figure was intended to indicate it. The size of the incisors in this
figure agrees exactly with the size of these teeth in specimens of Tho-
momys talpoides from Manitoba, and the size and shape of the fore feet
and claws are as in Thotnomi/s, thus dittering widely from the same
parts in Geomys, in which the teeth and claws are very much larger and
heavier.

The color of the body (which he describes as " pale greyish-brown,
rather lighter beneath "), the size and shape of the forefeet and claws,
the size of the incisors, the absence of the deep median furrow so con-
spicuous in Geomys (which could hardly have been overlooked both in
the description and figure), together with the statement that the animal
came from the interior of Canada, all indicate that the species now
known as Thomomys talpoides was the animal Shaw had before him.

The only point mentioned by Shaw in either of his descriptions of
the type specimen of ^ Mus hursarim^ that does not apply strictly to
Thomomys, to the exclusion of Geomys, is the length of the animal,
which he gives as 9 inches. This is easily explained on turning to the
figure, which shows the specimen to be greatly overstuffed— a common
error in taxidermy resulting from the exceedingly loose and distensible
skins of these animals, which are nearly always stretched in taking off
from the body.

Eecapitulating, Shaw's description and figures seem to establish the
following points:

(1) The type specimen of 31us bursarius came from the "upper parts
of interior Canada," the home of Thomomys. No member of the genus
Geomys reaches Canada, its northernmost known point being Warren,
Minnesota.

(2) The type specimen of i¥<6s bursarius was "ash coloured" or "pale
greyish-brown, rather lighter beneath," exactly as in Thomomys. The
color of the only species of Geomys inhabiting the Upper Mississippi
Valley is dark chestnut or liver-brown, both above and beloAv.

(3) The detailed description given by Shaw in his General Zoology
makes no mention of grooves in the incisors, though these teeth are
described with particularity. In Geomys the upper incisors are deeply
furrowed; in Thomomys they are plane.

(4) Not one of the four figures of the type specimen of Mus bursa-
rius published by Shaw shows any trace of the grooved incisors of
Geomys, and two of these figures are front views, one natural size.

(5) The size of the teeth, fore feet, and claws in Shaw's natural-size
figure agree with these parts in Manitoba specimens of Thomomys and
are very much smaller than in Geomys.

From the above facts it would appear that the animal described by
Shaw under the nameilf«.s- bursarius is the gray pocket gopher of Man-

JAN., 1893] HISTORY OF GEOMYS BURSARIUS. 125

itoba aucl the Dakotas {Thomomys talpoides of recent authors) aud not
the red pocket gopher of the Mississippi Valley ( Geomys bursarius of
recent authors). This view would necessitate a slight change in nomen-
clature: Tli<)m(>)iiys taljwides Auct. would become Thomomys bursarius
(Shaw), aud Geomys bursarius Auct. would become either Geomys fuscus
(Eafinesque) 1817, or Geomys sacc((fus (Mitchill) 1821.* Fortunately no
change in the generic name would be required, since Kafinesque based
his genuL,> Geomys on G.pinetis [=■ G. tuza] of the pine barrens of Georgia.

Clear as the case seems to have been left by Shaw, it became shrouded
in obscurity by the writings of subsequent authors.

In 1820 Heinrich Kuhl published his Beitriige zur Zoologie, in which
he described the genus Saccophorus, basing it on the Mus bursaritts of
Shaw. lie states that the specimen examined by him was formerly
in Bullock's Museum, but then in Paris (''in Museo Bullokiano, nunc
Parisieiisi," p. 6.")), but does not intimate that it was Shaw's specimen.
In the diagnosis of the genus he states that the upper incisors have
two sulci, of which the external is broader and deeper, thus describing
the condition in typical Geomys.

Lichtenstein, in a paper written in 1822, but not published until 1825,
says: '• When I was in London in the summer of 1819 I saw in the
Bullock collection the specimen described by Shaw" (UberiiussereBack-
entaschen an Nagethiereii, Abh. K. Akad. Wiss. Berlin [for 1822], 1825,
14-15). He then goes on to describe another specimen, assumed to
belong to the same species, which he says he had recently received
from North America.

The iirst positive statement I have been able to find to the eftect that
Shaw's specimen had grooved incisors was made by liichardson more
than a cpiarter of a century after the publication of Shaw's last descrip-
tion. Richardson states that the engraving of Shaw's Mus bursarius
published in the Linnean Transactions was drawn by MaJ. Davies,t
and that ''the specimen figured by Major Davies, in the Linnean Trans-
actions, was of a pale gray colour, and 9i inches long from the nose to
the root of the tail, which measured 2i inches. The belly was ])aler
than the back, and the cheek-pouches were covered with very short
pale hairs. Its superior incisors were deeply grooved in the middle,
aud more faintly close to their inner margins" {Ibid., 203). As to the
final disposition of this specimen he says: "The identical specimen

* Diplostoma fitsca Kafinesque, Am. Monthly Mag. ii, 1817, 45, is little more than
a noiiien nudum, the only specific description being "entirely brown, length 12
inches." But the generic (liagno.si.s, though full of errors, leaves no doubt as to the
animal; aud the locality assigned, "Missouri Territory," is sufficiently exact in con-
nection with the size and color of the species. If, however, this name is not con-
sidered available, the next in point of date seems to be Mus saccaius Mitchill, Medi-
cal Repository, vol. vi, 1821, 248-250; type "from the region bordering on Lake
Superior," doubtless Minnesota, where the animal is abundant. The bisulcate upper
incisors are described in detail by Mitchill.

t Fau^ia Boreuli- Americana, 1829, 199.

126 NORTH AMERICAN FAUNA. [no. 8.

described by Shaw, * * * on the dispersion of Mr. Bullock's col-
lection, passed into the hands of M. Teniniinck" {Ibid., p. 199).

That this x^ai'ticular specimen is now in the Leiden Museum is
certain, for it is mentioned by Dr. F. A. Jentink, the able director of
the Eijks Museum, in his Catalogue Systematique des Mammiferes, xii,
1888, J). 93. In response to a letter of inquiry. Dr. Jentink has had
the kindness to write me as follows: "On the underside of the stand
[of the si)ecimeii above mentioned] I see the following words written
with pencil: 'Mus bursarius, Cabinet Bullock, Londres.' So you may
be sure of the fact that this specimen truly has been bought from Bul-
lock's auction. As to the animal itself and its identity with Shaw's
description, you may judge if 1 tell yoTi that it has the cheek pouches
turned inside out and distended, but not in the extraordinary way as
represented in Shaw's figure 138, vol. ii, p. 1. The incisors are deeply
grooved. Shaw's figure represents, without doubt, an overstuffed speci-
men; meanwhile our specimen seems to be in excellent proportions and
very well-preserved condition. Length of the animal, 9.8 inches, meas-
ured from the upi3er lip along the dorsal line of the l)ody ; tail about
2.8 inches. The color of our specimen is a desert color, more reddish
toward head and hinder j)arts of the body."

In 1857 Baird made the following statement, evidently based partly
on the remarks of Richardson, already quoted, and partly on an erro-
neous translation of the statements of Kuhl and Lichtenstein. Baird
says: '-The same skin referred to by Shaw was subsequently investi-
gated by Kuhl, and then by Lichtenstein. It was for a time in the
celebrated museum of Mi-. Bullock, of London, and is said to have
been purchased by Temminck at the sale of this collection, and is doubt-
less now in the Leyden Museum." (Mammals of North America, 1857,
370.) But Kuhl does not say that his specimen was the same as Shaw's,
and Lichtenstein distinctly states that the animal described by him was
not Shaw's specimen (which he says he saw in London in 1819), but one
that he " received a short time ago with other North American mam-
mals."

If it is true that the specimen described by Kuhl is really the same as
that described by Shaw twenty years earlier, and afterwards mentioned
by Lichtenstein as having been seen by him in London in 1819, it would
be certain that no other animal thfin the furrowed-toothed pocket gopher
of the Upper Mississippi Valley {Geomy.s) could be meant. But unfor-
tunately Kuhl says nothing on this point, and it must be admitted that
the conspicuous discrepancies between his description and Shaw's are
hard to reconcile on the assumption that they refer to the same speci-
men. Sliaw says the body (jf his animal as stuffed measured 9 inches,
and the tail 2 inches. Kuhl says the body measured 7^ inches, and tail
2^ inches. Shaw described his animal as "ash-coloured," and "pale
greyish-brown," while Kuhl says that his inclined to rufous ("rufes-
cens ") — the proper color for Oeomys.

JAN., 1895.] GEOMYS LUTESCENS. 127

Is it not i)0ssible that Uicbiu'dsoii, in translating" the Latin of Knbl
or the (Jerman of Lichtenstein, fell into the same error as Baird? At
all events it .should not be forgotten that liichardsoii wrote nearly
thirty years later than Shaw — an interval sufficiently long -to allow
additional specimens to reach England and also to favor slips of mem-
ory. It should be further remembered not only that Lichtenstein had
a specimen additional to that described by kShaw, but also (and much
more important) that there a])j)ears to be no ground for the assump-
tion that Kuhl's description was taken from Shaw's specimen; in fact
the marked discrepancies between them seem to jirove the contrary, as
pointed out above.

Shaw\s and Richardson's descriptions are utt^irly irreconcilable on
the assumption that they refer to the same specimen, but would be per-
fectly intelligible if it can be shown that a second specimen found its
way into the Bullock collection between the years 1800 and 1819.

The matter is still further complicated by Richardson himself, who,
writing in 1831, says: " We lately received several specimens of the
Mus burftai'liis of Shaw (which is a true Gcomys^ with pouches oiiening
internally) from the banks of the Saskatchewan." (Zoology of Beechey's
Voyage of the Blossom, 1839, 9,) This statement shows that Richard-
son's ideas respecting- the status and distribution of the several mem-
bers of the group were badly confused, for it is now well known (as
before stated) that no species of Geomys reaches the plains of the Sas-
katchewan ; indeed the genus has not been found to enter Canada at
all. The use of the generic name Geomys by Richardson, however, has
no significance, since he applied the name to Thomomys as well as
Geomys^ and it is certain that his Saskatchewan animal is Thomomys
talpoides Auct. His identification of the species with Mus bursarius
of Shaw would be in accord with my belief that Shaw's animal could
have been no other than the common Thomomys of Manitoba and the
northern plains generally, except for his previous statement, already
quoted from Fauna Boreali-Americana, that the Bullock specimen had
grooved incisors and was the identical specimen described by Shaw.
These conflicting statements by the same author I am utterly unable
to reconcile.

GEOMYS LUTESCENS Merriam.
(PL 9, figs. 5 and 7; pi. 14, fig. 14.)
GeomyKburmrins lutescens Merriam, N. Am. Fauna, No. 4, Oct. 8, 1890, .51.

Tyjye locality. — Sand hills on Biedwood Creek, Lincoln County,
Western Nebraska. (Type in U. S. National Museum.)

Geoymphic distribution.— The Upper Sonoran belt of the Great Plains
from southwestern South Dakota southward to Colorado, Texas, cover-
ing the sand-hill region of western Nebraska, extreme eastern Wyom-
ing (between the North Platte and Cheyenne rivers) western Kansas^

128 NORTH AMERICAN FAUNA. [no.8.

■eastern Colorado, western Oklahoma, and western Texas, ranging east
to or a little beyond the ninety-ninth meridian (map 4, C).

General characters. — Size medium or rather large; coloration jjale;
tail moderate; scant haired; skull short.

Color, — Upper parts in winter drab, liberally mixed with black-tip-
ped hairs along the median line, forming a distinct dorsal band from
end of nose to rump; in summer pale buffy-ochraceous or very pale dull
fnlvons without dorsal band. Under parts buffy, usually white in the
young and sometimes white in adults. Along the eastern and south-
ern limits of its range the upper parts are decidedly more fulvous than
in the typical animal.

Cranial characters. — Skull intermediate in size between hrerlceps and
bursarius; zygomata broadly and squarely spreading, strongly diver-
gent anteriorly; nasals normally elongate w^edge-shaped, as in hursarius,
but sometimes broadening in posterior third ; tem])oral impressions nor-
mally uniting, at least posteriorly, in a low sagittal ridge (pi. 9, tig. 7),
but sometimes remaining apart, separated by an interspace 1 to 3 mm.
broad (pi. 9, tig. 5) [this form is connnonest in the southwestern part of
the range of the species] ; interparietal varying from subquadrate in
the young to subtriaugular in adults, its size decreasing with age and
the posterior suture becoming obliterated by ankylosis with the supra-
occipital; palatoi)tery golds usually lingniate and tapering posteriorly
as in hvrsarins, but somewhat narrower and sometimes strap-shaped.

Skulls of Geomys late^scens differ from those of G. hursarius chiefly in
smaller size, greater relative breadth and flatness (the braincase as well
as the rostrum being considerably shorter than in true hursarins from the
Mississippi Valley), and in lacking the high sagittal crest of hursarius.
Old skulls of lutcsccns have strongly spreading zygomatic arches which
are very much broader anteriorly than posteriorly, and as a rule the
premaxilla extends a little further back than in hursarius.

Sknlls of lutrscens bear a strong resemblance to those of breviceps,
from wdiich they difl'er in having the frontal region less depressed; the
zygomatic arches more squarely spreading and more decidedly angular
anteriorly; the nasal Ijones broader posteriorly; the ascending branches
of the premaxilla longer and less blouutly rounded posteriorly; the
temporal impressions normally meeting posteriorly in a low sagittal
ridge instead of remaining distant; the occiput more truncate (less
bulging) j)osteriorly ; the rostrum normally broader.

The cranial characters that distinguish lutescens from texensis, aren-
arius, und 2)er so nat us are mentioned under tbe heads of these species.

Measurements. — Average of 28 specimens of both sexes from western
Nebraska: Total length, 256; tail vertebrae, 77; hind foot, 32. Aver-
age of 12 males: Total length, 270.5; tail vertebr;e, 84; hind foot, 33.5.
Average of 10 females: Total length, 210; tail vertebrae, 72; hind foot,
31.5.

For cranial measurements, see Table A, p. 204.

JAN., 1895] GEOMYS BREVICEPS. 129

Specimens examined. — Total number of tyj)icalor nearly tyj)ical speci-
mens 118, from the following localities:

South Dakota: Pine Ridge Agency, 2; llosebud Agency, 3.

Il^ebraska: Chadron, 1; Kennedy, ISj Valentine, 3; E wing, 2; Oak-
dale, 2; Crawford, 1 ; Snake River, Cherry County, 1; Chirks Canyon,
Cherry County, 7; Dismal River, Thomas County, 1; Xiobrara River,
Sheridan County, 1; near North Platte, Lincoln County, 4; Birdwood
Creek, 1; Myrtle, 3; Sidney, 1; Calloway. 4- Kearney, 1.

Wyoming: Lusk, 3; Uva, 1.

Colorado: Las Animas, G; Denver, 1; Pueblo, 4 j Limon, 3; Burling-
ton,!; Chivington, G.

Kansas: Trego County, 3.

Oklahoma: Woodward, 3.

Texas: Canadian, 5; Tascosa, 4; Newlin, 3; Childress, 12; Vernon,
9; Colorado, 3.

Number of non-typical specimens 18, from the following localities:

Kansas: Garden Plain, 4; Belle Plain, 5; Cairo, 6; Kiowa, 2; Ellis,!.

General remarks. — Geomys lutescens is a pallid species inhabiting the
arid plains west of the ninety-ninth meridian. Its characters are very
constant throughout most of its range, and if it intergrades with bur-
sariiis it must do so in the narrow strip between the ninety-eighth and
ninety-ninth meridians. In southeastern Kansas an aberrant form
exists that seems to be an intergrade between the three types, hursarius,
lutescens and brevicepSj but a larger series of specimens than at present
available is needed to prove it. This animal is smaller than httescens,
nearly as dark above as hursarius, and paler below than either. Some
specimens indeed have the belly pure white, as in texensis. Specimens
of this ajiparently intermediate form (mostly immature) have been
examined from Cairo, Kiowa, Garden Plain, and Belle Plain, Kansas.

Mr. Vernon Bailey states that in western Nebraska, where typical
lutescens is abundant, the light sandy soil is probably improved by their
diggings, but that they do considerable damage in grain fields and to
young trees on the tree claims.

GEOMYS BREVICEPS Baird.
(PI. 9, fig. 6.)
Geomys hrevxceps Baiid, Proc. Acad. Nat. Sci. Phila., vii, April, 1855,335.

Type locality. — Prairie Mer Rouge, Morehouse Parish,
Louisiana.

Geographic distribution. — The alluvial lowlands of the Mississippi
Valley and Gulf coast in southern Arkansas, Louisiana, and Texas, and
the valley of the Arkansas River; north nearly to southern Kansas,
and west to near the ninety-eighth meridian, where it is replaced by G.
lutescens. It is therefore a memberof the Austroriparian fauna (maplD).

General characters. — Size small; color very dark both above and
below; tail of medium length, its distal half nearly naked.
7433^.^No. 8 9

130 NORTH AMERICAN FAUNA. [Na8.

Color. — Upper parts dark russet brown, darkest along the middle of
the back (but no trace of dorsal band in Louisiana specimens) ; nose
and front of face to above eyes dusky, more or less tinged with rus.^et;
sides Avashed with pale fulvous; belly dark plumbeous, more or less
obscured by pale bufify-fulvous tips to the hairs; feet and throat white;
hairs on base of tail dusky (remainder of tail practically naked). The
color of the back is hard to describe, aud the term used (' russet-brown ')
is intended only as roughly indicating the general effect. The indi-
vidual hairs are dark plumbeous, with a narrow subapical zone of dark
fulvous, tipped with sooty.

Cranial characters. — Skull similar to G. lutescens in general appear,
ance but smaller; zygomata broadly s[)roading; frontal flat, depressed;
nasals narrow, emarginate posteriorly, their sides nearly parallel for
posterior two-tliirds, abruptly divergent anteriorly : ascending branches
of premaxilla broad and bluntly rounded posteriorly; interparietal
small, very irregular, and much cut up with tortuous windings of the
sutures as in true 'Wormian' bones; temporal impressions never
uniting in a sagittal crest but permanently distant, the interspace
elevated, forming abroad convex band (3 to 5 mm. in width) along the
top of the skull posteriori}-; jugal longer than basioccipital, bluntly
rounded anteriorly; occiput bulging behind lambdoid suture, but not
so far as in fcxensis; i)tery golds narrow, tapering posteriorly.

Skulls of ?;>Tr/c('j9.S' may be distinguished from those of lutescens by
the following characters (pi. 9, fig. G) : Size smaller; nasals narrower,
shorter, and strongly emarginate posteriorly; ascending branches of
premaxilla normally shorter and more bluntly rounded iwsteriorly;
temporal impressions persistent, distant, the bone thickened between
them; iaterparietal 'Wormian 'like; zygomata more rounded; inter-
orbital region more depressed. Nevertheless, the cranial resemblances
are striking in view of the dissimilarity of the animals in size aud
external appearance. Moreover, skulls of hreviceps from the western
part of its range have broader nasals ; and skulls of lutescens from adja-
cent territory have a narrow sagittal area (resulting from permanently
distant temporal impressions). It is probable, therefore, that the two
forms will be found to intergrade.

Skulls of hreviceps differ from those of texensis in larger size, much
more spreading zygomata; longer and very much narrower nasals;
broader, flatter, and more depressed frontal interorbitally; much longer
jugal; smaller and more irregular interparietal; less bulging occiput;
broader and more bluntly rounded ends to ascending branches of pre-
maxdla. Viewed in profile, the skull of^hrericeps is flat and somewhat
depressed or concave between the orbits; that of texensis is normally
convex.

Average measurements of -40 specimens of both sexes from type locality
(Mer Rouge, Louisiana): Total length, 219; tail vertebra^, 64; hind
foot, 27. Average of 15 males from same place: Total length, 231; tail

I

JAN., 1895] GEOMYS BREVICEPS. 131

vertebrae, 70; hind foot, 28. Average of 23 females from same place:
Total leugtli 212; tail vertebra, (51; hind foot, 2G.5.

For cranial measurements see Table A, p. 205.

General remarhs. — Tlie type form of Geoniys hreviceps inhabits
northern Louisiana, east of the Red River, the exact type locality
being" Prairie Mer Rouge in Morehouse Parish, near the northern
boundary of the State and only a short distance west of the Mississippi
River. The species as a Avhole is an inhabitant of the dark alluvial
soils of the lowlands bordering the Lower Mississippi and its tributa-
ries and the Gulf coast of Texas, whence it spreads westward nearly or
quite to the ninety-eighth meridian. To the southward it reaches
Nueces Bay. On the west it probably iutergrades with texensis and
lutescens. On the north there seems to be a hiatus between its range
and that of bursar ins; but if pocket gophers are ever found in northern
Arkansas, southwestern Missouri, southeastern Kansas, or north-
eastern Indian Territory, they are likely to prove intergrades.

Departures from the type. — Specimens from extreme points in the
range of the species differ much from the type. Two of these forms
are here named as subspecies {G. hreviceps sagittalis and G. hreviceps
atticateri). Others are regarded as slightly aberrant forms not merit-
ing recognition by name; others still as intergrades. The following,
contained in the Department of Agriculture collection, seem worthy of
mention :

(1) A large dark form inhabiting the valley of the Arkansas River.
The skulls point toward intergradation with the interior animal.
Specimens from Tulsa and Fort Gibson, Indian Territory, and Fort
Smith, Arkansas, resemble hreviceps in coloration, while those from
Pouca Agency, Indian Territory, are redder, shading strongly toward
lutescens.

(2) A form from the valley of the Red River of the South, along the
boundary between Texas and Indian Territory (specimens from Gaines-
ville, Tex., and from Indian Territory opposite Arthur, Tex.). A small
reddish form resembling hreviceps externally, but with dark belly and
a short tail. The skulls are more like texensis in general form (full
brain case and narrow zygomata), and in the shortness and breadth
of the nasals; but the ascending arms of the premaxilla are even
shorter and more bluntly rounded posteriorly than in hreviceps. The
frontal and interparietal are intermediate between the two,* Regarded
as an intergrade.

(3) A form from Slireveport, Louisiana. Much redder than true
oreviceps, resembling texensis in coloration of upper parts, but with
dark beily. The skull differs from typical hreviceps in more angular
zygomata, broader nasals, and less depressed frontal. Regarded as a
slight local departure from hreviceps.

* Skull No. 47590 <? ad. from Gainesville, Texas, is an excellent example of this
form.

132 NORTH AMERICAN FAUNA.

[NO. 8.

(4) A foi'iii fVoiu Galvestou Bay, Texas (specimens from Clear Creek
abd Arcadia), A small, dark, llighly-colored form with the head nearly
black, and the throat and fore feet ilsnally Avholly or partly white, in
sharj) contrast with the dark of the surrounding parts. The skull differs
from that of typical brevic&ps in smaller size, and in having shorter and
broader nasals, llegarded as a subspecies and described under the
name mgittalis. (PI. 9, fig. 4.)

(5) A fbt'm from thft Coastal plane of Texas (si)ecimens from Brenham,
Milano, Hearne, Marquez, and Palestine.) Usually has a well-marked
dark dorsal band, and the skulls differ from typical hreviceps in having
shorter and broader liasals. Skulls of old males from these localities
are unusually short and have broadly spreading zygomata. The nasals
are very broad posteriorly in comparison with true hreviceps. Eegarded
as an aberrant form, perhaps shading toward texensis on one side and
toward atticateri and saf/ittalh on the other.

(()) A form from the extreme southern limit of the range of the spe-
cies on and near the Gulf coast of Texas, (Specimens from Eockport,
Aransas County; Tallj^s Island, Aransas County, and near San
Antonio.) A large dark form with a dark dorsal band in some pelages,
and peculiar cranial characters: angular and strongly divergent zygo-
mata, very broad ascending arms of premaxilla, and so on. Eegarded
as a subspecies, and described under the name attwateri (pi. 9, fig. 3).

Specimens examined,— Total number, 274, from the following locali-
ties :

Typical or nearly typical. — Mer Eouge, Morehouse Parish, Louisiana
(type locality), 42; Pineville, Eapides Parish, Louisiana, 2; Provencal,
Natchitoches Parish, Louisiana, 4 ; Shreveport, Caddo Parish, Louisi-
ana, 8; Camden, Ouachita County, Arkansas, 1; Benton, Arkansas, 7;
Fort Smith, Arkansas, 7; Fort Gibson, Indian Territory, 16; Tulsa,
Indian Territory, 2.

Not typical. — Gainesville, Cook County, Texas, 5; Decatur, Texas,
1; Indian Territory, near mouth of Boggy Eiver (opposite Arthur,
Texas), 4; Pouca xVgency, Oklahoma, 0; Oklahoma City, Oklahoma, 3.
The following, all from Texas: Longview, 4; Mineola, 14; Terrell, 7;
Troup, 1; Palestine, 5; Marquez, 5; Hearne, 9; Milano, 12; Brenham, 7;
Victoria, 1; Inez, 3; Navidad Eiver, 1; Houston, 9; Matagorda Bay, 9.

Subspecies sagittal is. — INIouth of Clear Creek, Galveston Bay, 4;
Arcadia, Galveston Bay, 22.

Snbsjwcies atticateri. — Eockport, Aransas County, 40; Tallys Island,
Aransas County, 3; Calaveras, Wilson County, 3; San Antonio (18
miles south), Bexar County, 7.

Mr. Vernon Bailey, chief field naturalist of the Division, visited the
type locality of Geomys brcvicejys, Prairie jMer Eouge, Morehouse Parish,
Louisiana, in June, 1892, for the purpose of obtaining a series of duplicate
types of the species. He found it common throughout the fields of the
open country and along roads and fields in the woods of the flat land,

JAN., 1895.]

GiPiOMVg feREViCEPg. 133

except wliere flooded, but uot in standing- timber or on hilly land. He
states: "They do uot seem to be so common in cultivated land as in
pastures and along fences and roadways. In one pasture of 20 acres
we caught fifteen and one remained. They were more abundant at this
point than elsewhere — probably twice as numerous to the area as they
would average over the whole prairie. The damage done in tlie pasture
by covering grass was trifling. This species does not seem to dig-
extensively, and the hills are small. Usually one or two are thrown
up in a night. In one place, where a gopher had run his tunnel in a
straight course, I counted sixteen hills in a line (> rods long (measured).
A hill of average size measured 24 by 15 inches in diameter and 5
inches in height. Probably the reason the gophers do not dig more
extensively is that food is abundant and the soil compact. The greatest
damage the tiirmers claim from gophers, or 'salamanders' as they are
called here, is that they carry the tubers of the troublesome cocoa or
nut grass from place to place, often bringing them from a roadside or
waste place and storing a large quantity in their burrows m gardens or
fields and leaving them to grow where they had been kept out with great
difticulty. This cocoa grass is one of the worst plants with which the
farmers are troubled and is very difficult to get rid of when once started
in the land. Small tubers are borne along the roots, and these are
carried by the gophers, though I have not found them in their pockets.
The stomachs examined contained green vegetable matter. White
clover seems to be a favorite food. Most of the specimens taken were
moderately fat. In June the young were half grown to nearly full
grown. Of 27 specimens which I examined, 12 were males and 15
females.''*

Mr. C. L. Newman writes me that at Camden, Arkansas, this sj)ecies
(specimen received for identification) is abundant in sections of the
Ouachita River Valley, where they are known almost exclusively as
'salamanders.' He says: "They seem to prefer old fields that have
grown up in pine. I know of a place about a mile from Camden where
the surface of about an acre of ground is mulched with loose earth
brought from their burrows. Last year (1893) I caught twenty-tliree
from about 6 acres of ground."

* Mr. Veruoii Bailey coutribiites the following notes ou a specimen examined in
the flesh at Mer Ronge, La., in June, 1892 : " Size small ; pelage very soft and silky ;
skin loose, as though much too large for the hody ; body soft and flabby; soles of
feet, nose, and end of tail hairless, smooth, shining, and Avhite Avhen clean. Lips
hairy over the edges, but roof of mouth not hairy all the way across, a narrow line
of smooth skin extending along the median line to the incisors; eyes small for a
Gcomyn; cornea relatively large, measuring 3 mm. across, nearly ecjualing diame-
ter of ball; no apparent lid, eye opening 3.5 mm. by 2 nun. (normally), its long
axis parallel to a line drawn from ear to tip of nose; color of eye appearing shiny
black; ears consist of a circular rim 1 mm. high and about 5 mm. in diameter;
opening of meatus 2 by 2.5 mm., slightly elongated vertically; mustache spreading
forward and back; distance from eye to end of nose 21 mm.; from eye to center of
ear, 17 mm."

134 NORTH AMERICAN FAUNA. [no. 8.

GEOMYS BREVICEPS SAGITTALIS subsp. nov.

(PI. 9, tig. 4.)

Ti/jic from Clear Creek, Galveston Bay, Texas. No. MtJir c? f^tl- U. S. Nat.
Museum, Department of Agriculture collection. C!ollected March 28, 1892, by
William Lloyd. (Original number llSl.)

Geographic distHhiition. — Gulf coast of Texas about (ralveston Bay.

General characters. — Similar to brericepn^ but smaller aud more biglily
colored; head very dark; throat aud fore feet i)ure white iu sharj) con-
trast to dark of surrounding- parts. The skull differs in having a dis-
tinct sagittal crest and in other particulars.

('o?or,— Upper parts rich, glossy, russet brown, strongly tinged with
fulvous, becoming dusky along the middle of the back and head (but
no distinct dorsal band) ; entire head and nose very dark, almost black,
but washed in j)laces with fulvous; inside of cheek pouches, chin,
throat (breast also in some specimens), and "fore legs pure white in
sharp contrast. On the upper side of the fore legs the dark color of
the sides reaches down about half way to the wrists and ends abruptly
with a sharp line of demarkation. The under side of the fore legs is
pure white to elbow. The belly varies from whitish, strongly washed
with buffy ochraceous, to fulvous. The Arcadia specimens are not
exactly like those from the mouth of Clea?" Creek.

Cranial characters (type specimen). — Skull similar to that of hreviceps
but smaller; zygomata more divergent anteriorly (in male); nasals
shorter and broader posteriorly, bringing the constriction much nearer
the middle; aiulital bulhe smaller; ascending branches of premaxilla
narrower posteriorly; temporal impressions meeting in a well marked
vSagittal crest in male. In the female the temporal impressions never
meet in a sagittal crest; the brain case is smoothly rounded, and the
interparietal persists as a relatively large bone.

In the Arcadia males the temporal impressions do not meet in a
sagittal crest as m the type.

Measurements (taken in Hesh). — Type: Total length, 225; tail verte-
brte, 70; hind foot, 27.

Average (of 5 males from Arcadia, Galveston County): Total length,
220; tail vertebme, G4; hind foot, 26.

Average (of 15 females from same place): Total length, 190: tail
vertebni?, 54; hind foot, 23.

For cranial measurements, see Table A, ]). 205.

Specimens examined, — Total number 24: 4 from Clear Creek, Galves-
ton Bay, and 20 from Arcadia, Galveston County, Texas.

General remarl's. — To the northwestward sagiftalUs passes into the
coastal plain form already mentioned under the head of G. hreviceps.
Old males of this form sometimes develop remarkably broad skulls.
The broadest skull that I have seen in the restricted genus Geomys is
an old male from Brenham, Wasliington County, Texas (No. 63G12). It
aft'ords the following measurements and ratios: Basal length, 40j

JAN., 1895.] GEOMYS BREVICEPS ATTWATERI. 135

basilar leiigtli ofHeiisel, 37; zygomatic breadth, 28.5, Ratio of zygo-
matic breadtli to basal length, 71; to basilar leugth of Ileusel, 77.

GEOMYS BREVICEPS ATTWATERI * snbsp. uov.

(PL 9, fig. 3.)
Tfipe from Rockpout, Aransas County, Texas. No. 51382 ^ ad. U. S. Nat, Museum,
Departmeut of Agriculture colU'ction Collected November IS, 1802, by H. H.
Keays. (Original No. 3G.)

Geograpliic (listribution. — Coastal plain and islands of Texas between
Matagorda and Nueces bays; penetrates the interior to within a few
miles of San Antonio. The south side of Nueces Bay is the home of
another form [G. i)ersonat'US fullax).

General characters. — Similar to G. br€vice2)s, but larger and less dark
in color; feet and basal third to half of tail moderately well haired
for a Geomys; terminal half to two-thirds of tail nearly naked ; zygomatic
arches angular, strongly divergent anteriorly.

Color. — Upper parts russet brown, becoming dusky on the head and
usually along the median part of the back; under parts varying from
soiled whitish to buflfy ochraceous. In some specimens the color of the
ujiper i)arts is less fulvous than in others, and the dark dorsal band is
variable; iu some specimens it is absent, sometimes the head is nearly
black from end of nose to occii»ut, the blackish area limited laterally
by the eyes and ears, the sides of the face being russet in rather strong
contrast. The type specimen is in this pelage, except on the hinder jiart
of the back and rump where the more fulvous summer pelage remains,
without trace of the dorsal band.

Cranial characters. — Skull similar to that of hreviceps, but frontal
less depressed interorbitally; zygomata less spreading, strongly diver-
gent anteriorly, more angular, more depressed, the maxillary arm slop-
ing strongly backward; ascending branches of premaxilla broader and
usually more abruptly truncate posteriorly; nasals shorter and normally
convex instead of emarginate posteriorly. The nasals are normally so
narrow posteriorly, and the premaxillje so broad, that iu some cases the
latter nearly meet behind the former (as in the type si)ecimen, pi. 9,
fig. 3). Normal skulls of attwateri differ markedly from those of fallax
in the form of the zygomata, the maxillary arms sloping strongly back-
ward instead of standing out at right angle, and the outer sides being
strongly divergent instead of nearly parallel. The nasals are narrower
and contracted posteriorly, the ascending arms of the premaxilla
broader, and the audita! bulhe less swollen. In the series of lifty-two
skulls of Geomys breviceps attwateri now before me, three depart from
the normal in general outline, as seen from above, and resemble /ai^aa;
in the form of the anterior part of the zygomatic arches, wliich stand
out sc^uarely from the cranial axis and have the antero-external angles

* Named in honor of Mr. H. P. Attwater, of San Antonio, Texas, who collected nearly
all of the specimens.

lo6 NORTH AMERICAN FAUNA. [no. 8.

broadly rounded. In other respects tliey are typical attwaterl. All
are very old males, collected at Kockport by Mr. Attwater (original
Nos. 102, 118 and 110), They now belong to the American Museum of
Natural History in 'New York.

Meamirements (taken in flesh). — Type: Total length, 250 ; tail vertebr.ne,
85; hind foot, 30.5,

Average of 10 males from type locality: Total length, 255; tail ver-
tebra, 80 ; hind foot, 30.

Average of 7 females from type locality: Total length, 220; tail ver-
tebrae, 08 ; hind foot, 28.

For cranial measurements see Table A, p. 205,

Specimens examined. — Total number 53, from the following localities
on or near the Gulf coast of Texas: Rockport, Aransas County (type
locality), 40; Tallys Island, Aransas County, 3; Calaveras, Wilson
County, 3; San Antonio (18 miles south), Bexar County, 7.

General remarks. — Geomys hreviceps attivateri is a medium-sized
species closely resembling its near neighbor G.fallax in color, though
somewhat darker, and with the hind foot shorter. The resemblance to
G. hreviceps is much closer in the plumbeous russet pelage than in the
fulvous pelage.

Mr. H. P. Attwater has kindly contributed the following memorandum
respecting the habits of this gopher at Rockport, Texas : "As soon as the
warm weather sets in, from about May to September, very few gophers
are observed working. The soil is sandy, and at all times damp,
dampness known as 'natural subirrigation.' In the hot weather the
dampness does not come as near the surface as in the cooler months.
I have thought that perhaps the gophers travel deeper in summer, but
now think the chief reason why they do not throw up hills in summer,
as they do in fall and winter, is that during the summer months the
soil is so full of roots, suckers, bulbs, etc., that they do not have far to
go before finding all they can eat, and that the reason they work so
nuich after the summer months are over is because they are hunting
around to fiud some bulb or root which was their favorite food in
summer, and which they commenced to find about the month of May,
and was over with in September. The animals are very abundant all
over the i)eninsulas in Aransas County, wherever the soil is sandy.
There is hardly a foot of laud that has not been 'plowed' several times
over by gophers, and I believe the fertility of some sections has been
greatly improved by them, by bringing the i)oorer soil up to the top.
I have noticed that the richer the land the richer the gophers. Of
course they do considerable damage to vegetable crops, especially to
young fruit trees and cuttings just rooting. The samples sent you of
mulberry trees cut by gophers were from the Faulkners' ranch, on St.
Charles j)eninsula, in the eastern part of the county. Mr. Sanmel
Walker, the manager of tlie ranch, told me that he killed over two hun-
dred and fifty gophers in his young pear orchard between the 1st of

JAN. 1805.1 GEOMYS TEXENSIS. 137

March and April 15, 1893. This orchard was set out where sweet
potatoes had grown the year before, and they came np again and cov-
ered the gronnd, and I think the potatoes attracted the gophers in the
first place more than the pear trees."

GEOMYS TEXENSIS sp. nov.

(PL 9, fig. 2; pi. 13, tig. 12.)

Type from Mason, Mason County, Texas. No. J^H 9 ad. Merriam collection. Col-
lected by Rev. Ira B. Henry, December 17, 1885.

Geographic distHbution. — Mason County, central Texas, and prob-
ably thence southerly to the Kio Grande; limits of range unknown
(map 4, E).

General characters. — One of the smallest known species; tail short;
terminal third nearly naked.

Color. — Upper parts liver-brown, finely mixed with black-tipped
hairs, much as m G. bursarius. Under parts and feet while. The hairs
of the belly are plumbeous at base in the type and other winter
specimens; in summer specimens they are white throughout. Throat
suffused Avith pale bufty fulvous, forming a complete collar. In some
specimens this collar is interrupted along the median line. The
color of the upper jiarts is darker in winter than in summer, as
usual in the genus. There is no trace of a dark dorsal band in adults,
but in the young the black-tipped hairs are sometimes concentrated
along the middle of the back, forming an ill defined dark streak.

Cranial characters. — Skull small (smallest of the known species),
smooth ; zygomata only moderately spreading and normally but slightly
divergent anteriorly; nasals short, rather broad and convex or trun-
cate behind; ascending branches of lU'emaxilla long, normally passing
plane of lacrymals, usually straight on inner edge behind nasals and
attenuate on outer edge; temporal imjiressions not forming distinct
ridges and not uniting in a sagittal crest, usually separated by inter-
space 1-3 mm. broad in adults; jugal short (shorter than basioccipital) ;
interparietal broader than long, normally oval or elliptical and project-
ing posteriorly behind plane of lambdoid suture; occiput bulging
posteriorly more than in any other United States species (resembling
Fappogeomys bulleri and some species of Thomomys).

Skulls of texensis differ consi^icuously from those of G. arenarius in
the following points: jSTasal branches of premaxilla longer and more
pointed posteriorly; jugal more slender; no distinct knob at end of
squamosal arm of zygoma; no distinct tem^joral ridges; inter j)arietal
projecting posteriorly behind ijlane of lambdoid suture; occiput more
bnlging posteriorly; mandible less heavy. G. texensis differs from G.
breviceps in the shape of the nasal bones which are usually short, very
broad posteriorly, with the sides nearly parallel. In G. breviceps they
are usually longer, strongly wedge shaped, very narrow posteriorly,
with the anterior third abruptly broader and flaring. In te.vensis the
nasal branches of the premaxilla reach or pass the plane of the orbital

138 NORTH AMERICAN FAUNA. [m.R.

fossa aud are pointed; in hreviceps they usually fall short of this plane
and are bluntly rounded. In texensis the'jugal is shorter than tliehasioc-
cijjhal; in brei^icips it is longer. In hreviceps the outer angle of the
zygomatic arch is evenly rounded ; in texensis it is angular and abruptly
flattened (or even excavated) on its infero-external face, beginning at
the angle and extending posteriorly under the jugal (as seen from the
side). The. inflated mastoids and audital bulhie are larger in hreviceps,
in which species the mastoids are conspicuously broader than in texen-
sis, the exposed part, viewed from behind, being as broad as high, while
in texensis the breadth is only about half the height. But the range
of individual variation is so great that much confidence can not be
placed on this character.* In hreinceps the frontal is flatter and
depressed interorbitally, forming a slight concavity in the plane of the
upper side of the skull when seen in profile; in texensis the profile is
convex at this point.

Skulls of Geomys texensis difter Irom those ot 6^. bursar ins, in addition
to their much smaller size, in shorter rostrum and brain case, less promi-
nent ridges and i^rocesses for muscular attachments, absence of sagittal
and lambdoidal crests fit all ages; much larger iiiter])arietal; uuich
larger andital bulhe (which are inflated and rounded antero-laterally
instead of flattened), and in the greater length of theascending branches
of the premaxilhi posteriorly. Tlie skull as a whole is not only much
smaller than that of hnrsariiis, but is relatively thin and smooth, like
that of Tltomomys. The arch of the brain case is low, but not so flat as
in hreinceps, and the temporal impressions never mret along the median
line.

Measurements. — Type specimen : Total length, 20."} (measured in flesh) ;
hind foot, 28 (in dry skin moistened to straighten the toes). Tail not
measured in flesh, but short; about 00 in dry skin. Average total length
of 28 specimens from tyi)e locality measured in flesh, 210.

For cranial measurements see Table B, p. 200,

Specimens examined. — Total number 31, from the following localities
in Texas: Mason, Mason 0(mnty (type locality), 28; Laredo, 1; Syca-
more Tree (on liio Grande), 1; Del Itio (on Eio Grande), 1.

General remarls. — Geomys texensis is a small white bellied species
inhabiting central Texas. Its back is chestnut-bvown or liver-brown,
much as in the large dark-bellied G. hursarius, with which it requires
no comparison. It is the smallest species in the United States, about
equaling Pappogeoniys hnlleri of Mexico. The only bisulcate species
of approximately the same size are G. hreviceps of Louisiana and its
subspecies saaittaUs of the Gulf coast of Texas, and G. arenarius of
the Upper Rio Grande Valley in extreme western Texas and south-cen-

* The actual size of the mastoid is often hidileu by the thin outer edge of the exoc-
cipital which overlies its iuner border, aud which is not always alike on the two
sides. Hence it sometimes happens that the exposed part of the mastoid is narrow
ou one side and broad ou the other.

jAN.,iR9r,.] GEOMYS ARENARIUS. 139

tral New Mexico, with all of which it may iutergracle, although it differs
widely from them all in color aud cranial characters, as elsewhere
shown. On the north, in Oklahoma and southern Kansas, it i)robably
intergrades with G. lutescens.

Three specimens of a small Geomys from as many points in the Eio
Grande Valley (Laredo, Del Kio, and Sycamore Creek) are provision-
ally referred to the jjresent species. The Laredo >specimen lacks the
skull and its upper parts are more drab than usual. The specimens
from Del Rio and the mouth of Sycamore Creek are too immature for
positive identitication. They differ from the young of texensis from
the type locality in having longer tails, somewhat darker backs, and
in lacking the chestnut tint on the sides. Their skulls seem to be
intermediate between texensis and arenarins. Mr. William Lloyd, who
collected the Sycamore Creek specimen, states that the species is
rare there and was found only in a belt of fine sand along the Rio
Grande. He found a species, presumably the same, on chalky soil near
Comstock. 'Slw Vernon Bailey collected the Del Rio specimen in the
river bottom, where the species was rather rare.

GEOMYS ARENARIITS sp. nov.

(PL 9, fig. 1; pi. 13, fig. 13.)

Type from El Paso, Texas. No. \r.wxl ,i ad. U. S. National Museum, Deiiartmentof
Agriculture collection. Collected Deci'ml)er 13, 1889, by Vernon Bniley (Orig-
iual No. 798).

Geouraphic distribution. — ^' alley of the Upi)er Rio Grande, from El
Paso, in extreme western Texas, and Juarez, Chihuahua (on the Mexi-
can side of the river opposite El Paso), north to Las Cruces, New Mex-
ico, and west to Deming, in the same state (map -1, a). It will prob-
ably l)e found to follow the valley somewhat further in both directions,
and to the east may intergrade with texensis. So far as now known its
range seems to be separated by a broad interval from that of the spe-
cies inhabiting central and southern Texas, the westernmost records of
which are Del Rio and Comstock, in the Rio Grande Valley. Curiously
enough the intervening region is inhabited by a widely different Pocket
Gopher, one belonging to the unisnlcate series, namely, Cratof/eomi/s
castanops. The ranges of all the other bisulcate species, except /«://,
are either directly continuous or contiguous. In faunal position G.
arenarins belongs to the upper edge of the Lower Sonoran Zone.

General characters. — Size medium; tail rather long and unusually
well haired, except near tip; coloration pale.

Color. — Upper parts drab-brown, finely mixed with black-ti])ped
hairs; under parts and feet white. In some specimens the color of the
sides encroaches on the belly and is only partly masked by the white
tips of the hairs.

Cranial characters. — Skull resembling Thomomys talpoides; size rather
small (intermediate between texensis ^nd hreviceps) ; zygomata normally

140 NORTH AMERICAN FAUNA. [no 8.

narrow and nearly parallel (in one S from El Paso, No. 58340, they are
exceptionally divergent anteriorly) ; no sagittal crest at any age; tem-
poral ridges prominent, distant, and nearly parallel or slightly diver-
gent anteriorly, usnally separated by a flat or concave interspace 4 to
5 """ wide, as in Thomomys talpoides ; squamosal arm of zygoma ending
in a prominent knob over middle of jugal (diagnostic of the species);
jngal short (shorter than basioccipital); interparietal rather large,
normally (but not always) broader than long, usually siibquadrangular
or with the corners rounded anteriorly, truncate posteriorly on plane
of lambdoid suture; occiput bulging posteriorly, but not solar as in
texensis; palatopterygoidsnormally abruptly narrow, their sides nearly
parallel (but form somewhat variable); mandible heavy fur size of skull.
The fenmles diifer fr<mi the males in having shorter nasals, larger par-
ietals, and less prominent temporal ridges. As a rule the interspace is
somewhat thickened and the ridge is evideut from the outer side only.

The skull of G. arenarius differs from that of texensis in the follow-
ing characters: Jugal heavier and broader; temporal ridges much
more prominent and distant; a prominent knob at distal end of squa-
mosal arm of zygoma; top of skull flatter; frontal broader and flatter
interorbitally; iuterparietal truncate posteriorly on plane of lambdoid
suture; occiput less bulging. It differs from lutesccns in much smaller
size, narrower and uiore parallel zygomata; shorter jugal; in the
presence of well-developed distant temporal ridges, ami of a prominent
knob at distal end of squamosal arm of zygoma; shorter and somewhat
narrower nasals, and shorter ends of ascending arms of premaxilla
behiud the nasals.

Measureynenis [taken in flesh). — Type specimen ( S ad.): Total length
258; tail vertebrae, 88; hind foot, 33. Average of 8 males from type
locality: Total length, 200; tail vertebr;Te, 83; hind foot, 32. Average
of 24 females* from type locality: Total length, 250; tail vertebne, 78:
hind foot, 32.

For cranial measurements see Table B, p. 207.

Sjyecimens examined. — Total number 43, from the following localities:
Juarez, Mexico, 3; El Paso, Texas, 30; Deming, New Mexico, 3; Las
Cruces New Mexico, 7.

General rcmarlxs. — In color and external appearance Geomys arenarim
closely resembles the typical form of G. Ititescens (from western Ne-
braska and eastern Wyoming), differing chiefly in smaller size and in
greater length and hairiness of tail. From its nearest ally in central
Texas (G. texensis) it differs both in color and proportions, having the
upper parts pale drab instead of reddish brown, and the tail long and
hairy instead of short and nearly naked. In cranial characters it may
be distinguished from all other species by the presence of distant tem-

* Some of tlae si)eciinens recorded as females are very largo and were probably
males; hence the averages here given for females are ijrobably too great.

JAN., 1895.] GEOMYS PERSONATUS. 141

poral ridges or ribs, wliicli are nearly parallel, in connection with the
develoi)inent of a prominent knob at the distal end of the squamosal
arm of the zygoma.

This fine cpecies was discovered by my assistant, Mr. Vernon Bailey,
at El Paso, Texas, in December, 1889, and was obtained by him at
Deming, New Mexico, also. Mr. J. Alden Loring-, who was sent to the
Upper Rio Grande Valley to work out its range, secured a large series
iVom Las Cruces, New Mexico, and Juarez, Chihuahua, Mexico, as well
as at the type locality. El Paso, Texas. Mr. Loring says: "They
are not very common on the Mexican side of the river, but extremely so
on American soil, where they seem to thrive and grow fat. The places
they most prefer are railroad embankments and irrigation ditches,
where they were found both in sand and wet, dark clayey soil. Two
were seen on February 5 just as they protruded their heads from their
holes. Their faces were covered with dirt, and as soon as they had
shaken it ott' they saw me and quickly dodged back. When these
Gophers were caught I noticed that they walked with the claws of the
front feet partially doubled under, which did not aUow the sole of the
foot to touch the ground."

GEOMYS PERSONATUS True.

(PI. 12, fig. 4; pi. 13, fig. 14; pi. 14, fig. 4.)

(ieomijs personatus True, Proc. U. S. National Museum, xi (for 1M88), Jau. 5, 1889,
159-160.

Type locaUty. — Padre Island, Texas.

Geographic distribution. — The Tamaulipan fauna of Texas, comprising
Padre Island and the adjacent mainland southwesterly to Carrizo on
the Rio Grande (map 4, f).

General characters. — Size large; coloration jjale; tail long, scant-
haired on proximal half and nearly naked on distal half.

Color. — Upper parts pale drab (darker in winter from more liberal
admixture of dark-tipped hairs) ; middle of face from nose to above eyes
inclining to dusky. Under parts white, sometimes obscurely clouded,
from the presence of irregular patches of hairs with plumbeous bases,
the hairs on other parts of the belly white to roots. Tail hairs white,
but too far apart to give color to the nearly naked tail.

Cranial characters. — Skull large, heavy, with well-developed proc-
esses and ridges and a high sagittal crest (pi. 12, fig. 4); zygomata
standing out at right angle to axis of skull; jugal bluntly and broadly
rounded anteriorly, and short, not longer than basiocciptal (measured
from condyle); nasals long and narrow, anterior third spreading;
frontal narrow interorbitally, the orbital borders rounded; basioccipital
with sides parallel, or nearly parallel. In profile the top of the skull
(including the sagittal crest) is nearly a straight line.

Adult skulls of Geomijs personatus may be easily distinguished from
those of bursarius and liitescens by the squareness of the zygomatic

I

142 NORTH AMERICAN FAUNA. [no. 8.

arches anteriorly, the shortness of the jugal bone anteriorly, with cor-
responding prodnction of the maxillary arm of the zygoma. The greatest
length of the jngal in personatus is only eijual to the length of the basi-
occipital bone (measured from the condjde). In both hursarius and
lutesccn.s the jugal is much longer than the basioccipital. In personatus
the skull as a whole is relatively as well as actually longer, and nar-
rower across the zygomatic arches, than that of IntcscenSj from which
it differs further in the following particulars: zygomatic breadth
usually less than distance from foramen magnuui to incisive fora
miua (the contrary being usually true in lufescens); ascending branches
of premaxilla extending much further iiosteriorly ; zygomatic arches
relatively long, only moderately spreading anteriorly (except in
extreme age), but standing out at right angle to longitudinal axis of
skull; orbital fossfe elongated antero-posteriorly instead of subtri-
angular; length of frontal along median line usually equal to length of
nasals (ommonly shorter in lutescens) ; audital bulhe longer, with outer
side flattened; inflated mastoid smaller. Skulls of personatus average
longer in proportion to the zygomatic breadth than those of any other
known bisnlcate species, except the Mexican Zygogeomys trichopus
(the ratio of zygomatic breadth to basilar length ranging from 68 to
72 percent), though in this respect they differ but slightly from Geoniys
hursarius.

Measurements. — Of 13 specimens (of Ijoth sexes) from type locality
(Padre Island): Total length, 399; tail vertebr;^, 103; hind foot, 37.
Average of 4 males: Total length, 315; tail vertebrje, 111; hind foot,
40. Average of 9 females: Total length, 293; tail vertebra} 100; hind
foot, 36.

For cranial measurements see Table B, p. 206,

Specimens examined. — Total number 33, from the following localities
on or near the Gulf coast of Texas: Padre Island (type locality), 15;
nenr Santa Kosa, 8; Sauz Kancho, 6; Carrizo, 3.

Number of subspecies /aZ/aj? 22, as follows: Nueces Bay and River
^south side), 6; Corinis Christi, 15; Las Mottes, 1.

Departures from the type. — The type locality of Geomys personatus is
Padre Island. Fairly typical specimens are at hand from points ou
the mainland west of the southern part of this island, namely, Santa
Rosa and the Arroyo Colorado (Sauz Rancho), and also from Carrizo
on the Rio Grande, though the latter depart somewhat from the type.
Singularly enough, specimens from the lower Nueces River and Bay,
and from Corpus Christi and Las Mottes, differ decidedly from the typ-
ical animal in smaller size, darker color, and in important cranial charac
ters. The skull is much smaller, more abruptly truncate posteriorly,
with more spreading zygomatic arches, and much more globular audital
bulla' (pi. 12, fig. 3). This form is here separated sub-specifically under
the name Geomys personatus fa llax (see p. 144). Intergradation between
personatus and/aZ/aa; probably occurs in the narrow strii) between Santa

JAN., 1895] GEOMYS PERSONATUS. 143

Rosa and Corpus Christi Bay, since the single specimen from Las
Mottes, a few miles south of Nueces Bay, is somewhat larger than the
Nueces Bay and Corpus Christi specimens.

Some of the specimens from Santa Rosa are fairly ty])i<ml jJer son at us,
though all have more swollen audital and mastoid bulhe. One adult
skull (No. 42,860) from the Arroyo Colorado (Sauz Rancho, about 50
miles north of Brownsville) has a very narrow rostrum, narrow zygo-
mata, projecting occiput, very much swollen mastoid and audital bulhe
(the latter almost subglobular) and abnormally short and narrow jugal.
Five other skulls from the same locality are young and apparently less
extreme. The adult skull may be regarded as abnormal, or as pointing
to the differentiation of an incipient race.

General remarJcs. — (Jeomys personatus resembles G. lutescens in sum-
mer pelage more closely than any other form. The typical animal may
be distinguished from lutescens at all seasons by larger size, longer feet
and tail, by important cranial characters (just described in detail), and
by the white of the under parts. In summer specimens of G. lutescens
the belly is sometimes pale, but rarely white except in the very young.
The color of the upper parts in summer pelage differs but little in the
two species, being drab in both, with the nose and middle of the fiice,
as far back as the eyes, inclining to dusky; but in winter and early
spring the two differ notably, the dusky face markings of lutescens
extending posteriorly over the head and back to the rump, forming a
distinct dorsal stripe. In this pelage, also, the under parts are much
darker, the fur being dark plumbeous, tipped with drab. While per-
sonatus is the larger of the two animals, the claws of the fore feet are
equally large (and relatively larger) in lutescens. In some specimens of
personatus the claws are remarkably long and slender — the result, doubt-
less, of the unresisting character of the sand in which the animals live.

The geographic distribution of Geomys personat\is (including sub-
species fallax) appears to coincide with the limits of the arid tropical
area of Texas — an area recognized and defined by me in 1892,* and sub-
sequently named the Tamaulipan fauna by Allen.t The range of the
species has been ascertained to terminate abrujitly both on the north
and on the west, specimens from a few miles north of Corpus Christi
Bay, and from Laredo on the Rio Grande, belonging to different species.

Mr. William Lloyd, who collected the si)ecimens, statesthat G. person-
atus IS abundant in a patch of fine sandy soil above (!arrizo, but was
not found elsewhere in the neighborhood. He sti.tcs further that in
traveling north from the mouth of the Rio Grande it was first met on
entering the great sand belt on the north side of the Arroyo Colorado
(at El Sauz), It continued throughout this sand belt, becoming more
abundant to the northward. On Padre Island he found the animals
living in colonies, perhaps a mile or more a]>art, and common from the

"Presidential Address, Proc. Biol. Soc, Washington, April, 1892, p. 33.
t Bull. Am. Museum Nat. Hist., Kew York, Vol. iv, Jan., 1893, 241-242.

144 NORTH AMERICAN FAUNA. [no. 8.

north end to tlio center of the island, but not within 20 miles of the
south end. Mr. Lloyd says: ''Their habits are in some respects pecu-
liar, o\Yin!4' i)erhaps to the soft sand that caves iu on them, or to fear
of the coyotes, or for both reasons; they fill uj) their tunnels for a yard
or two almost immediately after they throw out the dirt. They can
not go very deep iu the flats or they would reach water; in fact, the
water filled some of the tunnels for about a foot until they curved
upward. Not more than one is ever found in a hole. "

GEOMYS PERSONATUS FALLAX 8ul)sp. nov.

(PI. 12 fig. 3.)

Typo from south side of Nueces Bay_. Texas. No. f |?^i <? ad. Collected Novem-
ber 30, 1891; by William Lloyd. (Original No. 949.) •

Geographic distribution. — South shore of Nueces Bay and lower
Nueces Eiver, Texas; further south passing into G. personatus.

General characters. — Similar in external appearance to G. personatus
of Padre Island, but much smaller (only about half the bulk of that
species); somewhat darker; tail shorter and nearly naked.

Color. — Upper parts drab-brown, darker in winter; paler and more
fulvous in summer; nose and face between eyes dusky; sometimes an
ill-defined dusky band along the middle of the back. Under parts
usually marbled with pure white and i^atches of dark hair (the white
hair being white to roots).

Cranial characters. — Skull similar to that of personatus, but very
much smaller (pi. 12, fig. 3). The zygomata stand out squarely at right
angles to axis of cranium and are widely spreading, their outer sides
nearly x^arallel; the temporal impressions meet in the males in a well-
marked sagittal crest; in the females they remain apart, separated by
an interspace about 3 millimeters wide; nasals rather broad and blunt
posteriorly; jugals short (not longer than basioccipital) ; mastoid and
audita! bulhe swollen, the latter short and rounded; palatoptery golds
narrow, their sides nearly parallel. Skulls of fall ax differ from those
of personatus in very much smaller size, shorter (and usually blunter)
ascending arms of premaxilla, more squarely truncate occiput (lamb-
doid crest less convex posteriorly), and in much shorter and more
swollen audital bulhc.

Geomys personatus /aZ/aa? differs markedly from G.attivateri (which
it approaches in size) in the form of the zygomata, the maxillary arm
standing out at right angle instead of sloping strongly backward, and
the outer sides of the arches being nearly parallel instead of strongly
divergent anteriorly. It differs farther in having more globular audital
bullae, broader nasals, narrower ascending branches of the i)remaxilla,
and in the males a well-developed sagittal crest instead of permanent
temporal ridges.

Measurements. — Type specimen : Total length, 250; tail vertebrai,80;
hind foot, 35. Average of 9 males from south side of Nueces Bay : Total

JAN., 1895.1 GENUS PAPPOGEOMYS. 145

length, 263; tail veitebnu, 87; kind foot, 34. Average of 10 females
from same locality: Total length, 236; tail vertebrse, 75; hind foot, 31.

For cranial measurements see Table B, p. 20r).

S2)ecimens examined. — Total number 32, from the following localities
on or near Nueces Bay, Texas: oS^ueces Bay, 4; Nueces River, 10 miles
from mouth, 2; Corpus Christi 15; Las Mottes, 1.

General remarks. — Geomys fallax is a miniature of G. 2>ersonatus,
both in external appearance and in the general form of the skull. It
is hardly more than half the bulk and weight oi personatus^ from which
it differs further in somewhat darker coloration and in cranial details.
The geographic range of the typical form is remarkably restricted,
being limited, so far as known, to the south side of the lower Nueces
Kiver and Bay.

In his notes on mammals observed in southeastern Texas, Mr. William
Llo3'd states that this species "is abundant in all soils, although it
prefeis the black loam. On Nueces Bay they burrow in the sand close
to the water's edge, but are most at heme on the highest point attain-
able. I have seen an unbroken line of hills extending from 70 to 100
yards across patches of early pease and onions. They cause havoc
among the sweet potatoes, coming above ground to eat them in the
daytime. I shot a marsh hawk that was flying oft with a gopher which
had been thus engaged. While driving along the road cats may be
seen frequently a mile from the house intently watching the gophers'
holes. The gophers are known to be great pests to fruit and other
trees; in more than a dozen instances near the bay I have seen the
huisachi [Acacia farnesiana) leveled by their work in chewing the
rootlets and digging the earth away from the roots."

Genns PAPPOGEOMYS * nob.

(PI. 11, lig. 1; and text figs. 56, .57 and 58.)
Type Geomys hiiUcri Thomas, from Talpa, Mascota, Jalisco.

Dental characters. — Upper premolar with three enamel plates, the
posterior absent; m' and m^ with two enamel plates each, as in Geomys.
Last upper molar an imperfectly double prism; a single sulcus on outer
side, behind which the crown is narrowed, forming a moderately well-
defined heel; outer enamel plate bent slightlj' outward near its anterior
end. Upper incisor unisulcate, the sulcus median and deep (no trace
of minor sulcus; see fig. 21^).

Cranial characters. — Skull small, short, rather smoothly rounded ; a
broad sagittal area (no sagittal crest at any age, pi. 11, fig. 1); zygo-
mata slender, rather broadly and squarely spreading, without trace of
angular expansion; occiput bulging posteriorly; palatoi^terygoids little

''Pappof/eomys, from TrdTTTrof, grandfather, + Geomys, in reference to the apparent
antiquity of the type.

7433— No. 8 10

14G

NORTH AMERICAN FAUNA.

[NO. 8.

more tlian vertical lamelliv, slightly everted iiiferiorly ; orbital plates of
frontal separated interiorly by full breadth of cribriform plate as in
Tliomomys; orbitosphenoids broad, articulating firmly Avith alisphenoids
and sending a tongue upward to nearly fill the upper x)art of the sx)he-
uoidal fissure; mesetlimoid a nearly vertical plate much higher than
long, its inferior edge dipping down between wings of vomer posteri-
orly; endoturbinals as in PlatygeomySj the first sharply triangular and
the OS planum trimmed closely in front of the others.

Fi(i. 56. — I'appoij'MiHys biiUeri. Vault of cranium sawed off, showing floor of brain case.
(For key see tig. 9).

Fm. 57.—Pappo(jeomt/s bulleri. Vortical longitndiual section of skull, mesetbmoid and vomer
in place. (For key see fig. 7).

Fig. bS.—Pappogeomys bulleri. Mesothmoid and vomer removed to show endoturbinals.
(For key see fig. 10).

External eharacters. — iH/.e small; pelage soft; form Thomomine.

General remarks. — Pappogcomys holds an interesting position with

eference to the trunk line of the Geomyldw. Tn dental characters it

combines the inolariform enamel pattern of Geomys with the unisulcate

incisors of Cratogeomys and Platy geomys; and in cranial characters it

JAN., 1895] PAPPOGEOMYS BULLERI. 147

exhibits striking resemblances to both Geomys aud Thomomys on the
one hand, aud to Craiogeomys on theotlier. The endoturbinals are not
widely different from the Geomys type, while the orbitosphenoids depart
entirely from Geomys and surpass Craiogeomys in the extent of their
development aud articulations. They cut off" and shorten the sphe-
noid fossa^, which in Geomys reach forward to the orbital plates of the
frontal (pi. 17, fig. 3), The shape of the mesethmoid plate is unique.
The form of the skull as a whole is very like the simpler forms of
Thomomys and Geomys — as texensis aud arenarius — aud the permanently
distant orbital plates of the frontalis a decidedly Thomoiuine character.
The resemblances to Geomys aud Thomomys do not indicate that
Pappogeomys has descended from either of these genera, but that it
occupies a place near the trunk line and below the point from which
they branched off". On the other hand, the resemblances to Cratogeomys
and Platygeomys are prophetic, indicating a position near the base of
the great bram^h that afterward gave rise to these more specialized
types.

KEY TO SPECIES OF rAPPOGEOMYS.

Mastoids small, truncal e above hnUcri.

Mastoids large, rounded above albiiiasus.

PAPPOGEOMYS BULLERI (Thomas),

(PI. 11, fig. 1; pi. 13, tig. 15; pi. 14, tig. 11.)

Geomys huUeri Thomas, Auuals aud Magazine Nat. Hist., 6 series, Vol. x, August, 1892,

p. 196.
(leomijs neJsoni Merriam, Proc. Biol. Soc., Washington, vii, September29, 1892, 164-165.

Type locality. — IsTear Talpa, west slope of Sierra de Mascota,
Jalisco, Mexico (altitude, 8,500 feet). Type in British Museum.

Geographic distribution. — Lower slopes of Sierra Nevada de Oolima
and Sierra de Mascota, Jalisco, Mexico (map 3').

General characters.* — Size smallest of the known unisulcate species, •
of which it is a generalized type; skull small aud smooth, resembling
Thomomys; tail naked; a naked pad on end of nose, partly inclosed in
a pale patch.

Color. — Upper parts rich rusty chestnut; underparts paler. An
innnature but full-grown specimen (Xo. 33585) is dusky in color, aud
one in the molt has the anterior parts chestnut and the posterior dusky.

Cranial characters. — The skull of Pappogeomys bulleri is small and
smoothly rounded, with broadly distant and rather feeble temporal
ridges. The maxillary arms of the zygomata stand out at right angles

" The following description is based wholly on specimens from the north slope of
the Sierra Nevada de .lalisco. They are larger than Thomas's type and only speci-
men of (i. bnlleri, and may prove subspecihcally separable, in which case the name
nehoni will be available.

148 NORTH AMERICAN FAUNA. [no. 8.

to the axis of the skull; the zygomata are slender, rather widely spread-
ing, without trace of expanded angle, and their outer sides are nearly
parallel (sometimes broader posteriorly than anteriorly). The occiput
bulges far behind the lambdoid suture and is smoothly rounded (except
in old males, in which it is less inflated and is marked by a median ver-
tical ridge). In all of these respects it agrees with the closely related
P. alhinasus and differs from all other known Mexican species. The
frontal is broad and rather tlat interorbitally ; the nasals narrow and
truncate posteriorly; the ascending branches of the premaxilla short,
bluntly rounded posteriorly, and barely reaching plane of orbits. The
pterygoids are parallel lamelhe, their inferior edges slightly everted —
a transition step in the development of the horizontal shelf of Crato-
geomys from the simple lamella of Thomomys. The hamular processes
articulate directly with the audital bulliie. P. bullerl differs from the
nearly related P. alhinasus in smaller size, smaller mastoids (which are
truncate above instead of rounded), narrower rostrum, narrower and
longer nasals, narrower ascending branches of premaxilla, and much
shorter angular process of mandible.

Dental characters. — Upper incisors narrow, with a single median fur-
row; molariform series only slightly heavier than in G. texensis; last
upper molar with a large heel, which equals or exceeds the anterior
prism in antero-posterior diameter.

Measurements. — Average of 2 males from north slope of Sierra Nevada
de Colima, Jalisco (measured in flesh) : Total length, 236 ; tail vertebra",
81.5; hind foot, 33. Average of 4 females from same locality: Total
length, 215.5; tail vertebr;B, 72.5; hind foot, 30.*

For cranial measurements see Table F, p. 214.

Specimens examined. — Six, all from the north slope of the Sierra
Nevada de Jalisco, Mexico.

General remarks. — This species Avas described almost simultaneously
by Mr. Oldfield Thomas and myself, but his description has priority of
publication by about a month. Hence his name, hulleri, has precedence
over my nelsoni. Mr. Nelson states that the species " was found only
in some fields at the upper ranch at the foot of the main north slope of
the Sierra Nevada de Colima, Jalisco, in the upper border of. the lower
pine belt, at about (3,500 feet altitude, where it was common, and was
found in company with the large species, Geomys gymnurus.^^

Pappogeomys hulleri greatly resembles the bisulcate Geomys texensis,
from which its dental characters distinguish it at a glance. It is evi-
dent that both bulleri and texensis have undergone but little modifica-

* lu my original description of (f. nelsoni, the measurements were taken " from
dry skin of type [ (? ], slightly overstaffed," the field measurements not having been
received (Proc. Biol. Soc, Washington, vii, Sept. 29, 1892, 164.) The measurements
as published were: Total length, 250; tail vertebr;ie, 80; hind foot, 30. The flesh
measurements of the same specimen are: Total length, 238; tail vertebrse, 83; hmd
foot, 33. Mr 'J'homas' measurements of his type specimen of bulleri are: Head aud
body, 135; tail, 63; hind feet, with claw, 27,6.

JAN., 1895.]

PAPPOGEOMYS ALBINASUS. 149

tioii since they left the main trunk Hue of the group, aud that both
branched off from points not very remote from the place where Tho-
momys left the same stock.

PAPPOGEOMYS ALBINASUS sp. nov.

Type from Guadalajara, State of Jalisco, Mexico. No. i^^f 9 ad. U. S
National Museum, Department of Agriculture collection. Collected at Atema-
jac, a suburb of Guadalajara, May 21, 1892, by E. W. Nelson (Original No. 2654).

Geographic distribution. — The plain of Guadalajara; limits of range
unknown, Mr. Nelson states: "This species occurs very sparingly on
the open plain about Guadalajara, and diggings of a small gopher,
l^robably the same species, were seen near Ahualulco, some 35 miles
farther west. The range in altitude of these locahties lies between
4,000 and 5,100 feet.''

General characters. — Size small; naked nasal pad well developed;
tail naked. Animal similar to P. huUeri of Thomas, but somewhat
larger; nasal pad and white patch above it more elongated; color paler;
whiskers finer and less conspicuous.

Color. — Uniform pale plumbeous above and below, irregularly washed
with pale chestnut, palest below; a small dark patch around each ear;
an elongated white i)atch on nose inclosing nasal pad and reaching
posteriorly nearly to plane of eyes.

Cranial characters. — Skull small, smoothly rounded like Thomomys;
zygomatic arches parallel, .slender, angle not expanded; temporal
impressions widely distant; zygomatic breadth slightly exceeding
greatest breadth of cranium posteriorly. Skull similar to that of P.
hnUeri,hnt differing in larger size; much larger mastoids, wliicli are
rounded above instead of truncate; broader muzzle; shorter and
broader nasals; broader ascending branches of premaxilla, and more
elongated angular processes of mandible.

Measurements in flesh. — Type specimen 9 ad. Total length, 226; tail
vertebrte, G8; hind foot, 31.

For cranial measurements see Table F, p. 211.

General remarks. — The only known species requiring comparison with
P. alhinasus is the related P. hulleri of Thomas, a smaller and much
more highly colored animal, differing in the cranial characters above
pointed out. ' Future investigations may show that the ranges of the
two meet, and that the animals intergrade, in which case alhinasus will
become a subspecies of hulleri.

Unfortunately, only a single specimen of Pappoyeomys alhinasus is at
hand. But since its type locality, Guadalajara, is an attractive and
accessible locality. It is probable that a large series of specimens will
be obtained in the near future.

150 NORTH AMERICAN FAUNA. [no. 8.

Genus CRATOCJEOMYS ' nob.

(PI. 2; pi. 10, fig. .5; pi. 12, figs. I and 2; pi. 13, fig.s,4-8, and 17; pi. 14, figs. 6 and 7;

pi. 15, figs. (5 and i»; pi. 17, tig. 5; pi. 18, fig. 4 ; pi. 19, fig. 6.)
Type Geomys merriami Thomas, from the Valley of Mexico (pi. 2).

Dental characters. — Upper premolar with three enamel plates (the pos-
terior absent), its shaft strongly convex forward; upper and lower pre-
molars subequal in leugtli. First and second
upper molars with one enamel plate each (pos-
terior absent) ; posterior curvature of m' and m'^
and anterior curvature of nii and m2 strong.

Last upper molar an imperfectly double prism ;
a deep sulcus on outer side; no sulcus on inner
side; crown of tooth normally broader than
long, variable in form, usually more or less ob-

FiG.59. — Cratogeomysiner- , . -, , • i • -, , ,

riami. Crowns of moiari cordatc or subtriaugular ; inucr and outcr cnainel
form teeth: a, upper; b, platcs Variable; inner plate normally at least
^""^^^ two-thirds as long as anterior plate, obliquely

transverse, normally covering posterior face of tooth.

Upper incisor with a single sulcus, median or slightly on inner side,
and usually rather open (fig. 21', 2P, and pi. 15, fig. 9).

Cranial characters. — Skull large and massive; zygomata heavy and
rather broadly spreading ; orbitosphenoids short and broad, articulating
with alisphenoids anteriorly; niesethmoid a half crescent, its apex
pointing to presphenoid; endoturbinals together forming a compact
plate, strongly convex below, straight above, its anterior border sloping
strongly backward without any extension of the os planum in front of
the folds (pi. 19, fig. C) ; first endoturbinal moderately expanded and
elongated; second, third, and fourth subequal; vomerine edge of os
planum curving down below plane of roof of narial passage; floccular
fossa circumscribed and separated from internal auditory meatus by a
distinct ridge; ridge separating inner from superior face of petrous
sharp and incurved, and sometimes rising high posteriorly (pi. 17, fig.
5, and pi. 18, fig. 4).

The following additional characters, of more or less weight, are intro-
duced with special reference to antithesis with Platygeomys: t Breadth
of cranium posteriorly (above mastoids) much less than zygomatic
breadth ; breadth of occipital plane not more than twice its height ; lamb
doid crest broadly convex i^osteriorly ; squamosal expansion chiefly
toward the median line (in C. merriami in advanced age they comx)letely
cover and conceal the parietals, above which they meet in a median
crest) ; mandible longer than broad (including incisors) ; angular process

* Cratogeomys, from uparo^, strong, powerful, -f- Geomijs, in reference to the great
size and strength of the animals.

t Many of the characters already given in the generic diagnosis are also in strong
contrast to those of riuti/geomys.

f

JAN., 1895-1 KEY TO SPECIES OF CRATOGEOMYS. 151

of maiulihle short, nearly sessile, truncated externally, and forming a
shelf (•onii)letely around the base of the outer side of the incisor knob;
S(iuaniosal arm of zygoma covering nearly or quite two-thirds of jugal,
which latter tills but a narrow gap in zygomatic arch (except in one
species, C.fulvescens, in which the jugal is abnormally short posteriorly,
its anterior relations being normal); free part of ui)per edge of jugal
half or less than half the length of basioccipital on median line: paroc-
cipital j)rocesses relatively light; incisors heavy in contrast to those
of riaty(/comys (except in fulvescen,s and castanops); antero-posterior
diameter of incisors greater than transverse (except lufiilvescens and
castano2)s) ; enamel face of lower incisors forming a conspicuous bead
on outer side of tooth, behind which the tooth is strongly beveled, the
transverse diameter being much greater through the enamel face than
posteriorly (except in fulvescens and castanops).

In Cratogeomys a marked depression extends obliquely across the
sq^iamosals from the root of the zygoma to the occiput near the median
line. In the gymmirus series no such depression exists, but, on the con-
trary, a distinct bulge or elevation occupies this part of the skull.

Cratocjeomyti splits naturally into two sections: The merriami series,
comprising merriami, perotensis, estor, oreocetes, audperegrinus; and the
castanops series, comprising castanops and fulvescens. In the merriami
series the top of the skull seen in profile is a nearly straight line; the
zygomata are not strongly decurved, and the outer angle is only mod-
erately expanded. In the castanops series the top of the skull is decid
edly convex, the zygomata are strongly decurved, and the outer angle is
broadly expanded. Numerous other cranial differences exist, and it is
probable that the castanops series will be eventually separated, at least
subgenerically, from Cratogeomys proper.

KEY TO SPECIES OF CRATOGEOMYS.

(1) Basioccipital )'w/rtHr/i«/«r, its sides parallel

Rostrum and brain case long castanops

Rostrum short ; brain case broad gohlinani

(2) Basioccipital truncate wedge-shaped (sides approximating anteriorly),
rt' Sagittal crest well devoloped.

¥ Lower incisor strongly beveled on outer side merriami

If Lower incisor not beveled on outer side.

c' Top of skull strongly convex in i)rofile fulvescens

c- Top of skull nearly dat in profile.

Nasals normal (rather long and narrow) ])erotensi8

Nasals short, narrow posteriorly and broad anteriorly estor

a^ No sagittal crest.*

Outer face of upper incisor strongly beveled oreocetes

Outer face of upper incisor not beveled peregrinus

* The only specimens seen of oreocetes and pe)-egrinua are females ; it is possible
that the old males may have a crest.

152 NORTH AMERICAN FAUNA. [no. 8.

CRATOGEUMYS MERKIAMI (Tlioiiiits).

(PI. 2; pi. 10, fig. 5; pi. 13, fig. 4; pi. 11, (ig. 7; pi. 1.5, figs. (> ami 9; pi. 17, fig. 5;
pi. 18, fig. 4; ].l. 19, fig. 6).

Geornys merriami Thomas, Aunals & Magazine Nat. Hist., Ser. 6, Vol. xii, October,
1893, 271-273. (Type in l^ritish Museum.)

Type from "southern Mexico" — probably the Valley of Mexico.

Geofp-aphic (listrUmtion. — South end of Valley of Mexico and adjacent
mountain slopes from just below the lower edge of the lower pine belt
up to an altitude of 10,000 or 11,000 feet; east to Atlixco (Puebla), north
to Irolo (Hidalgo), and west to Lerma, in Toluca Valley (map 4, i).

General characters. — Size largest of the genus Gratogeomys; tail and
hind feet moderately haired but not so well covered as in G.fulvescens;
skull massive; incisors huge.

Color. — Upper parts dull chestnut brown, mixed with black-tipped
hairs, varying to glossy slate black; underparts similar but paler; the
rusty specimens have a dark patch around and behind each ear, which
is not apparent in the slate-black ones.

Granud characters. — Skull large and massive, the zygomatic arches
widely spreading anteriorly and rapidly Jiarrowing posteriorly (pi. 2);
incisor teeth larger and heavier than in any known Mexican species, not
excepting Flatygeomys gymnurus ; anteroposterior diameter of incisors
much greater than transverse; lower incisors with a strongly marked
bevel on the outer side immediately behind the enamel; behind the
bevel the tooth is abruptly narrower; outer edge of eimmel forming a
conspicuous bead. In adult males the squamosals completely cover the
parietals and meet in a median crest above the sagittal crest proper.
The mandible of the Lerma skull (No. 50110) is longer and narrower
across the angular processes than that of specimens from the slopes of
the Valley of Mexico. Skulls from Irolo differ from the typical form of
the Valley of Mexico in having the mastoids considerably larger and
fuller posteriorly, occupying more of the occipital plane. The audital
bullae also are somewhat more swollen. The mastoids do not extend
out so far laterally as in typical merriami; the postpalatal pits are not
so deep; the coronoid processes of the mandible are more spreading
(directed more strongly outward), and the heel of the last upper molar
is shorter. The Irolo skulls agree with typical merriami and differ from
the Atlixco specimens in having the frontal reach further forward along
the median line than on the sides. Skulls from Atlixco differ from t^'pi-
cal ?He>Tia'Wuntlie following particulars : The nasals extend farther back,
reaching or passing plane of frouto-maxillary suture; the frontal
reaches as far forward laterally as on median line (in merriami it resiches
much further forward on median line) ; as a rule the coronoid processes
of mandible are lower and more abruptly curved backward, with the
coronoid notch correspondingly narrower.

The massiveness of the incisor teeth in true merriami is much more
extreme than in any of the other species, and is coihdinated, as already

.,AN.,lH9i.] CRATOGEOMYS MERRIAMI. 153

poiuted out, with ;i much greater development of tlie squamosal and of
the various prominences and ridges for muscular attachment.

Variations in pelage. — Gratogeomi/.s merriami exhibits both the melan-
istic and chestnut color phases, and also intermediate pelages. In four
adult specimens from Tlalpam, three are dark brown, faintly washed
with fawn color or very pale fulvous; the fourth is bright chestnut or
reddish-brown on the rump and sides, while the newer hair of the back
is intimately mixed with blackish. One specimen from Amecameca
has a white spot above the tail, as in the Irolo specimens.

All of the three specimens from Irolo have an irregular white patch
at the base of the tail above, and one has a small irregular patch on the
rump and another on the belly between the hind legs.

In the Irolo specimens the tail is less hairy and the hind feet more
hairy than usual, and the hairs of the hind feet are white.

One of the eight specimens from Atlixco has the white spot at the
base of the tail, though not so large as in the Irolo and Las Vigas speci-
mens. The hind foot is scant haired in the Atlixco specimens, which
peculiarity is jirobably seasonal, since the Atlixco specimens were col-
lected in July, while those from Irolo were collected in March. The
tails are less hairy than usual in the Irolo and Atlixco specimens.

Measurements (taken in flesh). — Average of 11 males from the south
end of the Valley of Mexico and adjacent slopes (Amecameca, Tlalpam,
Ajusco, Salazar, Huitzilac, and Lerma) : Total length, 380 ; tail vertebme,
112; hind foot, 50. Average of 7 females from same localities: Total
length, 344; tail vertebrae, 105; hind foot, 46.

For cranial measurements see Table D, p. 210.

Speeinie)is examined. — Total number 31, from the following localities:
State of Mexico, Tlalpam, 4; Amecameca, 9; Ajusco, 2; Salazar, 1;
Lerma, 1; State of Morelos, Huitzilac, 3; State of Hidalgo, Irolo, 3;
State of Puebla, Atlixco, 8.

General remarks. — Mr. Nelson states that this large and j)owerful spe-
cies is common in the south end of the Valley of Mexico, where it inhabits
the soft soil of the bordering slopes and ranges on the west, south, and
east sides of the southern two-thirds of the basin. Owing to the hard
rock and clayey character of the middle and northern parts of the
valley it does not occur there. On the west side it ranges up to the
summit of the Sierra de Las Oruces (where he secured a specimen at an
altitude of 11,000 feet near Salazar), and thence down the west slope
into the border of the valley of Toluca, where a specimen was taken at
Lerma. South of the Valley of Mexico it ranges up over the Sierra de
Ajusco to an altitude of 10,000 feet, and across to Huitzilac on the
south slope within the borders of the state of Morelos. On the east
side of the valley it ascends the basal slopes of Mounts Popocatapetl
and 1/taccihuatl. On the southeast slope of Poi)ocatapetl it occurs at
Tochimilco and on the adjacent plain about Atlixco, Puebla. It was
also found at Irolo, Hidalgo, at the extreme north end of tlie Sierra

154 NORTH AMERICAN FAUNA. [no. 8

Nevada de Iztaccihuatl. Wherever found in aiuriciiltiual land it is very
destructive to (!orn, wlieat, an<l other crops.

('RATO(}E()MY!S PEROTENSIS sp. iiov.
( PI. 8, ii-r. fi. )

Type from Cofre de Perote, Vkua Cuu/ (altitude l),r>00 feot). No; 54299 9 ad.
U. S. Nat. Museum, Departmeut of Agriculture collection. Collected May 28,
1893, by E. W. Nelson. (Original No., 4889.)

Geographic distribution. — Cratogeomys perotensis inhabits the west
and higher slopes of the Cofre de Perote, which are wooded, and prob-
ably descends to the northward to meet tlie range of C. estor. Mr.
Nelson's specimens were obtained at the altitudes of 0,500 and 12,000
feet (map 4).

General characters. — Size rather large (smaller than merriami but
larger than estor); no naked nose pad; hind feet and tail rather well
haired.

Color. — Upper parts dark russet fulvous, everywhere finely mixed
with black-tipped hairs; a small dusky patch behind each ear; an
irregular white patch at base of tail in some specimens (in eight out of
thirteen); under parts dark plumbeous, more or less washed with
fulvous; hind feet usually dark proximally and white distally, but
sometimes all white (and not always symmetrical on the two feet). Not
one of the thirteen specimens is in the slaty- plumbeous ijelage so com-
mon in G. merriami. Tliis species has the tail more hairy than in the
others of the merriami series, and in a number of specimens it is irregu-
larly blotched with dusky and white, a iieculiarity not observed in any
other species.

Cranial characters. — Unfortunately the maleoH perotensis is unknown,*
all of the thirteen specimens collected by Mr. Nelson on the Cofre de
Perote being females. The skull of the female, however, furnishes
excellent characters. It agrees witli merriami in general form, in
having the i)rofile of the top of the skull a nearly straight line (not
convex as in fulvescens and castanops) and in having a well developed
sagittal crest. Whether or not the S(juamosals completely overlap the
parietal in the adult male, as they do in merriami, is not known, but
they probably do. Aside from its much smaller size, the skull of the 2
perotensis may be distinguished at a glance from that of merriami, and
from all other known species of Cratogeojnys, by the slender ness of the
jugal anteriorly. The jugal is not at all enlarged anteriorly, and is
deej^ly mortised into the maxillary arm of the zygoma (see pi. 13, fig. 5)«

* Unless one of the specimens obtained near Las Vigas (No. 54311) belongs to this
species instead of estor. It is an immature male, too young to place the identity
beyond question, but has the characters a young male perotensis would be expected
to possess. The skull as a whole is larger than the adult female o{ perotensis (and
hence considerably larger than estor); the rostrum and nasals arc longer; the jugal
is broader anteriorly, and the si|uamos:ils have already crei>t up over part of the
parietals and would undoubtedly meet in advanced age.

TAN, 1895] CRATOGEOMYS ESTOR. 155

111 some instances the squamosal arm of the zygoma reaches so far for-
ward and the maxillary arm so far backward that the two nearly meet
above the jugal. The nasals end on or near the plane of the front of
the zygoma, and the ascending branches of the premaxilla reach back
past the plane of the lachrymals, thus leaving a long median projec-
tion of the frontal between the hinder ends of the premaxillaries.
Skulls of jyerotensis may be distinguished from those of estor (from the
lower northeast slopes of the same mountain) by larger size, much
greater length of rostrum and nasals, slenderness of Jugal anteriorly,
greater length of sagittal crest, and by the form of the frontal between
the orbits, which is broadly rounded instead of flat.

Measurements (taken in flesh). — Type : Total length 300; tail vertebrpe
79; hind foot 40.

Average measurements of twelve females from type locality: Total
length 310; tail vertebra) 88 ; hind foot 41.5.

For cranial measurements see Table D, p. 210.

Specimens examined. — Thirteen, all from Oofre de Perote, Vera Cruz.

CRATOGEOMYS ESTOR sp. nov.

(PI. «, figs. 4 and 5.)

Type from Las Vigas, Vera Cruz (altitude 8,000 feet). No. 54308 ^ ad. U, S. Nat.
Museum, Department of Agriculture collection. Collected June 12, 1893, by E.
W. Nelson. (Original No. 500.5. )

Geograpliic distribution. — The pine-covered hills and flats forming
the extreme northeastern foothills of the Cofre de Perote, and also
the belt of i)ine forest connecting the timber of the mountain witli the
wooded hills of the north. Its range is chiefly east and north of that
of perotensis. G. estor thus reaches the extreme eastern edge of the
table-laud. Mr. Xelson's specimens were obtained at an altitude of
about 8,000 feet (map 4, k).

General characters. — Size medium (smaller than perotensis); naked
nasal pad small or absent; hind feet and tail rather well haired, as in
perotensis.

Color. — Upper parts dark russet fulvous, everywhere finely mixed
with black-tipped hairs; a small dusky patch behind each ear; an irreg-
ular white patch at base of tail above (on all ten specimens) and some-
times one below also; under parts dark plumbeous, more or less washed
with fulvous; hairs of hind feet whitish, usually to ankle. Not one of
the ten specimens is in the melanistic or slaty-pluinbeoiis pelage so
common in merriami.

Cranial characters. — Skull similar to that of perotensis in general form
and profile, the top of the skull a nearly straight line — not strongly
convex as in fulvescens and castanops. Contrasted vf\t\i perotensis (the
only species witli which it requires comparison) G. estor ditters in the
following characters: Size smaller ( <? of estor about equaling 9 of
perotensis); rostrum much shorter; nasals shorter ami broader ante-

156 NORTH AMERICAN FAUNA. [no. 8.

lioily; juji'al broader anteriorly and less deeply i^mbeded between forks
of maxillary arm of zygoma; frontal broader iiiterorbitally on top of
skull, and tiat instead of broadly rouuded; sagittal crest shorter ante-
riorly and perhaps not present iu the female. The female with distant
temporal impressions (No. 54300) figured on pi. 8, fig. 4, is not fully
adult; in advanced age the sagittal area is probably nearly or quite
obliterated by union of the temporal ridges.

Measurements (taken in flesh). — Type ( £ ad.): Total length 315; tail
vertebme 94; hind foot 41.

Average measurements of four males from type locality: Total length
313; tail vertebnp. 89; hind foot 42.

Average measurements of four females from same place: Total
length 277 ; tail vertebra? 75 ; hind foot 37.

For cranial measurements see Table D, p. 210.

Specimens examined. — Ten, all from Las Vigas, Vera Cruz.

General remarks. — C. estor resembles C. perotensis so closely in color
and external characters that the two are practically indistinguishable
except in size, estor being decidedly the smaller. In cranial characters,
however, they are quite distinct, as pointed out above.

Mr. Nelson states that wherever the pine forests are cleared away
and the ground cultivated within the range of this species, the animal
nuiltiplies rapidly and becomes exceedingly destructive to crops.

CRATOGEOMYS OREOCETES sp. nov.

(PI. 8, figs. 1 and 2.)

Typo from Mount Popocatapetl, Mexico (altitude, 11,000 feet). No. 57963 9 yg.
ad. U. S. National Museum, Department of Agriculture collection. Collected
January 7, 1894, by E. W. Nelson and E. A. Goldman. (Original No. 47.)

Geographic distribution. — The boreal higher slopes of Mount Popo-
catapetl, above the range of Cratogeomys merriami (abo/e 11,000 feet
altitude.)

General characters. — Incisor sulcus broadly open and wholly on inner
side; size rather large; pelage soft; nasal pad small; hind feet and tail
sparsely haired.

Color (of type specimen). — Dusky, darkest on head and along median
part of back; tips of hairs washed with i^ale brown; a golden brown
patch under each eye; forefeet dusky ; hind feet white. Apparently
the specimen is just beginning the change from the plumbeous to the
brown pelage.

Cranial characters. — Zygomatic arches narrow, their sides nearly par-
allel; anterior angle moderately expanded (about as iu Heterogeomys
/tis/>?rfws); temporal ridges strongly developed; nasals wedge-shaped,
not inflated anteriorly, ending posteriorly in front of jflane of anterior
face of zygoma; ascending branches of premaxiila just reaching plane
of orbit, not divaricating behind nasals; frontal flat (orbital edge
rounded), rather broad interorbitally and posteriorly, reaching forward

..AN.,1S05.] CRATOGEOMYS OREOCETES. 157

between premaxillte much further than laterally; supraorbital promi-
nences not strongly developed; temporal ridges anterior to interpari-
etal straight, inclosing an elongated wedge-shaped interspace (but very
diiierent from the interspace between the strongly curved ridges of
H. hispidus)', interparietal elongated autero posteriorly, very much
longer than broad; jugal long and large, forming an important part of
arch; lambdoid crest strongly and evenly convex iiosteriorly ; occipital
plane flat, sloping slightly forward from below upward ; j^osterior ends
of palatals excavated laterally; pterygoids uarrow linguhe with parallel
sides, as in C. merriami; audital bulhe relatively short and swollen,
more subglobular than in H. Jmpidus; brain (;ase rising abruptly from
posterior roots of zygomata, much as in hispidus (not flatly rounded as
\\\i\x<A merriami group and in peregrinus). Under jaw short and rather
narrow, as in hispidus; angular processes short.

Dental characters. — Face of upper incisors unisulcate, the groove
wholly on inner side and broadly open, as in merriami — not narrow and
deep as in H. hispidus and M. heterodus; breadth of enamel face of upper
incisor slightly greater than anteroposterior diameter of tooth; outer
side of tooth strongly beveled immediately behind enamel, as in the
lower incisor of merriami. Lower incisor narrow, the transverse diame-
ter less than the anteroposterior. Crown of last upper molar much
broader than long; no distinct heel ; the inner side convex, the outer
side eniarginate and longer. The curvature of the prism of this tooth
is much less than in the merriami series and less than in H. hispidus.

The premolar is the longest tooth and is slightly convex anteriorl}^;
ni' and m^ are hardly shorter and are subequal (or nV^ maybe slightly
the shorter); both are strongly convex anteriorly; m^ is more than
two-thirds the length of m'^ and is only moderately convex anteriorly.

ilAeaswremew/s (taken in flesh). — Type specimen: Total length, 318;
tail vertebrfB, 91'; hind foot, 43.

For cranial measurements see Table D, p. 211.

General remarks. — Cretogeomys oreocetes does not require close com
parison with any known species. From its nearest neighbor of the
lower slopes of the same mountain {C. merriami) it differs conspicuously
in smaller size, narrower zygomata, shorter and more globular audital
bullip, and in the presence of strongly developed temi^oral ridges.

From C. peregrinus, which inhabits the corresponding boreal slopes
of the neighboring mountain, the lofty Iztaccihuatl, it may be distin-
gnished by its narrower and higher craninm, by the beveled outer face
of the upper incisor, the convex (instead of notched) inner border of
crown of last upj)er molar, and other characters mentioned under that
species.

The measurements of the sknll of C. oreocetes (see table D) show that
the posterior breadth of the cranium is nearly equal to the zygomatic
breadth. This is due to the narrowness of the zygomatic arches — not
to any unusual breadth of the cranium posteriorly.

l.")8 NORTH AMERICAN FAUNA. [no 8.

CRATOGEOMYS PEREGRINUS sp. uov.

(I'l. S, fijr. 3.)

Ty]tc from Mount IzTACCiiirATL, Mexico (altitude 11,500 foet). No. 57964 9 old.
U. >S. National Museuiu, Department of AgTiculture colh^ction. Collected Jan-
nary 9, 1894, by E. W. Nelson and E. A. Goldman. (Original No. .50.)

Geographic distribution. — The boreal liiglier vslopes of Mount Iztacci-
biiatl, above the rauge of (Jratogeomys merriami (above 11,500 feet alti-
tude).

General characters. — Size medium or rather large; hind foot and tail
scant haired; nasal pad small; forefoot large (with claws nearly equal-
ing hind foot with claws). Color i^eculiar.

Color (of type and only specimen). — Steel gray from the intimate
adnnxture of dusky and whitish hairs; under parts paler than upper;
thjoat, sides ot face, and fore feet darker. The hairs of the hind foot
are whitish ; of the tail dusky.

Cranial characters. — The skull of the type, a very old female, has the
posterior part of the cranium very Hat and broad, and the zygomata
broad and bowed outward, suggesting Flati/geomys fiimosus. In other
respects the resemblances are more in the direction of Cratogeomys
merriami, with a few characters pointing toward Heterogeomys. The
zygomatic arches are widely spreading, not divergent anteriorly but
broadest across the middle (breadth anteriorly slightly less than great-
est breadth of squamosals jjosteriorly) ; tlie anterior roots stand out at
nearly a right angle; the antero-external angle is moderately expanded
and sharply angular when seen from the side; rounded as seen from
above. Thejugal is rather large and forms an important part of the
arch as in C. merriami. The muzzle and nasals are short, the latter
broad anteriorly and truncated posteriorly about on the plane of the
anterior face of the zygomata. The ascending branches of the premax-
illa are broad and blunt ^posteriorly, barely reach the plane of the
orbits, and do not approximate or divaricate behind the nasals. The
frontal reaches furthest forward along the median line; the suture at
base of maxillary root of zygoma (on top of skull) is nearly a straight
line. There is no sagittal crest, but the temporal ridges approximate
immediately in front of the interparietal, from which jDoint they divar-
icate in both directions; anteriorly they slope slightly outward in nearly
a straight line to a point about opposite the i^osterior part of the post-
orbital prominences of the frontal where they become less distinct
and curve abruptly outward. The iutersi^ace is an elongated wedge,
as in C. oreoeetes, and is not depressed below the level of the temporal
ridges, a result perhaps of the extreme age of the animal. In shape
it differs widely from that of the genus Heterogeomys. The great
breadth of the cranium i)osteriorly is due to lateral expan.sion of the
squamosals, as iu Platygcomys. The greatest breadth across squa-
mosals (over mastoids) is slightly greater than the zygomatic breadth
anteriorly. The interparietal is not covered by the parietals and is

jAN.isnvi CRATOGEOMYS CASTANOPS 159

elongated aiitero-posteriorly. The plane of the occiput is moderately
smooth and slopes forward; it is low and broad, the breadth being
about two and-a half times the height. The mastoid bulhe are much as
in merriami, except that the inferior border is shorter and the inner side
is armed Avith a short blunt spine projecting inward and slightly back-
ward. (This may be abnormal, but the points are symmetrical on the
two sides.) The audital bulhe are rather short and tumid (much as in
oreocetes) and the anterior projection which abuts against the basi-
sphenoidis sharply set oft' by a deep notch on the upper side. The palato-
pterygoids are lingulate, slightly broader than in merriami, the sides
nearly parallel ; mandible short and narrow, resembling that of oreo-
cetes, from Avhich it ditters in having the angular processes even shorter
and the coronoids more hooked.

Dental characters. — Upper incisors with a single very broad and open
groove (broader even than in oreocetes), its deepest point on the iuaer
side of the median line; breadth of enamel face greater than antero-
posterior diameter of tooth. Lower incisors narrow, the breadth of the
enamel face being considerably less than the anteroposterior diameter
of tooth. Crown of last upper molar not distinctly heeled, its inner
border about half the length of outer and deeply notched; outer side
broadly concave.

Measurements (taken in flesh). — Type specimen : Total length, 304; tail
vertebra^, 87 ; hind foot, 42.

For cranial measurements see Table J3, p. 1*11.

CRATOGEOMYS CASTANOPS (Baiicl).
(PI. 12, iig. 1; pi. l.S, fig. 17: pi. U, fig. 6.)

Pseudostoma castanops Baird, Report Stansbury's Exp'd. to Great Salt Lake, June
18.52, 313. (Type from near Bents Fort, Colorado.)

Geomys castanops Baird, Mammals of North America, 1857, 381-386.

Geomijs clarUi Baird, Proc. Acad. Nat. Sci., Phila., vii, 18.5.5, 332. (Ty])e from Pre-
sidio Del Norte, on tlie Rio Grande, Chihuahua, Mexico.)

Type locality: " Prairie road to Bents Fort," near the present town
of Las Animas, Colorado, on the Arkansas Eiver. (Type in U. S.
National Museum.)

Ueographic distribution. — Isolated areas on the Great Plains from
the Arkansas River in Colorado, southward through eastern New Mex-
ico (west to Albuquerque), and western Texas to Santa Rosalia, Chi-
huahua, and Jaral, Coahuila (map 4, h).

General Gharacters. — Size, medium; coloration, yellowish-brown; tad
of medium length; rather scant haired.

Color. — Upper parts yellowish brown or buffy ochraceous tinged with
yellowish, more or less mixed with black- tip])ed hairs, which are much
more numerous in winter pelage; under parts butfy.

Cranial characters. — Skull very broad and heavy; zygomatic arches
widely spreading anteriorly and strongly decurved; profile of skull
convex on top; end of maxillary root of zygoma greatly expanded,

160 NORTH AMERICAN FAUNA. [no. 8.

forming a broad plato, into which the euhirged head of the Jugal is
received; sides of basioccipital parallel. C. castanops differs from C.
fulvesccns m havinij tlie basioccipital narrow, its sides excavated and
j)arallel ; the nasals and nasal branches of the premaxilla more produced
posteriorly; the hitter cutting the plane of the orbits, and in lacking
the thickened sockets of the upper incisors.

Measurements (taken in flesh.) — An adult male from Las Animas,
Colorado (practically type locality): Total length, 295; tail vertebrae,
95; hind foot, 37.

Average of 3 females from same locality: Total length, 256; tail
vertebra', 77 ; hind foot, 33.

For cranial measurements see Table D, p. 211.

Specimens examined. — Total number 43, from tlie fcdlowing localities:
Olney, Colorado, 2; Las Animas, Colorado (type locality), G; Chico
Springs, New Mexico, 2; Albuquerque, New Mexico, 3; Eddy, New
Mexico, 3 ; Sierra Bianca, Texas, 1 ; Marfa, Texas, 3 ; Eagle Pass, Texas,
13; Samalayuca, Chihuahua, Mexico, 2; Gallego, Chihuahua, Mexico, 2;
Santa Rosalia, Chihuahua, Mexico, 4; and Jaral, Coahuila, Mexico, 5.

General remarks. — Coues has already shown that clarTcii can not be
distinguished from castanops, and the examination of a much larger
series than heretofore available confirms this determination. The
peculiar line of demarkation in the type specimen* described by Baird
as separating the color of the head and neck from that of the rest of
the upi)er parts, is now well known as the molt line (which progresses
from before backward) ; and the alleged differences in the feet and skull
donot hold good in the ample series (forty-three specimens) now at hand.
The species i^resents considerable geographic variation in size (mostly
sporadic), as usual in members of the family having an extensive range.
The only notable departure from the type observed in the present series
is in two specimens from Chico Springs, N. Mex. These specimens are
smaller than the type form, brighter and more ' yellowish-chestnut'
in color, and the fore feet, hind feet, and tail are distinctly blackish.
The tail furthermore is well covered with hair for its entire length.

Mr. Vernon Bailey tells me that Gratogeomys castanops is a very inju-
rious species to orchards and nurseries. Along' Onion Creek, 30 miles
southwest of Marfa, in Presidio County, Texas, he found them eating
the roots of fruit trees where "two or three soon sjioil an orchard if
left in it; the owners did not know how to get rid of them."

GRATOGEOMYS CASTANOPS GOLDMANI subsp. iiov.

Type from. Canitas. Zacatecas, Mexico. No, 57965 9 yg. ad. U. S. National Museum,
Department of Agriculture collection. Collected December 24, 1893, by E. A.
Goldman. (Original No. 286.)

* The type specimen, formerly in the Patent Office, is now in the National Museum,
but is in very poor condition, ha\'iug been exposed to the light for nearly forty years,
as a result of which it is so faded that no trace of the original color remains.

JAN., 1895.] CRATOGEOMYS FULVESCENS. 161

General characters, — Similar to C. castaiiops in size and external
appearance, but differing in cranial characters. Tail and hind feet
rather well haired for a Pocket Gopher.

Color. — Upper parts dull butiy-ochraceous, moderately mixed with
black- tipped hairs; under parts paler.

Cranial characters. — Unfortunately all of the five specimens at hand
of this form are females. Compared with females of C. castanops the
skulls diHer in being- broader, shorter, and flatter, with less decurved
zygomata, and decidedly shorter and broader nasal bones. The short-
ening is chiefly in the rostrum; the broadening chiefly in the brain
I case. The basioccipital averages longer and somewhat larger and its
' sides are less truly parallel, being a little broader jjosteriorly than
anteriorly. The plane of the occiput is narrow and much elongated
transver.sely. The coronoid process of the mandible is long, depressed,
and reaches far back.

Measurements (taken in flesh). — Type: Total length, 270; tail verte-

I brje, 90; hind foot, 35,

Average measurements of three females from type locality: Total
length, 257; tail vertebrie, 83; hind foot, 34.3.
For cranial measurements see Table D, p. 211.
Specimens examined. — Total number 5, all from Oaiiitas, Zacatecas.

CRATOGEOMYS FULVE8CENS sp. uov.

(PI. 12, fis. 2.)

Type from Chalchicomula, State ok Pueisla, Mexico. No. 58168 <? ad. U. S.
Natioual Museum, Department of Agriculture collection. • Collected January
15, 1894, by E. W. Nelson and E. A. Goldman. (Original No. 5651.)

I Geographic flistrihution. — The basin-like plain of eastern Puebla,
Mexico, from Esperanza north to Perote and west to the northeast
base of Mount Maliuche in Tlaxcala (map 4, j).

General characters. — Ui^per incisors unisulcate; forefeet shorter than
hind feet. Similar in general ap])earance to G. castanops, but larger;
color darker; tail rather longer, darker, and slightly more hairy.

Color. — Upper parts grizzled yellowish brown, liberally mixed with
dark-tipped hairs; under parts buffy-fulv^ous or ochraceous-bufl'. Com-
pared with castanops the general color is darker, owing to more bounti-
ful admixture of dark-tipped hairs.

Cranial characters. — Skull rather massive; zygomata squarely S])read-
ing, angles broadly expanded; alveoli of upper incisors thickened;
profile of top of skull very convex ; rostrum decurved anteriorly.

The fronto-maxillary suture is peculiar, its anterior end usually
reaching or nearly reaching the plane of the front of the zygoma — in
all the allied species the frontal ends about opposite the middle of the
anterior root of the zygoma.
7433— :No. 8 11

162 NORTH AMERICAN FAUNA. [no.«.

The lieigbt of the roof of the cranium above the palate, and of the
brain case above the posterior roots of the zygomata, are much greater
than in any other member of the genus, and the breadth of the skull
posteriorly is much less.

Contrasted with C. castanops the skull of fulvescens differs in the fol-
lowing particulars: size larger; rostrum broader; sockets of upper
incisors thicker, bulging externally; nasals and ascending branches of
l^remaxilla shorter i^osteriorly, the former hardly reaching plane of
front of zygoma, the latter not reaching jilane of orbits; basioccipital
much broader and wedge-shaped, as usual in the genus (in castanops
the basioccipital is narrower and its sides are parallel, see pi. 12, flgs. 1**
and 2^).

MeasuremenU (taken in flesh). — Type specimen ( t? ) : Total length,
318; tail vertebra^, 102; hind foot, 43.5.

Average of three males from type locality : Total length, 327 ; tail ver-
tebra?, 105; hind foot, 43.

Average of six females from type locality: Total length, 302; tail ver-
tebra?, 97 ; hind foot, 30.0.

For cranial measurements see Table I), p. 211.

Specimens examined. — Total number 11, from the following localities:
Chalchicomula, Puebla, 9; Perote, Vera Cruz, 2.

General rem^arlcs. — C. fulvescens does not require close comparison
with any known species except C. castanops, which it resembles in the
grizzled yellowish-brown color of its upper parts. It is more fulvous
than castanops, from which it difl'ers further in larger size and in the
cranial characters above i)ointed out. Specimens from Perote are more
-ellowish and less fulvous than those from Chalchicomula.

Mr. Nelson states that this species inhabits the sandy open i)lain from
an altitude of 8,000 feet in the lower parts of the basin up to 9,000 feet
on the west slope of Mount Orizaba. He states further: "In this dis-
trict its range is almost identical with that of Dipoilomys phiUipsi.
Like the latter species it follows up the cultivated land into the low^er
border of the pine forest on Mount Orizaba, and is common also about the
northeast base of Mount Malinche. These gophers are particularly
numerous in cultivated ground, and are very destructive to corn and
grain of all kinds."

Genus PLATYGEOMYS " uob.

( PI. 3; pi. 10, tig. 8; pi. 13, tigs. 1-3; pi. 14, tig. 9; pi. 15, tig. 7; pi. 17, tig. 4; pi. 18,

fig. o; pi. 19, fig. 7.)

Type Geomi/s (ji/mintnis Merriam, from Zapotlan, Jalisco, Mexico.

Dental characters. — Upper premolar Avith three enamel plates (the
posterior absent), its shaft nearly straight. First and second upper
molars with one enamel plate each (posterior absent).

* PJatijgeoniijs, from TrXatvi, broad, wide, + Geomys, with reference to the great
breadth of the cranium.

JAN., 1895] GENUS PLATYGEOMYS. 163

Third upper molar an incomplete double prism, the outer side abruptly
narrowed behind the anterior prism; axis of heel anteroposterior;
inner enamel plate nonnally less than two-thirds as long as anterior
plate; not covering posterior face of tooth; outer plate normally as long
as inner and usually reaching posterior edge of heel.

Upper incisor strongly unisulcate^ the sulcus median or slightly on
inner side (fig. 2P).

Cranial characters. — Skull large, heavy, and flat; hinder part of
cranium extraordinarily broad and flat, the great breadth due chiefly to
lateral expansion of the squamosals, which not only project as a thin
shelf beyond the brain case, increasing the size of the glenoid fossa
both anteriorly and posteriorly, but also completely arch over and
conceal the postglenoid notch, curving with only a shallow con-
cavityfrom the posterior angle of the zygomatic arch to and beyond
the extreme tip of the transversely elongated mastoid; zygomatic
arches massive, broadly spreading anteriorly, the antero-external angle
expanded vertically into a triangular plate between the strongly pro-
duced and decurved external angle and the evenly rounded orbit (the
resulting plate made up in part of the distal end of the maxillary arm
of the arch, and in part of the anterior end of the jugal, which is
usually expanded); jugal normally large and broad, forming an impor-
tant part of the arch; pterygoids vertical lamelhe with inferior border
everted; orbitosphenoids larger than in Cratof/eonii/s but not normally
articulating with alispheuoid; mesethmoid a little more than a half
crescent, its anterior border strongly rounded above (pi. 18, fig. 5);
endoturbinals together forming an elongated oblique plate which is
sharply pointed antero-superiorly, owing to the elongation of the upper
endoturbinals (pi. 19, tig. 7) ; no extension of os planum in front of lower
endoturbinals and no curving down of vomerine edge of os planum
below ijlane of roof of narial passage; floccular fossa ill defined and not
separated from internal auditory meatus by a distinct ridge; ridge
separating superior from inner surface of petrous only feebly developed
(pi. 17, fig. 4; and pi. 18, fig. 5).

In addition to the above-described generic characters, most of which
are in strong contrast to those of Cratogeomys, the following points are
selected with special reference to antithesis with Crafof/eomyfi (which
see) : Breadth of cranium posteriorly (above mastoids) equal to or
greater than greatest zygomatic breadth ; breadth of occipital plane at
least two and a half times its height ; lambdoid crest sinuous, presenting
three posterior concavities; squamosal expansion chiefly away from
median line — not covering inner part of parietals; mandible very much
broader than long* (including incisors); angular processes of mandible

* The extraordinary breadth of the mandible across the angular processes is not due
alone to the great length of these processes, but in part to tlieir position. They
are higher and more nearly on a level with the mcisor protuberance than in any
other torm, and the jaw as a whole is Hatter.

164 NORTH AMERICAN FAUNA. [no. 8.

extraordinarily long and spreading, reaching ont so far laterally that
the knob over root of incisor is midway between condyle and end of
angular process fpl. 3) ; squamosal arm of zygoma covering about half
(iu fumosus more than half) of upper edge of jugal, which latter enters
broadly into formation of zygomatic arch; free part of upper edge
of jugal equal to length of basioccipital on median line (except ni
fufnosus); paroccipital processes large and expanded, forming recurved
flanges; incisors slender in contrast to those of the merriami series;
anteroposterior and transverse diameters of incisors subequal ; enamel
face of lower incisors forming an inconspicuous bead on outer side of
tooth, behind which the tooth is not beveled, the transverse diameter
through the enamel face being inappreciably greater than posteriorly.

KEY TO SPECIES OF PLATYGEOMYS.

1" Zygomatic arches parallel or hatred outward in the middle fumosus,

1" Zygomatic arches strongly divergent anteriorly:

Jugiil only slightly expanded auteriory planiceps.

Jugal broadly expanded anteriorly :

Nasals strongly wedge-shaped; narrow posteriorly; reaching

plane of zygoma gymnurus.

Nasals not wedge-shaped; broad posteriorly; not reaching

plane ol zygoma tylorh inus.

PLATYGEOMYS GYMNURUS Merriam.

(Pl.3; pi. 10, fig. 8; pi. 13, fig. 2; pi. 15, fig. 7; pi. 17, fig. 4; pi. 18, fig. 5; pi. 19, fig. 7.)

Geomys gymnurus Merriam, Proc. P.iol. Soc. Washington, vii, Sept. 29, 1892, 166-167.

Tyjie locality. — Zapotlan, Jalisco, Mexico. (Type in U. S.ISratioual
Museum.)

Geographic distrihution. — Valley of Zapotlan and adjacent slopes of
the Sierra Nevada de Colima, Jalisco, and the volcano of Colima down
to the upper edge of the plain of Colima, Mexico.

General characters. — Size very large; a naked pad on end of nose;
tail naked; feet sparsely haired; hinder part of cranium extraordiua.
rily broad.

Color. — Upper parts dark reddish-brown or chestnut, varying to sooty
plumbeous or slate-black, slightly paler below. The rusty specimens
have a dusky xiatch about each ear and a larger one on the nose. The
depth of the chestnut seems to increase with the age of the hair, speci-
mens in the molt having the new hair very dark and only washed on
the tips with chestnut. The hairs of the hind feet are scattered and
nearly colorless. Tlie young are glossy slate-black, with the sides and
rump consi)icuously sprinkled with whitish bristles.

Cranial characters. — The skull of Platygeomys f/ymnurus differs from
all others of the family (except the related P. tylorhinus and planiceps
here described) in the extraordinary breadth and flatness of the hinder
l^art of the brain case, the result of lateral exj)ansion of the squa-

JAN., 1895.] PLATYGEOMYS GYMNURUS. 165

raosals, which completely arch over and conceal .the postglenoid notch,
curving with a shallow concavity from the posterior angle of the zygo-
matic arch to the extreme tips of the transverely elongated mastoids,
which they overreach. The breadth of the cranium here equals or
exceeds the greatest zygomatic breadth. Corr elated with this unpre
cedented breadth of the posterior part of the cranium is an even more
extreme lateral extension of the angular processes of the mandible.
The zygomatic arches are widely spreading anteriorly, with broadly
expanded subtriangular outer angles. The jugals are large, broadly
expanded anteriorly, enter largely into the outer wall of the orbital
fossa, and, as a rule, terminate anteriorly in a straight edge, which
articulates witli the lower third of the ascending or maxillary arm of
the zygoma without being mortised into it as usual in the group;
still the front of the jugal rests on a strong shelf of the maxillary
arm, and is commonly overtopped by a sliort spicule. The exposed
part of the upper edge of the jugal forming part of the outer wall of
tbe orbital fossa is usually, though not always, as long as the basi-
occipital (on median line), and as a rule the posterior half of the jugal is
overlapped by the squamosal arm of the zygoma. Tlie fronto-maxillary
suture is straight or slightly convex outward, while its continuation as
the premaxillo-maxillary suture (on top of the skull) is strongly con-
cave inward, the result being that the suture at the base of the maxil-
lary arm of the zygoma, taken as a whole, is shaped like the letter S
somewhat drawn out. In tylorhinus and planiceps it is broadly and
uniformly convex inward. The nasals end posteriorly on or a little
behind the anterior plane of the zygoma, and are strongly wedge-
shai^ed and much narrower posteriorly than in tylorhinus. The nasal
branches of the j)reniaxilla may or nmy not reach the plane of the
orbits; they a])proximate slightly behind tlie nasals, «

The occipital plane is exceedingly broken and irregular; the lamb-
doid crest overhangs it as a sinuous ledge throughout its entire
lengtli; the greatly enlarged paroccipital processes stand out like
broad flanges from the exoccipitals, projecting strongly outward and
backward, forming, in conjunction with the middle part of the lamb-
doid crest, a remarkable basin-shaped inclosure, outside of which, and
far anterior to the great paroccipital flanges, are the transversely-
elongated mastoids (pi. 15, fig. 7). In striking contrast is the smoothly
planed- off occiput of Heterof/eomys his2)i(lus (pi. lo, fig. 4).

The shape of the lambdoid crest is peculiar; it is deeply sinuous,
with three concavities directed forward (of which the median is deep,
the lateral shallow), and two strong convexities directed backward;
at each end it terminates in a club-shaped knob directed outward.
Looking at the skull from above there is nothing to indicate the
limits of the brain case, the broad squamosals being convex upward
behind the zygomata, without trace of the lateral depression that
marks off" the brain case in Cratogeomys and most other members of
I the family.

166 NORTH AMERICAN FAUNA. [no. 8.

Measurcniciits (tiikeii.iii flesli). — Average of three males from type
locality (Zapotlan, Mexico): Total leiigMi, o5li.G; tail vertebra', 105.3;
bind foot, ~)3.'.'>. Average of three females from same place: Total
length, 341; tail vertebne, 91 ; hind foot, 40.G.

For cranial measurements see Table E, p. 211*.

Specimens examined. — Total number, 10, from the following localities
in Jalisco, Mexico: Zai^otlan, 7; Sierra Nevada de Colima, 3.

General remarks. — Plati/f/eomys (jymnnnis may be regarded as the
type (for it is the largest and most extreme in cranial peculiarities) of
a remarkable series of Pocket Gophers inhabiting southern Mexico
from the Sierra Nevada de Colima of Ja.lisco eastwarTl to the north
slope of the Volcan Toluca in the State of Mexico, and Tula in
Hidalgo. Externally these animals difler so little from the larger spe-
cies of Gratof/eomys as to be distinguished with difficulty, but in cranial
characters they may be told at a glance. The number of recognizable
forms now known is four, of which one {fumosus) is very distinct from
the others; the remaining three are closely related {(jyninurus, tylo-
rliinns, and planiceps) and two of them {tylorhiniis and planiceps) may
be found to intergrade when specimens are obtained from intermediate
localities along the line of their distribution, in which event the latter
must be reduced to subspecific rank. Still another form that might
be deemed worthy of separation is the Patzcuaro animal mentioned
under the head of P. tylorhinus.

All the members of the gymnurus series have the upper parts more
or less plentifully sprinkled with long, slender, bristle-like hairs which
protrude far beyond the ordinary fur. In fumosus these hairs are very
conspicuous, owing to the marked contrast of their whitish color with
the blackish-slate of the body; the same is true of the young in {/ym-
nurvs, but in the adult they harmonize so well with the prevailing
reddish-brown or chestnut tints that they may be easily overlooked.
They are most abundant in the Patzcuaro specimens of iylorliimis.

Mr. Nelson states that the rangxi of Platygeomys {/ymnvrvs, so far as
determined by him, is limited to the valley of Zapotlan and slopes of
the Sierra Nevada de Colima and base of the adjacent volcano of Colima,
and the immediate vicinity. On the north slope of the Sierra Nevada
de Colima he found them up to an altitude of 11,000 feet, among the firs
and alders, where a specimen was secured. Thence to the base of the
mountain they are rather common on open grassy slopes, and range
out over all of the adjacent valley of Zai)otlan. In this latter district
they were usually found in fields, where they do much damage to corn
and wheat. Zapotlan Valley has an altitude of about 4,500 feet, and
is an open basin-like ijlain just below the pines and oaks of the moun-
tains. On the extreme upper border of the phiin of Colima, near the
southwest base of the volcano, at an altitude of about 3,500 feet, he
saw numerous diggings of a gopher, which was probably this species.

1

JAN., 1895.] PLATYGEOMYS TYLORHINUS 167

PLATYGEOMYS TYLORHINUS sp. nov.

(PI. 13, fig. 1.)
Type from TrLA, Hidalgo, Mkxico. No. 51883 <? ad. U. S. National Museum, Depart-
ment of Agriculture collection. Collected March 13, 1893, by E. W. Nelson.
(Original No. 4442.)

Geographic distribution. — Tula, Hidalgo, and tbeiice southwesterly
along the north slope of the Sierra Madre to the vicinity of Patzcuaro,
Michoacan.

General characters. — Size, large; tail nearly naked; a naked pad on
end of nose; coloration dark. Similar to P. gymnnrus, hut smaller,
with shorter and more hairy hind feet, which are distinctly white in
contrast to dark of ankles and legs; .skull remarkably broad and flat,
as in P. gymmirns. but lighter and differing further in important
characters.

Color. — Upper parts chestnut or liver-brown, as in Geomyshursorius;
under parts similar but slightly paler, the plumbeous showing through
in places; legs and ankles ccmcolor with body; hind feet white in
contrast.

Cranial characters. — Skulls of P. tylorhinvs differ from those of
P. (lymnurns in smaller size, narrower rostrum, and shorter nasals,
which do not reach plane of zygomatic arches. The most conspicuous
difference is in the shape of the nasals: instead of being wedge-shaped,
as in fjymnnrus, they are much broader i)osteriorly and abruptly
truncated behind, and the premaxilhe do not approximate behind
them. The skull as a whole is much less massive and the maxillary
arm of the zygoma less thickened than in gymnnrus. The jugal is
enlarged throughout and expanded anteriorly into a broad plate which
abuts against the sides of the maxillary part of the zygomatic arch,
which latter is hardly excavated to receive it, sending out a small shelf
below and a short spicule above, much as in gymnurus. The suture
at the base of maxillary root of zygoma is broadly convex inward; in
gymnurus it is shaped like a drawn-out S. As usual, the skull of the
female is much smaller than that of the male, and the jugal is narrower.

Measurements [t'A\L&\\ in flesh). — Type specimen, 5 ad.: Total length,
345; tail vertebric, 100; hind foot, 45. Average of two 9 specimens
from type locality: Total length, 298; tail vertebrae, 91.5; hind foot,
39.5.

For cranial measurements see Table E, p. 212.

Specimens examined. — Total number 9, from the following localities
in Mexico: Tula, in Hidalgo, 4; Patzcuaro, in Michoacan, 5.

General remarlis. — Sjiecimens from Patzcuaro, State of Michoacan,
are intermediate in size and form of nasals between gymnurus axid tyi)i-
cal tylorhinns from Tula, but exceed the latter in the expansion of the
jugal and whiteness of the hind feet. The hind feet are more hairy,
and the ankles are dark plumbeous instead of chestnut, causing the
white to stand out in stronger contrast. Skulls of the Patzcuaro

168 NORTH AMERICAN FAUNA. [no. 8.

.inimal differ further from those froui Tula in having smaller and shorter
pterygoid lamelliie (as seen from the side), leaving more space between
their posterior edge and the audital bullte. The posterior ends of the
palatals are smaller, thicker, and have the outer edge straighter. In
the Tula skulls the palatals are thinner and broader, with the outer
edge irregularly sinuous. In the Patzcuaro animal the jugals are con-
spicuously broader anteriorly than in those from Tula, but as in the
latter they are much less expanded in the female than in the male.

There is an average difference in external characters by which tlie
Patzcuaro specimens may be distinguished from specimens from Tula
and the Volcano of Toluca. They are darker and richer in color (the
chestnut being more ferruginous), and the head is mainly slate black,
more or less faintly washed with rusty. This color does not cover the
head uniformly but is disposed in a tolerably regular pattern from
which there is little variation in the series of specimens at hand. The
slate-black covers the muzzle, reaching back along the median line as
far as the plane of the eyes, and sends a broad arm backward on each
side to the shoulders, inclosing the eye and ear. The chestnut of the
back comes forward over the top of the head to about the plane of the
eyes, and on the sides of the face below the eyes to and sometimes
including the cheeks. Possibly there is something seasonal in this
pelage; all of the Patzcuaro specimens were collected at the same
time — the latter half of July.

Mr. Nelson contributes the following information respecting the local
distribution of P. iylorhinus: "I found this species common along the
north slope of the mountains about Lake Patzcuaro and thence to the
vicinity of Lake Cuitzeo, in Michoacan. All of this district lies in the
zone immediately below the pines (from about 5,500 to 6,800 feet alti-
tude), and is largely cultivated to wheat and corn. The gophers are
particularly numerous in the fields, where they do considerable damage
to crops. They range up into the lower border of the forest where
Zygogeomys trichopns is found. Beyond Lake Cuitzeo no work was done
to the northeast until Tula, Hidalgo, was reached. There these animals
were found in small numbers at an altitude of about 6,000 feet, in tlie
vicinity of the town. They were only noted about the borders of small
basin-like sinks, where the land was under cultivation. Not being "
numerous here their depredations in the grainfields were of little
moment. The district from Lake Cuitzeo to Patzcuaro has a cool cli-
mate, with abundant rains during the summer months. Tula lies in a
much more arid and warmer zone."

PLATYGEOMYS PLANICEPS sp. nov.

(PI. 13, fig. 3; pi. 14, fig. 9.)
fAscomys mexicanns, Liclit., Brants Muizen, 1827, 27-31 (in part).
Type from north slope Volcan Toluca, Mexico. No. 55906 ^ U. S. National Museum,

Department of Agriculture, collection. Collected September 12, 1893, by E. W.

Nelson. (Original No. 5466.)

JAN., 1895.]

PLATYGEOMYS PLANICEPS. 169

Geographic lUstribution. — N'orthern and eastern slopes of tlie volcano
of Toluca and adjacent part of tbe valley to the city of Toliica, from an
altitude of 8,600 feet up to tbe vicinity of timber line.

General characters. — Similar to I*, tiflorhinus, from wbicb it differs
inappreciably in external appearance except in tbe greater lengtb of
tbe tail. Ui)per incisors unisulcate; skull broad and flat; size large;
tail nearly naked; a naked pad on end of nose; forefeet with claws
shorter than bind.

Color. — Upper parts chestnut, as in Ujlorhlnus from Tula; under
parts similar but paler, the plumbeous basal fur showing through in
places; legs and ankles concolor with body; hairs of hind feet whitish,
but scant. Nose below eyes blackish ; a large blackish spot around each
ear. One specimen is dark plumbeous, washed with chestnut, and has
the head markings described under the Patzcuaro specimens of tijlor-
hinus.

Cranial characters. — Skull similar to that of tylorhinus, from which it
differs chiefly in the form of the jugal bone, which is narrow throughout
or very slightly expanded anteriorly — not broadly expanded as in
tylorhinus. It differs further from tylorhinus in having the nasals less
squarely truncate posteriorly (and ending about on plane of middle
of maxillary root of zygoma); the ascending branches of premaxilla
rounded posteriorly and ending near anterior plane of orbits — not
passing nasals so far as in tylorhinus; the cranium very broad and
flat; occipital plane more than two and a half times as broad as high.
The rostrum is narrow, but not narrower than in some specimens of
tylorhinus from Tula.

Measurements (taken in flesh). — Type specimen $ : Total length, 372;
tail vertebnie, 121 ; hind foot, 40. Average of two females from type
locality: Total length, 336.5; tail vertebme, 100; hind foot, 43.

For cranial measurements see Table E, p. 212.

Specimens examined. — Three, all from the north slope of the Volcan
de Toluca, State of Mexico.

General remarlcs. — This animal may prove to intergrade with tylo-
rhinus of Tula, in which case it must be reduced to subspecific rank.
The number of specimens at hand (only three) is not sufficient to deter-
mine the constancy of the characters that distinguish it from tylorhinus.
The chief differences, as above stated, are the longer tail and narrower
jugal. The jugal is always narrower in females than in males, and two
of the three specimens are females. The raa,le (type specimen), while
full grown, is not. old, and its jugal may be abnormally slender, though
there is nothing about the skull to suggest this belief. In the light of
the present material no course seems open but to recognize the animal
as a distinct species. It may be remarked, however, that it is the
poorest species described in the present paper.

Respecting its local distribution Mr. Nelson states : " On the slopes
of the Volcano of Toluca this species is not very numerous, but is found

170 NORTH AMERICAN FAUNA.

[no. 8.

scattered in small uiiinbers continuously from the base of the mountain
up to the vicinity of timber line, usually in open parts of the pine forest
and in small grassy parks. It is more common in the valley of Toluca,
where it inhabits fields and grassy meadows and is very destructive
to crops,"

PLATYGEOMYS FUMOSUS Merriam.
(PI. 11, fig. 4, and pi. 14, fig. 8.)

Geoniys fnmosiin Merriam, Proc. Biol. Soc. Washington, vii, September 29, 1892,
165- IfiG

Type locality. — Colima City, Mexico. (Type in TJ. S. National
Museum.)

Geographic (Ustribution. — Plain of Colima, Mexico. (Altitude 1,500 to
2,000 feet.) '

General characters. — Size medium, about equalling Geomys hursarius
(smaller than the other species of Platygeomys)', pelage rather soft,
sparingly mixed with long Avhitish bristles, which are most abundant
on the rump; tail and hind feet nearly naked; nasal pad not strongly
developed; color very dark.

Color. — Upper parts everywhere plumbeous slate or dark sooty-
brown, faintly washed in places, particularly along the sides, with pale
reddish-brown; color of upper j^arts fading in worn pelage to pale dull
liver brown, usually in irregular patches; underparts scant haired,
pale plumbeous, sometimes indistinctly wa.shed with pale brownish. A
young specimen, about half grown (No. 341SG S ), is rich slate black
above, conspicuously lined with whitish bristly hairs, which are most
abundant on the rump, and more so on the sides than along the middle
of the back. There is also a fiiint brownish tinge on the sides of tlie
neck. The scant hairs of the belly are very i^ale plumbeous or even
soiled whitish.

Cranial characters. — Skulls of Platygeomys fumosus agree with those
of the other members of the gymnurus group in the extreme breadth
of the hinder part of the cranium, due to the expansion of the squamo-
sals beyond the parieties of the brain case, and in the great lateral
production of the angle of the mandible. P. fumosus departs from
the gymnurus series markedly in the form of the zygomatic arches,
which, when looked at from above, are rounded instead of sharply
angular anteriorly, and have the sides nearly parallel or bowed out-
ward, so that they are broadest across the middle instead of anteriorl}^
In gymnurus they are usually widely divergent anteriorly. P. fumosus
differs further from the other members of thi^, gymnurus series m greater
interorbital breadth of frontals; strongly wedge-shaped nasals; more
elongated postpalatal pits (which reach the plane of front of last
molars), and in having the anterior end of Jugal more deeply embedded
between the terminal forks of the maxillary arm of the zygoma.

,AN..l895.] PLATYGEOMYS FUMOSUS. 171

The jugals are but slightly (sometimes not at all) expanded anteri-
orly, in which respect the species agrees with P. planicejys, from tlie
A^olcauo of Toluca. It differs from the latter greatly in the extent to
which the Jngal enters into the formation of the zygomatic arch; the
jugal being so far overlapped by the maxillary and squamosal roots of
the arch that its free upper border is short — less than half the length of
the basioccipital iu median line. It differs from planiceps further in
broader rostrum, less spreading and more strongly decurved zygomata,
and shorter and broader ascending arms of the premaxilla, which are
bluntly rounded off opposite the middle of the maxillary root of the
zygoma.

Measurements. — Average of seven males from type locality: Total
length, 287.5; tail vertebrae, 82.2; hind foot, 42. Average of three
females from type locality: Total length, 277; tail vertebrae, 75.3; hind
foot, 39.C.
I For cranial measurements see Table E, p. 213.

Specimens examined. — Total number, eleven; all from Colima City,
Colima, Mexico.

General remarks. — Platygeomys fumosushelongfi, to the gymnurus series,
3f which it is the smallest species yet described. It differs markedly
from the other members of the series in having the zygomatic arches
rounded and nearly parallel instead of sharj)ly angular and strongly
diverging anteriorly; and differs further in having the sides and rum])
beset with whitish bristles that ])rotrude far beyond the fur.

The original description of this species was faulty in several respects
and is here corrected. The material collected by Mr. Nelson since the
original description was published has thrown a flood of light not only
on the affinities of this species but also on the whole group. It is now
dearth at /Vwo.sns is not related in any way to hispidus, authentic skulls
of which are now available for the first time.

Mr. Nelson found this species limited in distribution. His notes state
that it was rather numerous in damp saline flats overgrown with cocoa-
nut palms, wild fig trees, mesquites, and acacias, in the valley of the
Colima River near the city of Colima. In the vicinity of Armeria, at
m altitude of about 200 feet, a few hills were seen but none of the
inimals were caught. Thence up the course of the Armeria river, on
the plain of Colima the hills became more and more numerous, especially
between the altitudes of 800 and 2,500 feet. The animals seem to live
m isolated and limited colonies, between which, in apparently equally
favorable ground they occur singly and rarely. One colony of con-
siderable size occupies an open grassy area in the limestone belt between
Colima and the volcano ; others were seen along the sandy border of tlie
Armeria river bottom in a growth of low bushes, and in some thick
thoruy woods on a dry bench bordering the Colima river a few miles
below the city.

172

NORTH AMERICAN FAUNA.

[no. 8,

Genus ORTHOGEOMYS* nob.
(PI. 19, figs. 1 and 2; text figs. 60-64; map 8\)
Type Geomi/s scalops Thomas, from Teiiuaxtepec, Mexico.

Dental eharactcr.s. — ITpper premolar with three or four enamel plates,
the posterior when present restricted to inner fourth; tm' and m^ with
two enamel plates each. Third upper molar with an elongated heel and
deep outer sulcus; inner sulcus variable; both inner and outer enamel
plates normally reaching posterior end of heel, the inner plate usually-
covering the posterior half of the inner side of the tooth, leaving a broad
cement band in front of it (fig. 34, ", «, and »). In 0. scalops the outer
plate is often divided, presenting an anomalous condition in the family
(fig. 62). Posterior curvature of m' and m- and anterior curvature of
mi and m., strongly developed. Shaft of upper pm straight.

Upper incisor unisulcate, the sulcus widely open and slightly on inner
side, but sometimes reaching middle.

Fifi. &0.—Orthof)eomys .icalops. Longitudiual vertioal median aoction of skull, mesetlimoid and rome

in place. (For key see tig. 7.)

I

Fig. r,i.—Oitho<ienm>is xcalops. Mesetlimoid and vomer removed, showing endoturbinals.
(For key see fig. 10.)

* Orthogeom)i«, from ofjfioc, straight, -\- Geomi/s, in reference to the unusual shap
of the skull.

tThe posterior plate is present in both upper premolars of the type and only know
specimen of 0. latifrons, but is altogether absent, or present as a very narrow stri
on one side only, in 0. scalops an<l O. nelsoni.

ljAN..l895.] KEY TO SPECIES OF OKTHOGEOMYS. 173

Skull as a whole miicli elongated; froutal extraordinarily broad and
dat, much broader than muzzle, with sides nearly i)arallel (not exca-
vated or concave laterally between the orbits, fig.
17^); orbital plates of frontal not meeting inferiorly
behind cribriform, but broadly separated by orbi-
tosphenoids, as in Pappogeomys and Thomomys.
Zygomata narrow or only moderately spreading.
Brain case subcylindric, as seen from above, in ¥\(i.Q2.-orthogeomys
I'ontinuation of^ the general form of the froutal f^eaiopa. Last upper mo-

, , 1 1 r, 1^^ ^ 1 i /'A 1 -j^ 1 lar. h, divided outer

ind nnizzle. Angle or mandible short. Urbitosphe- ^,nau^el piate.
loids rather large, articulating with the anterior
:)art of the alisphenoids and sending a tongue upward, partly filling
]he upper i)art of the sphenoidal fissure (fig. 60). Mesethmoid a half
•rescent, as in Cratogeomys; endoturbiuals as a whole quadrangular,
he anterior border essentially parallel to cribriform plate; first endo-
urbinal only slightly expanded and rounded anteriorly, as in Geomys;
hird endoturbinal larger and much broader than second — a unique
;oudition in the family (fig. 61). The palatopterygoids are long and nar-
ow, and of nearly equal breadth throughout; the basal third or less, is
)alatine; the distal two-thirds or more, pterygoid. The foramen rotun-
limi and foramen ovale are nearer together than usual, and sometimes
nerge into a single large opening which communicates directly with
:be alisphenoid canal.

External characters. — Size large; pelage very coarse, hispid or setose;
lasal pad present or absent.

Cranial characters. — The chief cranial characters that distinguish
h-thogeomys from the other genera having essentially the same enamel
)attern of the molariform series (Heterogeomys and Macrogeomys)* are
he great breadth of the frontal interorbitally, absence of interorbital
constriction, absence of conspicuous postoibital prominences or ridges,
arge size and extended relations of orbitosphenoids, peculiar form of
■ndoturbiuals, and shape of the palatopterygoids. The great length
md narrowness of the cranium as a whole is matched by Macrogeomys
htlichocephalus^ but the nearly uniform breadth of the upper part of
lie skull and the form of the zygomata and palatopterygoids are very
lifl'ereut. Tlie posterior position of the lateral enamel plates of m^,
)0th of which normally reach the end of the heel, is a distinctive
•haracter.

KEY TO SPECIES OF OKTHOGEOMYS.

^elage setose; muzzle short latifrons.

^elage not setose; muzzle long:

i Frontal inflated ou orbital margin anteriorly ; m^ normal —

j Nasals broad posteriorly grandis.

I Nasals narrow posteriorly nelsoni.

I Frontal inflatioa slight or absent; m' with outer enamel plate divided.. scaZo^^s

* It has been stated in the preceding footnote that the upper premolar of Ortho-
'eomys normally has only three enamel plates, while in Heierofieomys and Macro-
•eomys four are always present. Hence the enamel pattern can hardly be said to be
he same.

174 NORTH AMERICAN FAUNA.

[NO. 8.

ORTIIOGEOMYS SCALOPS (Thomas).

(PI. 19, figs. 1 and 2, aucl text tigs. 60-62.)

Geomysscalops Thomas, Auuals aucl Mag. Nat. Hist., Othsenes, XIII, May, 1894, 437-438.

Type from Tehuantepec, Mexico. (Type in British Museum).

Gcof/raphic distribution. — Extreme southern Mexico, in State of
Oaxaca, and probal)]y adjacent part of Chiapas.

Mr. Nelson states that on the pme-covered slopes of the Cerro kSan
Felipe, a few miles north of the city of Oaxaca, he found the diggings
of this gopher extending ui)ward from an altitude of about 7,000 feet
to the summit (altitude about 10,500 feet), always in pine or oak timber
or in the small openings that occur in the forest.

General characters. — Size rather large; pelage hispid j naked nasal
pad large (measuring 20 mm. in length in fresh specimen); tail naked;
hind feet naked, except for a few scattered colorless hairs ; ear opening
surrounded by a brpad, thickened rim.

Color. — Type specimen in worn, faded i^elage: "Smoky-brown, tend-
ing rather toward rufous (very near 'Front's brown' of Eidgway)."—
Thomas. An adult specimen from Cerro San Felipe, Oaxaca, col-
lected June 21, 1894, by E. W. Nelson, is in good pelage and is dark
seal-brown (almost black in places) with an evident gloss.

Cranial characters. — Skull of adult 9 very long and narrow; frontal
very broad interorbitally, not constricted in front of postorbital pro-
cesses; zygomata little spreading, flattened, elongated autero-poste-
riorly, the outer sides ])ara]lel; occipital plane sloping forward; paroc-
cipital flanges turned backward, but not reaching plane of occipital
condyles; palatopterygoids narrow, of nearly uniform breadth through-
out, the pterygoids forming distal two thirds, but not reaching base of
notch (see pi. 19, fig. 2). Interiorly the ])remaxilla reaches far behind the
incisive foramina. Contrasted with latifrons, which it greatly resem-
bles, scalo2)s diflers in having the rostrum much longer, the nasals
broader, more arched anteriorly, and longer, and the jugal broader
anteriorly. The resemblances and difterences are such as to at once
suggest sexual variation — the skull of 0. latifrons differing from that of
scalops in the way that female skulls usually differ from males in the
Geomyidw — smaller size, shorter rostrum, and narrower jugals. But,
unfortunately for this hypothesis, the specimen of scalops is an adult
female, as shown both by the collector's label and by the conspicuous
teats on the dry skin. Furthermore, the grooving of the upper incisors
is very different and the external characters are marked.

Since the above was written I have received nine additional speci-
mens of 0. scalo2)s from Mr. Nelson, all collected in the Cerro San Felipe,
near the city of Oaxaca, during the last week of August and 1st of Sep-
tember, 1894. Two of these are adult males. Their skulls differ from
those of the female in slightly larger size; more spreading and some-
what heavier zygomata, which divaricate anteriorly instead of being
parallel; in a more decided tendency to inflation of the anterior part of

JAN., 1895.] ORTHOGEOMYS GRANDIS. 175

the border of the frontal : the development of a long sagittal ridge, and
of much larger jiaroccipital processes, which reach backward behind
the plane of the condyles.

]Male skulls of scalops from Cerro San Felipe, Oaxaca, differ from
males of nelsoni from Totoutepec and Mount Zempoaltepec, Oaxaca
in the following characters. Size smaller, muzzle mucii narrower, the
narrowness especially marked in the ascending branches of the pre-
maxilla 5 nasals decidedly broader posteriorly and less eveidy acum
inate, spreading more abruptly in front of the middle ; zygomatic arches
more slender and more divergent anteriorly; frontal intlation less
pronounced; paroccij)ital processes much larger ami directed more
strongly backward, exceeding the plane of the condyles ; occipital plane
less Hattened, and marked by three ridges, a median ridge and two
lateral; palatoptery golds shorter; groove of upper incisors narrower.

Dental characters. — Molars as in the genus. Upper incisors with a
single deep and rather broad furrow wholly on inner side: outer side
strongly convex. In latifrons the groove is relatively shallow and
median, or nearly so. The outer enamel plate of the last upper molar
is usually divided, making four instead of three plates for this tooth,
a condition not observed elsewhere in the family (tig. 62).

Measurements. — Type specimen (measured by Thomas from dry skin):
Head and body, 270; tail, 95; hind foot, 45.2 (without claw, 40).

Average of two males from Cerro San Felipe, Oaxaca (measured in
flesh): Total length, 369; tail vertebrae, 103.5; hind foot, 50.*

Average of eight females from same place: Total length, 360; tail
vertebrne, 109 ; hind foot, 50.

Cranial measurements. — Type specimen (measured hj Thomas) : Basal
length, 63; basilar length of Hensel, 56.7; greatest zygomatic breadth,
40.8; nasals, length 20, greatest breadth, S; least breadth of muzzle
above maxillo-premaxillary suture, 15; interorbital breadth, 14.2;
between tips of postorbital j)rocesses, 10.2; postglenoid breadth, 26.7;
greatest squamosal breadth, 39; basion to occipital crest, 18.4; between
tips of paroccipital processes, 27.5; palate from gnathion, 47; diastema,
24.5. Upper molar series on crowns, 12.6; breadth of m', 4; least
height of muzzle on diastema, 12.

For other cranial measurements see Table F, p. 214.

specimens examined. — Total number 13: 10 from Cerro San Felipe,
Oaxaca, Mexico; 3 from mountains 15 miles west of city of Oaxaca.

General remarks. — Orthogeomys scalops seems to be more closely
related to 0. grandis than to 0. nelsoni.

ORTHOGEOMYS GRANDIS (Tliouias).
(Text fig. (>3.)
Geomii>s (jratuUsThomtiH, Auuals aud Magazine Nat. Hist., 6 ,ser., XII, October, 1893,
pp. 270-271.

Type locality. — Duexas Guatemala. (Type m British Museum).

*A larger series of males would undoubtedly result in larger average measure-
ments, as neither of our specimens are very old.

176 NORTH AMERICAN FAUNA. [no. 8.

Oeographic distribution. — "Cominou all over the liighlaud« [of Guate-
mala!, and traces of their presence are to be met with almost every-
where in the neighborhood of Dueiias." — Biologia Centrali- Americana,
Mammalui, 1880, IGO.

General characters. — Size very large; upper incisors deeply unisul-
cate, the sulcus on inner side and widely open ; tail naked ; fore and hind
feet "very thinly haired, the few scattered bristles whitish;" pelage
coarse. The following quotation is from Mr. Thomas's description of
the type specimen:

Color. — "Smoky chocolate brown throughout, except on the muzzle,
cheeks, and chin, where the hairs are white or pale whitish brown.
A few white hairs scattered over the back."

Cranial characters. — " Skull large and heavily built. Ascending
processes of premaxillaries surpassing the nasals by about a quarter of
an inch ; the space between them behind the nasals less than the breadth
of one of them. Interorbital space broad, as broad as the muzzle, its
edges anteriorly rounded and inflated in a manner quite unique. Zygo-
mata not very widely expanded in proportion to the size of the skull.

"Incisors pale yellow or whitish, in marked contrast to the deep
orange found in the allied species. Their single groove deep and very
widely open, so that its greatest width on the cutting edge amounts to
2 mm.; in position the bottom of the groove is internal, the breadtb
of the inner portion of the tooth being about 13 to 45 percent of the
whole; owing, however, to the great breadth of the. groove itself, it
considerably overlaps the median line, but the above percentage is
taken strictly from the bottom of the groove. Molar teeth large."*

Measurements of type specimen (from dry skin). — Head and body, 320;
tail, 135; hind foot, with claw, 57 ; without claw, 50; longest foreclaw, 23.

For cranial measureinents see Table F, (i^. 214).

General remarks. — This animal, though long known from Guatemala,
had been confounded with Idspiilus until recently separated by Mr.
Thomas, who, struck by its larger size and some other external differ-
ences, removed the skull from one of Mr. Salvin's original Duefias
si)ecimens and discovered the remarkable cranial peculiarities above

mentioned.

ORTHOGEOM.Y.S NELSONI sp. iiov.

(Text lig. 63. )
Type from Mt. Zempoaltepec, Oaxaca, Mexico. (Altitude 8,000 feet.) No. 66751
(? ad. U. S. Natioual Museum, Department of Agriculture Collection. Collected
July 8, 1894, by E. W. Nelson and E. A. Goldman. Original No. 6376.

Geographic distribution. — Mt. Zempoaltepec in the State of Oaxaca,
Mexico, and the adjacent region, including Gomaltepec and Totontepec.

General characters. — Size, largest of the known species of the family,
slightly exceeding 0. grandis of Guatemala, which it closely resembles,
differing chietly in the fronto-nasal region of the skull. Ears larger
than in any other member of the family; naked nasal pad large; tail
naked except at base.

* Annals and Magazine Nat. Hist., XII, October, 1893, 270-271.

JAN., 189.0

OKTHOGEOMYS NELSONI.

177

Color. — liuiform dull dark-brown; hardly paler below.

Cranial characters. — Skull large, 'ong, and heavy, resembling both
scalops and gramUs, but differing from both in the shape of the nasal
bones, wliich are very much narroa-er pos-
teriorly. Mr, Oldfield Thomas has had
the kindness to compare his type of
grauflis with the type and other skulls
of nelsoni sent him for the purpose, and
has taken the trouble to give nie a sketch
of the fronto-nasal region of grandis,
with a number of detailed measurements
which show the differences between the
two forms. In addition to the striking
narrowness of the nasals posteriorly,
nelsoni differs from grandis further in
the following points : the ascending arms
of the premaxilla reach much further
backward, cutting the plane of the orbit;
the articular face of the maxillary root
of the zygoma (on top of the skull) is
much longer, measuring 11.5 instead of
8.7mm. ; the frontal is both narrower and
shorter between the nasal branches of
the premaxilla; the muzzle is narrower,
the frontal broader, and the frontal in-
flations are more anterior and less ex-
treme. The mandible differs, not only from grandis, but from all known
members of the family in the absence of the capsular inflation over the
root of the incisor, between the condyle and angular process. It is
entirely wanting in the type, and only family apparent in the adult
female from the same locality. It is larger, but still abnormally small,
in an old male from near Totontepec (No. CG753). The skull of the latter
specimen is the largest I have seen of the species and the jugal is
broader anteriorly than in the specimens from Mount Zempoaltepec.

Skulls of 0. nelsoni differ from those of O. scalops in larger size, much
broader muzzle, heavier zygomata, longer nasals, which are much nar-
rower posteriorly and truly cuneate in form; much broader ascending
branches of premaxilla; broader and decidedly more inflated frontal;
U-shaped, instead of V-shaped postglenoid notch; flatter occipital
plane, with less backward extension of the paroccipital processes.

Measurements. — Type specimen, an adult $ from Mount Zempoalte-
pec: total length, 397; tail, lL'3; hind foot, 53. Another male, from
near Totontepec, is even larger: total length, 435; tail, 140; hind foot,
55. An adult female from INIount Zempoaltepec measures: total length,
380; tail, 118; hind foot, 52.

For cranial measurements see Table F, p. 214.
7433— No. 8 12

Fig. 63. — Orlhogeom>/s nelaoni cT type
(natural size). From Mouut Zempoalte-
pec, Oaxaca, Mexico.

178 • NORTH AMERICAN FAUNA. [no. 8.

Specimens examined. — Five, all from the State of Oaxaca, southern
Mexico: Mount Zempoaltepec, 2; near Toton tepee, 2; Comaltepec, 1.

General remarks. — In color the specimens of 0. nelsoni (lifter mate-
rially from Mr. Thomas's description of grandis. They are in ^vorn
l)elage, and are very dark-brown, but the muzzle and cheeks are not
paler. In fresh pelage they would i)robably resemble 0. sealo2)s in
being rich seal-brown, almost black. The feet are evidently more hairy
than those of grandis^ and the ears are larger than in any other mcDi-
ber of the family, measuring about 5 mm. in height in the dry skin.

ORTHOGEOMYS LATIFRONS sp. nov.
(PI. 11, figs. 5 and 6; text fig. 64.)

T(/2)f/ro»i Guatemala. Exact locality unknown. No. . U. S. National Museum

(No. 2 World's Fair exhibit of Gnateniala).

General characters. — Size medium (rather small for the tropical
American species); incisor groove median or nearly so; tail long and
absolutely naked ; hind feet naked, except a few scattering hairs; fore-
feet scant haired ; nasal pad small or absent ; pelage hispid, scant and
unusually long, unlike any known species of the family. The indi-
vidual hairs are bristles, very much coarser and longer than those of
Geomys liispidns. There is no under fur. The belly is so sparsely
haired that the bare skin shows through.

Color. — Everywhere uniform dull sooty-brown.

Cranial cliaracters — Unfortunately the skull of the type and only
known specimen of this remarkable animal is defective, the entire
occipital region and the audita! bulhe being absent. The anterior part
of the skull is perfect, including all of the teeth and one of the zygo-
matic arches. The upper surface of the cranium is remarkably smooth
and free from lateral indentations or projections, and is of almost uni-
form breadth. Seen from above, the muzzle, frontal, and brain case
pass into one another without interruption or constriction, the frontal
being a trifle wider than the muzzle and the cylindrical brain case a
trifle broader than the frontal. There is only a faint attempt at a
postorbital prominenc^e, and it is below the level of the top of the
skull and is made up of ttie alisphenoid and squamosal. The muzzle
is short. The zygomata are narrow and slender, without any enlarge-
ment or expansion at any point; they are broader i^osteriorly than
anteriorly, and the maxillary arm slopes strongly backward. The
jugal is small and slender and the arch is incomi)lete without it. The
I)alatopterygoids are broken off. The ascending branches of the pre-
maxilla slightly surpass the plane of the orbits. Inferiorly the pre-
maxilla slightly passes the ])osterior end of the incisive foramina. The
nasals are small, short, and narrow, but slightly broader anteriorly
than posteriorly, and without trace of inflation. The angles of the
mandible are short and flat. Unfortunately the i)alatopterygoids and
audital bulhe are broken off, along with the whole of the occipital
region, hence additional important characters may exist that are not
apparent in the single specimen at hand.

JAN, 1895] GENUS HETEROGEOMYS. 179

Dental characters. — The single groove of the upper incisors is median,
open, and rather shallow, and the face of the tooth slopes toward it from
both sides. It thus differs widely from the deep and abrupt groove of
G. scalops, which is wholly on the inner side. The face of the incisors is
orange; in scalops it is pale yellowish or straw-color. The long axes of
the crowns of the individual molars are not quite
transverse, but slope slightly backward toward
the median line. In most species they slope for-
ward. The heel of the last upper molar is short,
but is sharply circumscribed. In addition to the
usual deep sulcus on the outer side, the inner '
side is abruptly narrowed (tigs. 34" and 64). The

. , Fig. 64. — Orthogeomys lati-

enamel plates are peculiar: Inner enamel plate /;oris(tyiie). downs of mo-
covering considerably more than half of inner lariform teeth.- « upper; h
side of tooth, its anterior end bent outward at ^'^^^'
nearly a right angle; its posterior end curved towar<l the median line
and reaching the hindermost part of the heel; outer enamel plate cover-
ing about five-sixths of the outer side of the tooth, its anterior third
bent outward at right angles, its posterior half sloping strongly back-
ward to the end of the heel, forming nearly a right angle with the mid-
dle part and thus making two sharp angles instead of one — a unique
condition. The posterior interspace is very narrow and is on the
median line of the tooth behind. The inner interspace is twice as broad
as the i)osterior.

Measurements (from dry skin, not overstuffed). — Total length, 320;
head and body, 235; tail. 100; hind foot with claw, 44; hind foot with-
out claw, 39.

General remarTis. — Externally Orthogeomys latifrons may be distin-
guished from all other known members of the Geonnjidw by the char-
acter of the pelage, M'hich is setose, the individual hairs being long-
bristles. In cranial characters it closely resembles 0. scalops, but dif-
fers in the much shorter muzzle and nasals (which latter are not at all
inflated anteriorly), and narrower jugal. The upper incisors are very
uidike. In latifrons the face is orange, the -groove median, or nearly
median, and relatively shallow, and both sides slope similarly into it.
In scalops the face is pale j^ellowish or straw color, the groove wholly
on the inner side and deep and abrupt, and the outer side is strongly
(roundly) convex.

Genus HETEROGEOMYS * uob.

(PI. 4; pi. 14, tig. 12; pi. 15, tig. 2; pi. 17, tig. 1; pi. 18, fig. 3; pi. 19, tig. 5; text

tigs. 6~) and 66; map 3^.)

T'lpi (leovii/n hispidiis LeConte, from near .Talapa, Vkra Cruz, Mexico.

Dental characters. — Upper premolar with four enamel plates, the pos-
terior restricted to inner or lingual half. Upper aud lower premolars

* Hetcrogeomys, from trepur, different, -f- Geomys.

180 NORTH AMERICAN FAUNA. [no. 8.

siibequai in length. First and second npper molars witli two enamel
plates each, the posterior complete. Third npper molar a double prism ;
crown much longer than broad; posterior loop or heel strongly devel-
oped; outer sulcus deep; inner sulcus slight; inner enamel plate cover-
ing half or more than half of inner side of tooth and falling short of
hinder end of heel; outer enamel plate very long, covering the whole
of the outer side of the tooth behind the anterior cement band, and
curving inward posteriorly to the median line of the tooth. At the
lateral sulcus the outer enamel band bends outward at right angles.
Posterior curvature of m' and m- and anterior curvature of mi and m2
slight. Shaft of upper pm straight or faintly convex forward. Upper
incisor unisulcate, the sulcus wholly on inner side of median line and
sometimes on inner third; deep and abrnj^t (fig. 20^).

Cranial characters. — Skull as a whole high and narrow; frontal broad
and fiat; its sides biconcave interorbitally; distance between orbits
much greater than length of basioccipital on median line; temporal
impressions anteriorly defining a marked frontal shield ( fig. 17') ;
orbital i)late of frontal usually perforated by a foramen above apex of
sphenoidal fissure; zygomatic arches variable, outer sides nearly
parallel, antero-exterual angle sharp and moderately expanded; infe-
rior surface of palatopterygoids cuneate-lingulate, long and slender,
the palatal arms much elongated, the pterygoid part small and ter-
minal; postpalatal pits deep; nasals nnich arched anteriorly to support
the large nasal callosity; occipital i^lane but little more than twice as
broad as high, very flat, sloping strongly forward from below upward,
squamosal part very high above mastoid bullae; orbitosphenoids shield-
shaped, rather narrow and long, not articulating with alisphenoids ; *
upper part of optic foramen disappearing in advanced life (pi. 17, fig. 1) ;
endoturbinals peculiar, the first greatly expanded, its anterior face
vertical or slightly emarginate (pi. 19, fig. 5). Mesethmoid rather
small and strongly convex anteriorly (pi. 18, fig. 3). Sijuamosal expan-
sion slight; fronto-maxillary suture reaching orbit in front of lachry-
mal (instead of behind, as usual). Mandible short and compact; angu-
lar processes short.

KEY TO SPECIES OF HETEROGEOMYS.

Zygomata broadly spreading, divergent anteriorly ; nasals short torridiis.

Zygomata not broadly spreading and not divergent anteriorly ; nasals rather

Ion"' hispidits.

* In immature skulls of S^fitero(7eomi/s the orbitosphenoid seems to articulate ante-
riorly Avith the maxilla as well as the frontal, but careful examination shovrs it to be
separated by the narrow descending arm of the frontal. In rare cases, irregular
absorption of the exceedingly thin plate may permit the orbitosphenoid to reach
the maxilla,

JAN., 1895. J HETEROGEOMYS HISPIDUS. 181

HETEROGEOMYS HISPIDUS (LeConte).
(PI. 4; textlig-. 65; pi. 13, lig. 20 ; i>l. U, tig. 12; pi. 15. fig. 4.'
Geomys hispidus LeConte, Proc. Acad. Nat. Sci., Pliila.,v, S eptember. 1852, 158.

Type locality. — N"ear Jalapa, Vera Cruz,* Mexico. (Type in Acad. Nat.
Sciences, Pliila.)

Geographic distribution. — The 'Tierra Templada,' or middle belt,
along the basal slope of the table-laud, in the State of Vera Cruz,
Mexico, between the altitudes of 4,000 and 4,500 feet. Mr. Nelson found
the species common about Jalapa and Jico, aud in less abundance from
near the city of Orizaba north to Huatusco. The U. S. National
Museum contains a specimen from Necostla (near Orizaba).

General characters. — Size large; upi)er incisors deeply unisulcate, the
sulcus wholly on inner side; tail naked; a large naked ])ad on end of
nose; forefeet with claws shorter than hind; pelage harsh and stift',
unlike any other species known to occur in Mexico except torridus.

Color. — Upper parts everywhere uniform dark seal-brown; i hardly
paler below.

Cranial characters. — Skull as a whole high and narrow; frontal
very broad and flat, depressed and biconcave interorbitally, concave
both longitudinally and transversely; distance between orbits much
greater than length of basioccipital on median liui^; temporal im^jres-
sions forming elevated semicircular ridges sei)arated in both sexes by a
distinct interval, and extending from postorbital prominences to outer
angles of interparietal, anteriorly defining a marked frontal shiehl, and
posteriorly inclosing a broad interparietal; zygomatic arches narrow,
the maxillary arms sloping strongly backward, outer sides nearly
parallel (sometimes broadest across the middle instead of anteriorly),
antero-external angle sharp and moderately expanded, but not in the
usual way; angle not produced downward; expansion oval in shape
and encroaching on orbital fossa, which is correspondingly narrowed
at this point; inferior surface of i:»alatine bones greatly elongated i^os-
teriorly, forming, on either side of the postpalatal notch, narrow Ungu-
late extensions which are terminated by short and narrow pterygoids;
postpalatal pits deep; ascending branches of premaxilla broad and
bluntly rounded posteriorly; premaxilla extending far enough posteri-
orly to inclose incisive foramina; nasals inflated anteriorly and then
contracted at nares; anterior nares larger than in the other groups;
occipital plane a little more than twice as broad as high, very flat (free

" The type specimen was collected by Mr. Pease in 1847 on the road followed by
Scott's army "between Vera Cruz and the City of Mexico," which road passes through
Jalapa. Mr. Nelson found the species abundant about Jalapa,, which is in the
'Tierra Templada,' about halfway down the slope from the table-land to the coastal
plain. He ascertained further that the species does not occur on the table-land,
which is inhabited by other genera.

tThis color maybe otherwise described as very dark plumbeous, faintly tinged
with purple.

182

NORTH AMERICAN FAUNA.

[no. 8.

from the projections and irregularities coinnioii to other forms), sloping
strongly forward from below upward; brain case larger, more clearly
defined, and higher above posterior root of zygoma than in any other
group; squamosal expansion minimum, neither extending out far later-
ally nor iucreasing length of glenoid fossa anteriorly — the usual shelf
like projection into the orbito-temporal fossa from the posterior root of
the zygoma being nearly obsolete; fronto-maxillary suture reaching
orbit in front of lachrymal (instead of behind it as usual). Tliis arrange-
ment broadens the frontal anteriorly, shortening and apparently weak-
ening the attachment of the maxillary root of the zygoma. ^landible
short and compact, little spreading posteriorly; angular j)rocess short;
prominence over root of incisor low and flattened posteriorly; condylar
process long and only slightly sloping inward.

(65.)

(66.)

Fig. 65.—Heterogeomyshispi(his. Jico, Vera Cruz, Mexico. (Nat. size.)

Fig. 66. — Heteroacomya torridus. Motzorongo, Vera Cruz, Mexico. (Nat. size.)

Dental characters. — Front face of incisors perfectly flat, not rounded
off on edges as in Gcomt/s, PlatygeomyN, and Zj/(/o(jeomi/s. Upper
incisors deeply unisulcate, the groove narrow and wholly on inner
side. Lower incisors without bevel or groove on outer face. Molars
larger, heavier, and less flattened antero-posteriorly than in Geomys
or Zygogeomys; crown of last upper molar elongated posteriorly and
abruptly narrowed behind lateral sulcus, tlie crown. of posterior prism
longer than anterior, to which it forms a distinct heel. Isthmus con-
necting anterior and posterior lobes of uj)per premolar decidedly on
inner side of tooth.

Measurements (taken in flesh). — Average of two males from near type
locality (Jico, 7 miles south of Jalapa, Vera Oruz) : Total length, 345;

fl

JAN., 1895.] HETEROGEOMYS TORRIDUS. 183

tail vertebme, 92.5; bind foot, 53. Average of three females from same
place: Total lengtli, 310.0; tail vertebrti', 85.3; hind foot, 47.3.*

For cranial measurements see Table F, p. 1*15.

Specimens eiamined. — Total number 9, from the following localities
in the State of Vera Cruz, Mexico: Jico, 0; Huatusco, 1; Necostla, 1;
locality unknown, 1.

General remarks. — Through the courtesy of Mr. Witmer Stone and
other officers of the Academy of ISTatural Sciences of Philadelphia, the
type specimen of Geomys hispidus has been sent me for examination.
In size, character of j)elage, and all other respects except color, it
agrees almost exa(;tly with Mr. Nelson's specimens. The color, Avhich
LeConte described as "reddish-brown" and Baird as "red dish- brown or
dull chestnut," was probably the result of museum exposure, the skin
being mounted and exposed to the light. It was collected by Mr. Pease
in 1817, during tlie march of Scott's army from VeraCruz to the City of
Mexico, and consequently had been in the collection five years before
it was described by LeConte. The fading has continued, the specimen
now being much paler than when seen by Baird in 1855.

In view of the large number of species of Pocket G-ophers now known
to inhabit southern Mexico, it is exceedingly gratifying to be able to
settle the status of his2)uliis by actual comparison of the type sijecimeu
with the series collected by Mr. Nelson at or very near the original type
locality. The skull of the type specimen has never been removed, and
the cranial characters of the species have remained unrecorded until
the present time. The series of skulls obtained by Mr. Nelson there-
fore were examined with unusual interest and the result was a complete
surprise. They show not only that the animal is a strongly marked
species, but that it is generically distinct from Geomys, as already
pointed out.

The naked nasal pad is more largely developed in this species than
in any of the others, and its large size is clearly correlated with the
inflated nasal bones. For this reason it shows to unusual advantage
in the type specimen, which is mounted with the skull inside, the arched
nasals keeping it stretched in its natural relations. In this specimen
it measures 12.5 mm. in length by 10 in breadth.

Mr. Nelson states that H. hispidus is confined to the district suitable to
the cultivation of coffee and sugar cane and is said to be very injurions
to cane i^lantations.

HETEROGEOMYS TORRIDUS sp. uov.
(PI. 15, fig. 2; pi. 17, fig. 1; pi. 18, fig. 3; pi. 19, fig. 5; text fig., G6.)
Type from Chichicaxtle, Vera Cruz. No. 63629 9 ad., U. S. Natioual Museum,
Department of Agriculture collection. Collected February 15, 1894, by E. W.
Nelson. (Original number, 5850.)

Geographic distribution. — Lowlands of Vera Cruz, from Chichicaxtle

*The measurements of the feet of the mounted type specimen as taken by me now,
nearly half a century after its capture, are : T'orefoot from basal pad to tip of longest
claw, 42.5; hind foot from heel to tip of longest claw, 45.5.

184 NORTH AMERICAN FAUNA. [no. 8.

and Motzoroiigo to Catemaco, aud tlieuce into Guatemala; penetrating
the interior to lieyes, Oaxaca, and (xLiatemala City, Guatemala.

Mr. Nelson tir^t observed this species on the way from Mirador to
the coast, from an altitude of about 1,500 feet near Santa Maria, down
to the border of the sand hills along the coast at Antigua, The next
l)oint where it was noted was on the route from the city of Cordoba to
the hacienda of Motzorongo. At an altitude of 800 feet at this latter
place it was again found in abundance. The easternmost locality at
which it was obtained, by Mr. Nelson is Catemaco, in the district of
Tuxtlas. He afterwards secured it at Reyes, in northern Oaxaca, at an
altitude of 6,700 feet. The range of the species is strictly tropical.

General characters. — Similar to H. hispidus. Size large; tail naked;
naked nasal pad large; hind feet nearly naked; fore feet scant haired.

Color. — Everywhere dark seal-brown, only slightly paler below; in
worn pelage chocolate brown.

Cranial characters. — Skull large, heavy and rather broad, resembling
that of H. hisjjidus^ from which it differs in the following particulars:
Pituitary fossa deeper aud (usually?) perforate; zygomata much more
squarely spreading anteriorly (the maxillary arm standing out at more
nearly a I'ight angle instead of sloping strongly backward); temporal
impressions uniting iwsteriorly in old of both sexes, but not rising in
a sagittal crest; audital bullii3 smaller, narrower anteriorly, and not
sending u}) a point or ridge toward hamular process of pterygoid; ascend-
ing arms of premaxilla averaging broader and shorter posteriorly. The
skull of the male differs from that of the female in larger size and greater
angularity. The zygomata reach out much further sideways, are much
broader anteriorly than across the middle, and the outer angle stands out
prominently (in the female it turns downward). The jugal is consid-
erably larger and broader anteriorly in the male.

Measurements (taken in tiesh). — Type specimen ( 9 ad. from Chichi-
caxtle): Total length, 323; tail vertebroe, 88; hind foot, 52.

Average of four adult males from Motzorongo: Total length, 348;
tail vertebrti?, 9(3.5; hind foot, 49.2.

Average often adult females from Motzorongo: Total length, 317;
tail vertebrfB, 81.5 ; hind foot, 45.5. The 9 from Eeyes, Oaxaca, is
decidedly larger, measuring: total length, 332; tail, 98; hind foot, 49,5.

The mounted specimen in the World's Fair exhibit from Guatemala,
which is considerably overstuffed, now measures: Total length, 380;
tail vertebrae, 85; hind foot, 46. It is a female.

For cranial measurements see Table F, p. 215.

/Specimens examined. — Total number 27: 2 from Guatemala; 1 from
Reyes, Oaxaca, and 24 from the following localities in Vera Cruz, Mex-
ico: Chichicaxtle (type locality), 1; Motzorongo, 22; Catemaco, 1.

General remarks. — Heterogeomys torridus differs but little externally
from true his^ndns. Even in color the type specimen, which is in worn
pelage, except on the head, is only a shade paler than si)ecimens of his-

JAN., 1895] GENUS MACROGEOMYS. 185

pidus in worn pelage. The differences in cranial characters, however,
are marked and constant. Still it is quite possible that mtergrades
maybe found in the exceedingly narrow belt separating the two forms.
It should be observed that the type specimen has a hind foot 4 mm.
longer than the largest female from Motzorongo, and that the skull,
also, is larger. The type is a very old individual.

Two specimens of a Heterogeojnys from (xuatemala, belonging to the
U. S. ^national Museum collection, are here referred to the present
species. One of these, a young adult (No. AV~A) "^^s collected many
years ago near Guatemala City by Dr. Van Patten; the other was
recently presented to the Museum by the Guatemala Commissioners to
the World's Fair. The exact locality where it was obtained is unknown.
It is an old female, and the temporal impressions meet over the middle
part of the sagittal suture (which is obliterated, as in all adults of the
species). The specimen obtained by Dr. Van Patten (probably also a
female) is younger, and the temporal impressions are still distant. The
two Guatemala skulls differ from those from Vera Cruz in having the
postorbital prominence obsolete or nearly so.

Mr. Nelson states that in Vera Cruz this species is one of the most
injurious of the genus to the agriculturist. At Catemaco he found it
in small numbers among the dry hills and plains on the western border
of the lake, but in the forest on the eastern shore it swarms in countless
numbers. At one point the ground was fairly honeycombed with their
tunnels, so that he sank to the knee at nearly every step.

Hefergeomys torrid ks becomes sexually mature at a remarkably early
age. Several of the young females were mothers, and one in particular,
though hardly half grown, has long ]^endant teats that have evidently
been nursed. This specimen (Xo. 03040) is still m the woolly pelage of
the very youiig, and its skull, barely half the size of the adult, has not
yet attained tlie mature form. The animal could hardly be more than
three months old. Its measurements m the tiesli are : Total length, 259 ;
tail vertebra^, 71; hind foot, 43.

Genus MACROGEOMYS * uob.

(PI 5; pi. 11, figs. 2 aud 3; pi. 13, figs. 18, 19, 22, and 23; pi. 14, figs. 3 and 10.)
Type Geomiifs heierodus Peters, from Costa Rica.

Dental characters. — Upper premolar with four enamel plates, the pos-
terior restricted to inner third; m' and m^ with two enamel plates
each. Last upper molar witli an elongated heel and deep outer sulcns;
inner emargmation variable (slight in heierodus; deep in doUchoceph-
alus); inner enamel plate covering half to two-thirds of inner side of
the tooth, its posterior end nearly reaching hinder end of heel. Outer
enamel plate variable, the posterior limb double the length of the ante-
rior. In ili. heterodus it covers half; in dolichoceiihalus aud eostari-

' Maciogeomys, from /uaKpw, large, great, -f Geomys, in reference to the large size
of the animals.

186 NORTH AMERICAN FAUNA. [no. 8.

censis^ three-fourtlis of the outer side of thetootli. The posterior loop
or heel is greatly developed, sittaiuiiig the maximuiu size known in the
family (about half or more than half the length of the tooth and uar
row, the constriction about half the breadth of tlie anterior prism).

Posterior curvature of m^ and ni'' and anterior curvature of mj and in?
strongly developed. Shaft of both upper and lower premolar strongly
convex forward and very large and heavy.

Upper incisor unisulcate, the sulcus wholly on iyner third of face,
narrow and deep; face of tooth flat on both sides of sulcus (fig. 20',
and pi. 15, tig. 8).

Cranial characters. — Frontal broad, flat, depressed or concave along
the median line, deeply excavated laterally between the orbits, the
notch immediately succeeded by a strongly developed postorbital proc-
ess (much larger than in any other member of the family, fig. 17').
Palatopterygoids broad, short, and truncated posteriorly, the horizontal
part composed almost wholly of the palatal, the pterygoid simply cap-
ping the end and abruptly upturned at right angles (fig. 11^). Nasals
moderately convex, slightly or not inflated. Brain case rising high
above posterior root of zygoma. Unfortunately there are no skulls of
Macrogeomys in the Department collection: hence I have been unable
to make sections to ex])ose the mesethmoid and turbiuals.

The lambdoid crest is straight or slightly convex posteriorly (not
sinuous as in riatygeomys) and the occipital jdane Is flat and slopes
strongly forward, as in Ileterogeomys.

External characters. — Size large; naked nasal pad well developed;
tail naked; pelage soft, almost silky, and with a tendency to become
wavy; color pattern unique-, bicolor : muzzle and sides of rump abruptly
whitish ; rest of upper parts dark chocolate or sepia in marked con-
trast. (The color pattern of the adult M. costaricensis and cherriei is
unknown.)

General remarlcs. — Macrogeomys requires comparison with only two
genera, Hcterogeomys and Orthogcomys, from both of which it may be
distinguished at a glance, whether viewed from above or below. The
most striking points of diflerence are the remarkably short and broad
palatopterygoids and the strongly developed postorbital processes.

KEY TO SPECIES OF MACROGEOMYS.

AudUal hidla normal, outer side not flattened.

Skull short and broad ; zy "omata divergent anteriorly heterodus.

Skull long and narrow ; zygomata parallel dolhlioeephalus.

AudUal huUa pccurmr, the outer side flattened and disk-shaped.

Jugal normal, entering largely into zygoma cherriei.

Jugal small, the zygoma complete above without it costaricensis

MACROGEOMYS HETERODUS (Peters).
(PL 11, fig. 2; pl.U, fig. 3).
Geomiis heterodus Peters, Monatsber. K. Preuss. Akad. Wiss., Cerlm (1864), 1865, 177.
(Tran.slation of original description appended to present article, p. 189.)

Type locality. — Costa Rica. Exact locality unknown.

,AN..1895.] MAOROGEOMYS HETERODUS. 187

GetK/raphic distribution. — The Ira/Ai range and perhaps other parts of
Costa Eica.

General eharacters. — Si/e large; face of upper incisors deeply unisul-
cate, tlie sulcus narrow and wholly on inner side of median hue; enamel
face of incisors orange; naked nasal pad large; tail absolutely naked;
hind feet naked, with a few stift' hairs about the toes; fore feet nearly
naked (shorter than hind); pelage moderately coarse, but not hispid as
[in G. hispidus; no external ears. Coloration peculiar, the muzzle and
^ides, including sides of rump, being conspicuously paler than rest of
ipper parts.

Color. — Upper parts uniform sepia or hair brown; muzzle, under
i)arts, and sides all round abruptly much paler, the pale color (a soiled
i-ray) reaching higher on the sides of rump than elsewhere and includ-
ing base of tail.

Cranial characters. — Skull large, heavy, and rather short; zygomata
jroadly spreading, their sides divergent anteriorly, maxillary arms
;loi)ing backward less strongly than in (lolichocephalus; antero external
ingle well marked, moderately expanded ; jugal large and broad, its
ipper surface not covered by squamosal and maxillary arms; frontal
jroad and flat, concave along tlie median line between the orbits and
leeply notched on the sides immediately in front of the large post-
)rl)ital processes, which latter are capped by the apex of the alisphenoid
md overlapped posteriorly by the anterior edge of the squamosal.
S^asalsbroadly wedge-shaped and not inflated. The ascending branches
»f the premaxilla slightly exceed the plane of the orbits. Inferiorly the
)remaxilla reaches but does not inclose the posterior end of the incisive
"or9jmina. The zygomatic breadth is considerably greater than the
■greatest squamosal or mastoid breadth. The occipital plane is flat
except a vertical median ridge) and slopes moderately forward ; the
ambdoid crest is straight, slightly incurved near median line. The
lalatopterygoids are broadly U-shaped and shortly truncate posteri-
orly, the pterygoids abruptly upturned at right angles to the palatals.
The basioccipital has the sides parallel for the anterior half and is
broadly wedge-shaped posteriorly. Audital and mastoid bullii^. normal.
The enamel face of the upper incisors is flat, with the sulcus deep, rather
narrow, and wholly on inner side. Traces of the fine inner sulcus may
ilso be seen in the only specimen at hand. The heel of the last upper
molar is narrow, much elongated, and slopes strongly outward.

Macrof/eomys heterodus differs from 3f. dolichoceplialus, the only known
sjiecies Avitli which it requires comparison, in the very different form
of the skull as a whole, it being nuich shorter and broader, and in the
following details: Jugal broadest anteriorly and not covered by squa-
mosal and maxillary arms of zygoma; zygomata divergent anteriorly
(mstead of ])arallel); nasals shorter and not inflated; orbital borders
of frontal not inflated anteriorly; muzzle and diastema much shorter;
palatopterygoids less broad at base; occipital plane broader and lower;

188 NORTH AMERICAN FAUNA. fms.

mastoid biillai iiiiiTower vertically. Mandible iiuich slioiter. Heel of
last upper molar longer and narrower, the outer eaamel plate reach-
ing little moie than halfway from sulcus to end of lieel; in fJolicho
cephalus it reaches all the way.

Measurements. — Peters recorded no measurements for his type speci-
men, but Dr. Matschie lias kindly measured it for me and finds the total
leugtli 325 mm. He states that the tail is defective. The specimen in
the U. S. ^National Museum, from the Irazu Mountains, wliich is the
subject of the foregoing description (a well made dry skin), affords the
following measurements: Total length, 325; head and body, 280; tail,
65; hind foot with claw, 45; hind foot without claw, 41.

For cranial measurements see Table F, p. 215.

General remarks. — The only species kn(>wn to me with which hetero-
dus needs comparison is dolichocrplialits, which agrees with it in the
abrupt paleness of the nnizzle and sides of the rump. But heterodm
differs from dolicliocephnlus m liaving the entire under parts and lower
sides of the same pale color as the muzzle and sides of the rump. It
differs further (in the specimens at hand) m the tint of the upper
parts, which is sepia or hair brown instead of chocolate brown, and in
the cranial characters above pointed out.

Unfortunately, Peters's description ot his G. lieterodus from Costa Rica
is brief and unaccompanied by measurements, cranial characters, or
exact locality (see next page). That his animal is the same as hispidm
of LeConte (from Vera Cruz), as assumed by Coues and Alston, is
exceedingly improbable on geographic grounds (in view of the remark-
ably restricted ranges of all the tropical American species row known)
and impossible in view of the wide difference in coloration. Peters
described heterodits as bicolnr, the upper i)arts "dark brown,"' the muz-
zle, rump, and underi^arts "brownish gray or white.'' Hispidus is eon-
color and uniformly dark. Fortunately the type of Peters's heterodus is
extant. It is still in the Berlin Museum, and Dr. Paul Matschie of that
museum has had the kindness to send me additional notes, accompanied
by full cranial measurements, which suffice to place its identity beyond
question.

Through the courtesy of Mr. F. W. True, Curator of Mammals in the
United States National Museum, I have been able to examine several
specimens of the G corny Ida- from Costa Rica and Guatemala. Among
those from Costa Rica is one which agrees in every way with Peters's
original description of heterodus, and also with the additional particu-
lars concerning Peters's type specimen kindly furnished me by Dr.
Matschie. This specimen was recently presented to the museum by
the Costa Rica Government through its commissioners to the World's
Columbian Fxposition at Chicago in 1893. It consists of a well-pre-
pared skill, from which Mr. True has kindly had the skull extracted.
It is the only specimen of heterodus I have seen, and is the subject ot
the foregoing description. Mr. George K. Cherrie, of the Costa Rica

JAN., 1895) MACROGEOMYS DOLICHOCEPHALUS. 189

National Museum, in res{)onse to a letter of iuqiiiiy, coutributes the
f'ollowiDjj important statement respecting this specimen: "It is No. 313
of tbe collection of the ' Museo Nacional,' an adult male; was collected
October 15, 181)0, near Rancho Iledondo, a point on the Irazu range
between the volcanoes Irazu and Barba, at an altitude of about 1,400
meters. The specimen was purchased from a 'peon' and mounted by
myself. October is the last month of the rainy season, and the month
in which it rains hardest. I might also add that the species is abundant
in the locality given above."

Peters's original description of hcterodus is as follows: "Our museum
has received through Dr. Hoffmann and Dr. v. Frantzius the skin with
the perfect skull of a new species of Geomi/s from Costa Rica, whereby
the geographical distribution of this genus in Central America is estab-
lished. This species agrees best with G. mexicanus Licht. in size, pro-
portion of tlie limbs, nakedness of the tail, and the nature of its hairy
covering, which latter, however, appears to be somewhat shorter and
stifter. The color is dark brown except on the belly, rump, and muzzle,
which are brownish gray or white. It is, however, readdy distinguished
by the position of the deep longitudinal groove of the upper incisors,
which does not run along the middle but between the inner and middle
thirds of the teeth, for which reason I propose to name the species Geoniys
heterodusy (Monatsber. K. Preuss. Akad. Wiss., Berlin, 1864, 177.)

Dr. Paul Matschie has kindly sent me the following cranial meas-
urements of Peters's type specimen of heterodiis, which is in the Berlin
Museum (No. 2864) :

Greatest basal length (condyle to front of premaxilla), 61; basal
length (basion to gnathion), 58 ; basilar length of Hensel (basion to alveo-
lous of incisor), 51.2; greatest breadth across squamosals, 38; least
breadth between postglenoid notches, 27.5; least interorbital breadth,
11; breadth across postorbital processes, 15.25; height of cranium above
palate, 24; height above basion, 17; length of upper molar series on
alveoli, 14; length of diastema, 22.5; length of single mandible (condyle
to front of jaw between incisors), 44; breadth across angular processes,
40; distance from condyle to end of angular process, 13; breadth of
muzzle just in front of zygoma, 15.

MACROGEOMYS DOLICHOCEPHALUS sp. uov.

(PI. 5; pi. 10, fig. 7; pi. 13, fig. 19.)

Type from. San Jose, Costa Rica. No. jH^tAV J ad. Collected Janiiary, 1866, by
Jo86 C. Zeledon.

Geographic distributum. — Vicinity of San Jose, Costa Rica. Range
unknown.

General characters. — Size large. Animal similar to .1/. hcterodus; face
of upper incisors deeply unisulcate, the sulcus narrow and wholly on
inner side of median line (pi. 15, fig. 8) ; enamel face of incisors orange;

190 NORTH AMERICAN FAUNA. |no.8.

naked nasal pad large;* tail absolutely naked; bind feet naked, with a
few stiff hairs about the toes; fore feet nearly naked (shorter than hind);
pelage moderately coarse, but not hispid ;is in Jleterof/comys hi.spuhis;
no external ears. Coloration ])eculiar, the muzzle and sides of rump
conspicuously paler than rest of ui)per parts, as in heterodus.

Color. — Upper parts dull chocolate browu, except muzzle and lower
part of rump, which are buffy iu couspicuous contrast, but without liue
of demarkation. (The buffy of the rump surrounds the base of the tail
and reaches further anteriorly on the sides than along the middle of the
back.) Under parts simihir to back but paler, without line of demar-
kation ; wrists and ankles pale. No dark patch around ears.

Cranial characters. — The skull of Macrof/eomys dolichocephalus, in
addition to the generic characters which associate it with. M. heterodm,
is remarkable for its length and narrowness, the zygomatic breadth in
an old male (the type specimen) beiug only 58 percent of the total
length (from condyle to point of premaxilla), and tlie greatest squamo-
sal or mastoid breadth only 57 percent. The opposite extreme is found
in the genus Platygeomys, in wh4ch the corresponding ratios in P. gym-
mtrus are 71 and 75.

The zygomata are not ouly very narrow, but present the appearance
of having been drawn out while in a plastic condition. The maxillary
arms slope strongly backward and are broadly rounded off' without
tra(.'e of angle or of angular expansion at the usual place, though
there is a shght expansion about the middle of the outer side of the
ai-ih, encroaching on the orbitotemporal fossa, which it constricts in
the middle opposite the large postorbital processes — a step toward the
differentiafion ot these two fossie from one another. The jugal is
broad, short anteriorly, narrower at both ends than iu the middle,
and is overlapped by the maxillary and squamosal arms of the
zygoma, which nearly or quite meet above it. The frontal is grooved
medially between the orbits and is somewhat inflated along the
margin of the orbits behind the lachrymal bones, in this respect
resembling 0 grandis of Thomas, though the inflation is much less
extreme. The sides of the frontal are deeply notched immediately in
front ot the large postorbital processes. The nasals are wedge-shaped
as iu heterodus, but longer and slightly inflated anteriorly; they are
broadest near junction of middle and anterior thirds (in the S only).
The ascending branches of the preuiaxilla barely reach the plane of
the orbits. Inferiorly the premaxilla reaches the posterior end of, but
does not inclose, the incisive foramina, as in hcterodus. The zygomatic
breadth is only a trifle greatei than the mastoid breadth. The 0(;<'i])ital
plane is flat, high, and slopes strongly forward; the lambdoid crest is
slightly convex posteriorly. The palatopterygoids are very broad and

''In an alcoholic specimeu (No 1466 U. S. Nationul Museum) the nasal pad or cal-
losity 18 broad and rather short, not reaching posteriorly behind plane of upper

incisors.

JAN, 1895] MACROGEOMYS BOIJCHOCEPHALUS. 191

short. (Ill the male skull the pterygoids are broken off; m the female
they are abruptly upturned, as in heterodns.] The basioccipital lias the
sides parallel in the anterior half and is broadly wedge-shaped poste-
riorly. The height of the cranium above the palate is unusually great,
and the zygomata do not descend below a plane drawn midway of the
height of the skull. The audital bulhe are normal and rather short,
l)lump, and well rounded anteriorly. The brain case seen from above
is subcylindric in shape, in which respect it resembles Ortliogeomys.
The nasals end in front of the plane of the zygomatic arches, while the
premaxillai reach the plane of the orbits, causing an unusual elongation
of the median part of the frontal in order to articulate with the nasals.
The mandible is long and narrow. The enamel face of the upper incisors
is flat, the sulcus deep, narrow, and wholly on inner side (fig. 20').

A young female of M. doUcliocephalus (Xo. 30820) differs from the old
male above described (36295) in the following particulars: The skull as
a whole is very much smaller (see table of cranial measurements) ; nasals
very much shorter, flatter, and broadest anteriorly (instead of at junc-
tion of middle and anterior thirds) ; temporal impressions distant (inter-
space 3 to 4 mm broad); brain case less cylindrical (owing in part to
gTeater depth of constriction running obliquely upward from posterior
root of zygoma to occiput, and in part to a slight bulging upward of the
middle of the brain case) ; basiocciiiital narrower. The top of the skull
in profile is not a straight line, tlie brain case presenting a slight con-
vexity behind the orbits, while the interparietal and occipital crest fall
below the plane of the upper surface as a whole. In both sexes the
ciuterior part of the nasals is strongly decurved.

M. (lolichocepha(u.s differs markedly from J/, heterodns, the only species
rtitii which it requires comparison, in the general form of the cranium,
.vhich is narrow and greatly elongated; in the narrow, drawn out
'Tgomata, without trace of angular projection or expansion ; in the nar-
•ower jugal, which is covered above by the anterior and posterior arms
'f the, arch, Avhich meet ot nearly meet above it; in having the zygo-
nata parallel (instead of divergent anteriorly); the nasals longer and
iomewhat inflated anteriorly; the muzzle and diastema much longer;
he palatopterygoids broader at base; the occipital plane higher and
ess broad; the mastoid bull* much higher vertically; and the mandible
'inch longer.

Measurements (of type specimen, $ ad., from dry skin): Total length,
ibout 380 (approximate, as the tail was not wired and is shrunken);
lead and body, 310; tail, about 75 (approximate only); hind foot, 48;
\ ithout claw, 45.

Measurement of a young female from Costa Rica, preserved in alcohol
^^' HiU ? yg. ad., U. S. National Museum, collected bv Jose C. Zeledon
ind received in October,1884): Total length,310; tail, 74; hind foot, with
law, 49; without claw, 43; forefoot, with claw, 45; without claw, 33.

For cianial measurements see Table F, p. 215.

192 NORTH AMERICAN FAUNA. [mg.

General rpmarl-s. — Extern;illy Macrogeomys ilolirhocephalns resembles
31. lirierodus in the peculiar paleness of the muzzle and sides of the
rump (in strong' contrast to the color of the rest of the upper parts), but
differs from heterodus in not havin.i;- the lower pait of the sides and belly,
of the same pale tint. On the other hand, the pale color of the rump
reaches a little further forward on the dorsal surface. There is a slight
difference also iu the tint of the upper parts, the color being dull choco-
late brown iu (loHchocephaJus, while it is sepia or hair brown in heterodus.
The important cranial differences have been ]>ointed out.

The alcoholic specimen already mentioned (No. 14GC6) is a female, and
although not fully adult, has borne young, as shown by the large pen-
dent ni])ples. The teats are: pectoral. y, inguinal | = |, as usual in the
group. The pectoral pair are situated on the sides immediately behind
the fore legs. The inguinal pairs are not on the belly at all, but on the
mner side of the fA<V///s just below and outside of the belly.

The great callosity at the hinder edge of the wrist is made up of two
large tubercles resembling kernels of corn placed side by side and cov-
ered by common integument.

M4.CROGEOMYS COSTARICEXSIS sp. nov.

(PI. 11, fig. 3; pi. 13, fig. 23; pi. 14, fig. 10.)

Type from Pacuare, Costa Rica. No. iMH j"'^'- U. S. National Museum. Collected
in 1876 by Juan Cooper. (Original No. 96.)

General characters. — Ui^per incisors with a single dee]) sulcus wholly
on inner side; pelage in type specimen (immature) short aud silky,
suggesting the fine crinkled pelage of Didelphi.s miirina; tail and hind
feet naked; a conspicuous naked pad on end of nose.

Color, — Upper parts uniform dark-brown, not paler on nose and
rump; underparts abruptly whitish. The type and only known speci-
men has a large symmetrical white spot on top of the head, occupying
about three-fourths of the area bounded by the eyes and ears.*

Cranial characters (of immature skull, pi. 11, fig. 3). — Similar iu a
general way to an immature 9 skull of M. dolichocephalus (Xo. 36820),
from which it differs in the foUowiug particulars: Nasals very much
broader throughout, particularly posteriorly; space between posterior
ends of ascending arms of premaxilla about twice as broad; zygomata
standing out more squarely, nearly at right angles to axis of skull,
with anterior angle abruptly rounded; jugal narrower; palatoptery-
goids shorter and broader; basioccipital very much broader and wedge-
shaped, its inferior surface not excavated by audital bullae ; audital

*The white crown patch of costaricensis was at first believed to be abnormal,
ailing in the same category with the irregular white blotches frequently fouud ou
the throat and sometimes at the base of the tail, iu various species of pot ket
gophers. But the fact that the spot is bilaterally symmetrical, and is repeated in
the only specimen known of a closely allied species, cherriei, points to its perma-
nence, at least as a mark of the young

JAN. 1895.] MACROGEOMYS COSTARICENSIS. 193

bulla peculiar, compressed, the outer side strongly flattened' more
smoothly rounded, somewhat disk-shaped, and separated from the
mastoid bulla interiorly by a distinct groove. The only other known
species of the family having a similar audital bulla is Macrogeomys
cherriei of Allen. Both are known from single specimens only, and
both are too young to show all of the characters of the adult. Their
specific distinctness will be apparent at a glance at the accompanying
cut (fig. 67) showing the differences in the jugals. The palatoptery-
golds also are different. The palatopterygoids of 71/. costaricensis are
shown on pi. 14, fig. 10, but the figure is inaccurate; in the specimen
they are sliorter and broader, more nearly as in fig. 3 of the same
plate. The pterygoids of cherriei are broken, but the remaining base
shows that they are considerably more slender.

In M. costaricensis the jugal is much shorter than the basioccipital
(measured from condyle) and is wholly inferior, the maxillary and
squamosal roots of the zygoma meeting above it and on its inner side,
so that when ^•iewed from the inner side it appears only as a narrow
edge with the apex upward (fig. 67, *). In position and relations,
therefore, it resembles Zygogeomys trichopus, though considerably
broader than in that species.

Fig. 67.— Zygomatic arches of Jilacrogcomyg costaricensis (3 and 4), and If. cherriei (1 and 2). 1 and 3

outer side ; 2 and 4 inner side.

Measurements. — Type specimen (probably not more than two-thirds
grown) from dry skin: Total length, 330; tail (apparently stretched),
100 from point assumed to be over first caudal vertebra, 80 from
apparent base; hind foot, 37 (without claw, 33).

For cranial measurements see Table F, i>. 215.

General remarlcs. — This singular species, for the privilege of describ-
ing which I am indebted to the courtesy of Mr. F. W. True, Curator
of Mammals in the U. S. National Museum, is represented in the
collection by an immature specimen only. At first it was supposed to
be the young of M. (lolic/iocephahis, but comparison of its skull with
that of dolichocephalus shows numerous points of specific difference, as
above mentioned. While the peculiar texture of its pelage may be
due in part to immaturity, this explanation fails when applied to the
cranial characters which, as described above, are numerous and strik-
ing and of such a nature that most of them would be accentuated by
age. In external appearance the animal bears a striking resemblance
to the young type of Macrogeomys cherriei.
7433— No. 8 13

194 NORTH AMERICAN FAUNA. [no.8.

MACEOGEOMYS CHERRIEI (Alleu^

(PI. 15, fig. 1.)
Geomys cherriei Allen, Bull. Am. Mus. Nat. Hist., V, 337-338, Dec. 16, 1893.

Type from Santa Clara, Costa Rica. No. G64 S im. Museo Nacional
de Costa Rica. Collected in October, 1892, by George K. Clierrie.

General characters. — Naked nasal pad large; tail and hind feet naked.
Similar to Macrogeomys costaricciisis in size and coloration, including
the white head patch, but differing in important cranial characters.

Color (of type, jnv.). — Upper parts very dark plumbeous or sooty
brown.; under parts abruptly paler, with distinct line of demarkation;
top of head between eyes and ears pure white.

Cranial characters [i^rom skull of type, but little more than half grown,
pi. 15, fig. 1).— The skull of ili. cherriei agrees with Heterogeomys
hispidus in general form, in the widely-seiiarated temporal imjiressious;
the broad and fiat frontal, depressed between the orbits; the flat
forward-sloping occipital plane; the form of the zygomata; the inflated
nasals, and the short and compact under jaw, with short angular pro-
cesses. But it is so young that one must be cautious in placing much
stress on characters that vary with age. It differs from H. hispidus
and agrees with 71/. costaricensis in the convexity of the anterior part
of the roof of the brain case;* m the peculiarly flattened and smoothly
rounded audital bulla?, which are separated from the mastoid bullae by
a distinct inferior transverse groove; and in the h>ng heel of the last
upper molar. It differs from costaricensis in the size, form, and rela-
tions of the jugal (as shown in fig. 07), in narrower palatopterygoid
lingular, and in a narrower gap behind the nasals (between posterior
ends of ascending branches of prcmaxilla). -The jugal is large and
long, and nearly half of its upper edge enters into the orbital fossa; it
is not covered anteriorly by the maxillary arm of the zygoma, and its
total length is greater than that of the basioccipital (measured from
condyle). In M. costaricensis the jugal is much shorter than the basi-
occipital (measured from condyle), and is completely covered by the
maxillary and squamosal arms of the zygoma, which meet above it (fig.
07). It differs further from costaricensis in the shape of the horizontal
part of the zygomatic arch, which is not strongly convex upward, and
lacks the constriction tending toward the separation of the orbital
from the temporal fossa. The large orbitotemporal fosstii are broadest
across the middle — ^just Avhere they are narrowest in costaricensis.

Measurements. — Hind foot, with claws, 39 mm. (in dry skin). No
measurements were recorded from the flesh, and the specimen is far
from full grown.

For cranial measurements see Table F, p. 215.

General re»mr7c,s.— Through the courtesy of Dr. J. A. Allen, Curator
of Mammals in the American Museum of Natural History of New

* It 18 probable that the saddle-shaped frontal ot costaricensis and cherriei is the
resxalt of immaturity, since a .young skull of G. trichopus (No. 50104) shows the same
peculiarity, though m less degree.

lAT.Jsns.] GENUS ZYGOGEOMYS. 195

. 1 k, I have been able to examine the only specimen known of this
sjKcies. It belongs to the Museo Nacional de Oosta-Kica, and was
loiiiicd Dr. Allen by Mr. George K. Cherrie, who collected it at Santa
Chira, Costa Eica, in October, 1892. It is a male, and, like the type of
cosfaricensis, is immature. It resembles the latter in having a large
pure-white patch on top of the head,* in the large size of the naked
nasal pad or callosity, and in the nakedness of the tail and feet. The
hind feet are absolutely naked; the forefeet are naked except for the
presence of a few long hairs about the toes. The color of the upper
parts is somewhat darker than in co.staricensis. The specimen is so
young that some hesitancy is felt in its generic assignment. It may
hen Ecterof/eomys instead of a 3Iacrogeomys, though this is exceedingly
improbable.

Genus ZYGOGEOMYS t nob.
(PI. 6; pi. 13, fig. 24; pi. 14, fig. 1; pi. 15, fig. 10; pi. 17, fig. 2; pi. 18, fig. 2; pi. 19, fig. 4.)

Type Zy(/ogeomys tncJiopus sp. no v., from Xahuatzin, Michoacan,
Mexico.

Generic characters. — Upper premolar with four enamel plates, the pos-
terior restricted to lingual third ; upper and lower premolars subequal in
length; shaft of upper premolar slightly convex forward.

First and second upper molars with two enamel plates each, the
posterior failing on outer side. Third upper molar an incomplete dou-
ble prism; crown nuich longer than broad; heel well developed, broad,
narrowed on outer side only; sulcus on middle of outer side; absent
ou inner side. Inner enamel plate covering two-thirds to three-fourths
of inner side of tooth, straight, reaching end of heel posteriorly; outer
enamel 2)lttte covering about half or a little less than half of outer side of
tooth, its anterior half bent strongly outward. Interspaces broadly
open, the posterior broadest, directed backward, and often forming a
sort of everted lip (fig. 27^).

Upper inclHors bisulcate; principal sulcus on inner side of median
line; minor sulcus on inner convexity (see fig. 22^ and pi. 15, tig. 10).

Cranial characters.^ — Cranium as a whole long and narrow, the zygo-
mata not widely spreading, slender, antero external angle rounded and
not expanded; zygomatic arch normally complete without jugal, the

* The white crown patch of cherriei and costaricensis was at first believed to ba
almoraial. lint the fact that the .spot is bilaterally symmetrical, and is repeated in
the only specimen known o{ Macrogeomys cosfariceiisis, which is likewise young, sug-
gests its possible permanence, at least as a mark of immaturity.

'\ Zy(io(ieomiis, with reference to the unique cliaracter of the zygomata.

! Owing to the extreme difficulty of discriminating generic from specific charac-
ters 111 animals presenting such extraordinary cranial variations as the Mexican
GeoDujida, it is thought best in descriptions of genera, of which only a single species
IS known, fo record all of the characters that seem entitled to more than specific
woiglit, The generic diagnosis here given, therefore, errs on the side of fullness.
The fill lire discovery of additional species will promptly reduce the number of
characters.

196 NORTH AMERICAN FAUNA. [no. 8.

maxillary and squamosal arms comiug in contact above it; jugal rudi-
meutary, inferior and chiefly external; rostrum long and narrow; tem-
poral impressions meeting in a short but well-developed sagittal crest;
palatine bones contracted at base of pterygoids; pterygoids vertical
lamellae as in Thoinomys, meeting or nearly meeting in median line
behind palate. Premaxilla not inclosing incisive foramina, which is
bordered posteriorly by the maxilla.

Mandible rather long and slender, much as in Geomys hursar'ms;
orbitosphenoids relatively larger than in any other genus of the fam-
ily, closing the upper part of the sphenoidal fissure (except a foramen
at apex) and ankylosed broadly with the alisphenoid (pi. 17, fig. 2), as in
some species of Thomomys; sphenoid fosste corresi^ondingly shortened,
reaching only halfway from horizontal part of alisphenoid to base of
cribriform plate; mesethmoid quadrangular, much longer than high
Cpl. IS, fig. 2) ; endoturbiuals collectively subquadrate, but with antero-
superior corner rather sharply elongated, projecting into posterior
emargination of nasoturbinal; the os planum spreading forward in
front of fourth endoturbiual about as far as length of latter {\A. 10, fig. 4).

General remarks. — Zygogeomys presents the unique combination of
distinctly bisulcate incisors with remarkably short sphenoid fossai and a
type of zygomatic arch heretofore unknown in the whole order Eodentia.
It presents further an exceptional degree of coossification of the
component elements of the skull. The occipitals, parietals, frontal,
ethmoid, squamosals, alisphenoids, maxilla, palatines, and pterygoids are
ankylosed together ; and the basisphenoid, presphenoid, and orbitosphe-
noids are ankylosed together. Furthermore, the two resulting complex
masses are firmly united by ankylosis of the orbitosphenoids with the
alisphenoids. The coossification is sometimes carried even further by
the fusion of the anterior and posterior arms of the zygoma, and the
union of the premaxilla with the maxilla and nasals. The sutures that
remain open are between the basioccipital and basisphenoid; between
the frontal on the one hand and the nasals, premaxillaries, and maxil-
lary root of the zygoma on the other; between the maxilla and
frontal anteriorly, and maxilla and alisphenoid posteriorly. The
result of these extensive ankyloses is that in old age all of the
bones of the cranium except the mandible are inseparably bound
together — if not directly in every case, then in a roundabout manner.
Zygogeomys thus occupies an anomalous position in the family.

ZYGOGEOMYS TRICHOPUS sp. nov.

(PI. 6; pi. 13, fig. 24; pi. 14, fig. 1; pi. 15, fig. 10.)

Tj/pefrom Nahuatzix, Michoacan, Mexico. No. 50107 i ad. , U. S. National Museum,
Department of Agriculture collection. Collected October 11, 1892, by E. W.
Nelson (original No. 3571).
Geographic distrihution. — The Sierra Madre of Michoacan, from Patz-
cuaro to Nahuatzin ; strictly limited to the pine zone, between the alti-
tudes of 6,800 and 9,500 feet (map 3 =).

jAN.,1895.] ZYGOGEOMYS TRICHOPUS. 197

General characters. — Size large; tail rather long, entirely naked from
base; a conspicuous naked pad at end of nose; fore feet and claws
shorter than liiud; upper surfaces of both fore and hind feet densely
covered with hair, completely hiding the skin; color very dark. Cra-
nial characters marked ; maxillary and squamosal arms of zygoma meet-
ing above the jugal, which is greatly reduced.

Color. — Upper parts varying from dark slate to rich seal-brown,
glossy, and finely mixed with a very thin wash of ferruginous, espe-
cially on the sides; underparts dark plumbeous washed with fulvous;
upper surfaces of hind feet slate-gray, sometimes varying to white; an
irregular patch of white on throat. Some specimens lack the ferrugi-
nous wash and are glossy slate-black. Some have an almost metallic
luster.

Cranial characters.* — Skull, as a whole, long and narrow; zygomatic
arches contracted, slender, not expanded at antero-exterual angle; com-
plete without jugal, which is much reduced in size, the maxillary and
squamosal arms meeting above itt ; rostrum and nasals long and narrow;
temporal impressions meeting in a short but well-develoj)ed sagittal
crest; palatine bones contracted at base of pterygoids; pterygoids ver-
tical lamellne as in Thomomys; occipital jjlane nearly vertical, about
twice as broad as high; mastoid bullne fuller and more rounded poste-
riorly than in Geomys; audital bullae of moderate size, similar to those
of Geomys hursarius; premaxilla ending below at middle of incisive for-
amina (instead of surrounding them, as usual in the family) ; postpalatal
pits rather narrow, elongated and shallow, reaching anterior plane of
last molar; mandible rather long and slender, much as in Geomys
hursarius; angular processes moderate; condylar process rather short;
coronoid process long, its tip overhanging front of condyle.

Measurements (taken in flesh). — Type specimen, $ ad. : Total length,
346 ; tail vertebrae, 115 ; hind foot, 40. Average of three adult males from
type locality : Total length, 342.6; tail vertebrae. 111; hind foot, 45.8.
Average of seven females from tyi^e locality: Total length, 322.7; tail
vertebrae, 105.8; hind foot, 42.8.

For cranial measurements see Table C, p. 209.

Specimens examined. — Total number 12, from the following localities
in Michoacan, Mexico: Nahuatzin, 10; Patzcuaro, 2.

General remarks. — Mr. Nelson found these remarkable animals pretty
generally distributed over the wooded mountain slopes except where
the timber is dense. They are most numerous about the borders of
small grassy parks and in the more open parts of the forest. In places
where the land has been cleared in these mountains they infest the culti-

* Owing to tlie circumstance that only a single species of this remarkable genua
18 known, it is unsafe to attempt to discriminate sharply between generic and spe-
cific characters. For this reason many of the characters given in the generic descrip-
tion are here repeated.
I tin some specimens the union is not quite complete.

198

NORTH AMERICAN FAUNA.

tNOig.

vated fields and do considerable damage to the corn, wheat, and pota-
toes of the Indian farmers.

Geuus THOMOMYS Max Wied, 1839.

(Text fit^s. 31% 32b, aud 68-71.)

Type TJwmomys riifescens Max Wied. Type locality iiiiknown.
Thomomya Max Wied, Nova Acta Acad. Caes. Leop. -Carol. Vol. XIX, pt. i., 1839,
377-384.

Upper and lower molars, including m^, with two enamel plates each,
one anterior aud one posterior (figs, 31^ aud 32''). Upper incisor with
sulcus normally very small and close to inner edge of tooth (fig. 23, p.
72), or absent. In a few species it is relatively hirge and deep, as in
T. monticola of Allen.

Orbital plates of frontal not meeting inferiorly behind cribriform plate
of ethmoid, but broadly separated by orbitosphenoids (fig. 71, /ro).

Figs. 68-71.— T7tomom?/s hxdbivorus. ? Salem, Oregon.

68. Vertical longitudinal section of front of skull, showing turbinated bones. For key see fig. 10.

69. Vertical longitudinal median section of skull, mesethmoid and vomer in place. For key see fig- 7.

The accompanying cuts (figs. 68-71) show the relations of the several
bones forming the floor of the brain case, and also those of the nasal
chamber, in Thomomys bulbivorns of Richardson. In this s])ecies the
incisors project much further forward than usual. The various species
differ considerably iu important cranial characters, as will be shown in
a special paper on the species of Thomomys. The geographic distribu-
tion of the group as a whole is shown on n)ap 1, A.

I

JAN., 1895.]

THE GENUC THOMOMYS.

199

Externally Thomomys differs from all tlie otlier genera of the Geo-
myida' in tlie relatively small size of the fore feet. In this respect, and
in the faint sulcation of the incisors, the presence of two enamel plates
oil each of the molars, above and below, and in numerous cranial char-
acters it is much less highly specialized than most members of the
family.

70. Thomomys bulbivonis, from Salem, Oregon. Skull from above ; vault of cranium sawed ofFto show
floor of brain ease. For key see fig. 9.

71. Anterior part of floor of brain case, much enlarged. (Same specimen as flg. 70.)
ale Anterior opening of alisphenoid canal.

as Alisplienoid l>oue.

bs Basisi)beiioi<l.

cr (/ribriform plate of ethmoid,

/(• Frontal.

fro Orbital or descending plate of frontal. It should be observed that this plate does not meet
its fi^llow inferiorly behind the cribriform plate as in most of the other genera.

of Optic fdriunrn.

OS Orl)itiis]ili(n<iid.
ptf Pterygoid fossa.

sf Upper iiart of sphenoidal fissure.

APPENDIX

(A ) Status of Geomys mexicanus Auct.

The earliest description that I have seen of any member of tlie
family GeomyUliv was ])ublislied by Fernandez in 1051, and relates to
a Mexican aninial called by him the Tucan or Indian mole.* Nearly a
century and a half later Kerr bestowed the name Borex mexicanus upon
Fernandez's Tucan without having seen a specimen (Kerr, Animal
Kingdom, 1792, 207-208). It is not surprising that Kerr followed
Fernandez and Button in placing the animal among the moles,t misled
by its projecting incisors and habit of throwing up little mounds of
earth along the course of its subterranean galleries.

The animal seems to have been first referred to the genus Geotmjs by
LeConte in 1852 (Proc. Phila. Acad. Kat. Sciences, 1852, p. IGO).

In 1827 Lichtenstein described, under the name Ascomys mexicanus^
three specimens of pocket gophers collected by Deppe on the table-
land of Mexico, but the exact locality whence they came is unknown
(Brants Muizen, 1827, 27-31). The specimens differed greatly among
themselves in color, as originally described by Lichtenstein, and their
cranial measurements, kindly famished me by Dr. Matschie, show that
they belong to at least two different genera. The case as it stands,
therefore, seems to be as follows : Lichtenstien's mexicanus is composite |

*Following is a trauslation of the original description: "On the Tucan, or a
certain kiml of Indian mole. Chap. xxiv. [The Tncan] is apparently a species of
mole 9 inches in length, and e(iualing the humerus of man in size; it is fleshy, fat,
and furnished with such short legs that it almost touches the ground with its helly ;
hair, fulvous; tail, short; claws and nails, long; snout, murine; ears, small and
round; front [teeth], two above and same in number below, considerably exserted
and curved inward; [the other teeth], though much smaller, are very strong. When
fat the flesh is edilde, of pleasant taste, but causes stupor. ' ■ *."— (Francisco
Fernandez, Historiic Auimalium et Miiicralium Nova' Hispania;, Liber i, 1651, pp. 7-8.)

tAU the American moles were at that time placed with the shrews in the genus
■S'ojTJT, the genera Scalops, Scapanits, and Condijlura not having been proposed until
souietimc later.

t From the cranial measurements kindly furnished me by Dr. Matschie, and now
for the tirst time published, it is evident that one of Lichteustein's specimens was a
Platygeomys closelj related to, if not identical with, the animal here described as
■P. planiceps.

201

202

NORTH AMERICAN FAUNA.

[NO. 8.

and is preoccupied by mexicanus of Kerr (1792). The latter is unident-
ifiable, the vague description applying equally well to several species.
It being- clearly impossible to use the name mexicanus, it should be
dropped from the group.

Cranial measurements of hco of Lichten stein's ti/pe specimens of Ascomys mexicanus.
[Measured by Dr. Paul M.itschie.]

1558.

1559.

Greatest basal length (condyle to front of premaxilla) .

Basal length (basion to gnathion)

Basilar length ot Hensel (basion to alveolus of incisor).

Greatest zygomatic breadth

Gre;;te.st breadth posteriorly across squamosals

Least breadth between postglenoid notches

Least interorbital breadth

Height of cranium aliove palate

Height of cranium above basion

Length of upper molar series on alveoli

Length of diastema

Length of single mandible without teeth

Breadth across angular processes

Distance trora condyle to end of angular process

Breadth across muzzle just in front of zygoma

14.5

25.5

42.5

54

20

13

55
52
48
35.5
40
28
9 5
20
17
13
21
36
37
16
11

(B.) Tables of Average Measurements of the Various

Species.

Average measurements of the species of Geomys.
[All measurements are in millimeters .and from fresh .specimens.]

Name of species.

G. bursarius .

O. hitescens . .
G. breviceps . .

G. sagittalis . .

6. attwateri. .

G. texensis . . .
6. arenarmt .
G. personatus
G. fallaz

G. tuza

G. mobilensis
G.Jiondanus.

Locality.

Southeastern North Dakota. . .

Elk River, Minnesota

Hunter and Williamsville, Mis-
souri

Western Nebraska*

Childress, Texas

Mer Eouce, Louisiana *

Benton, Arkansas

Fort Gibson, Indian Territory.

Miueola, Texas .*. .

Molano, Texas

Galveston Bay, Texas '

Houston, Texas

Kockport, Aransas County,
Texas * ." . .

Mason, Texas *

El Paso, Texas *

Padre Island, Texas*

South side Nueces Bay, Corpus
Christi, Texas *. .. '

Augusta, Georgia *

Butler, Georgia^

Mobile Bay, Alabama *

San Mateo, Florida

Number of
specimens.

! <^

Total
length.

Tail
VertebrsB.

Hind foot.

40

20

20 :

12

4

8 ■

22

12

10

10

4

0 1

38

15

23

7

4

3

14

5

9

14

6

8

11

5

6

20

5

15

9

5

4

17

10

7

''8

32

8

24

13

4

9

13

9

10

19

10

9

10

5

5

8

4

4

6

3

3

296

284

256
270
257
231
243
233

265
243

223
246
228
213
206
209

220.5, 193.
216.2 206
220 I 196
226 208

255
(t)
260
315

263
269
257
250
288

220

;250
293

236
249
241
229
235

78

37

74 63

84 72

81. 5 08

70 I 61

74 I 66. 3

68. 2i 61.7

67. 8, 57. 2

63. 8: 60. 3

64 I 54

64 57

80

83
111

33
33.5

30

28

29

27.8

26.3

26.4

26

25

30

34

30

31.5

28

26.5

27

26.3

24.1

24.5

23

24

28

32
40

34 '
34.4
33.8
33.6
35.5

t32
36

' 31
32

3i'.6
30.5
33

'Type locality.

(Average of 28 specimens of both sexes: total length, 210; hind foot, 28.

ISome of the .specimens of arenarius recorded as females are very large and were probably m.'ileB;
hence the averages here given for females ari' probably too gieat.
§The specnuens from Butler. Ga., are clearly intermediate between tuza and mobileiuis.

JAN., 1895.]

AVERAGE MEASUREMENTS.

203

Average measurements of the species of Craiogeomys.

LAll measurements are in millimeters and from fresh specimens.]

Locality.

Number of
specimens.

Total
length.

Tail
vertebrae.

Hind foot.

Name of species.

3

o

H

d

?

d

?

d

?

d

?

C. merriami

Valley of Mexico, Mexico

18
7
3

12
8
1
1
4
3

11
4

4

"i

7
3
3
12
4
1
1
3
3

380
328

'sis'
'295'

344
289
324
310
277
318
304
256
259

113
94.5

'89'"

"95""

105
85
91
88
75
92
87
77
77

50

47

"'42'

"37'

46
43 5

42 6

C. perotensis

Cofre de Perote, Mexico

41.5
37

C. oreoeetes

C. peregrinug

C. castanops

Mount Popocatapetl, Mexico . .
Mount Iztaccihuatl, Mexico...

43
42
33

Albuquerque, New Mexico

34

C. castanops gold

mani

C. fulveicens

Cauitas, Zaeatecas, Mexico....
Chalchicomula, Puebla, Mexico.

3
9

"3'

3

6

327

257
302

ios '

82.7
97

"43"

34.3
39.6

Average measurements of the species of Platygeomys, Orthogeomys, Heterogeomys, Pappo

geomys, and Zygogeomys.

[All measurements are in millimeters and from fresh specimens.]

Xocality.

Number of
specimens.

Total
length.

Tail
vertebr*.

Hind
foot.

Name of species.

1
^

cT

9

d

?

d ?

d

9

Platygeomys gymnu-
run.

P. tylorhinus

P. planiceps

Zapotlan, Jalisco, Mexico . . .

Sierra Nevada de Colima,
Jalisco, Mexico.

Tula, Hidalgo, Mexico

Patzcuaro, Michoacan, Mex-
ico.

N. slope Vole. Toluca, Mex-
ico, Mexico.

Colima City, Mexico

Cerro San Felipe, Oaxaca,
Mexico.

Mount Zempoaltepec, Oax-
aea, Mexico.

Jico, Vera Cruz, Mexico

Motzorongo, Vera Cruz,

Mexico.
Chichicaxtle, Vera Cruz

(type), Mexico.
Sierra Nevada de Colima,

Jalisco, Mexico.
Guadalajara, Jalisco

Nahuatzin, Michoacan

6

2

3
5

3

10
10

3

5

14
1
6
1

10

3

1
3

1

7
2

2

2

4

2

3

3
2

2

2

2
3

8

1
3

10
1
4
1
7

353

345
348

372

287.5
369

416

345

348

236

343

322

298
331.5

336.5

277
360

380

311

317

323

216

226

323

105 9:

85

100 91.5
101.5 91.5

121 100

82 7.1

53.5

45
49.5

46

42
50

54

53

49

33

46

49.5

49

39.5
45.5

43

39 5

Orthogeomys sealops . .

Orthogeomys nelsoni. .

Heterogeomys hispi-
dus.

103.5
131

92

96. 5

81
111

109
118

85

81.5

88

72

68
106

50
52
47
45.5

Pappogeomys bulleri .

Pappogeomys albi-

nasus.
Zygogeomys trichopus

52
30
31
43

204

NORTH AMERICAN FAUNA.

[no. 8.

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CRANIAL MEASUREMENTS.

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210

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CRANIAL MEASUREMENTS.

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214

NORTH AMERICAN FAUNA.

[NO. 8.

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INDEX.

[Kames of genera and species are in heavy type.]

Adaptation to subterranean life. 15.
Alispbenoid bone, 43-45.
canal, 36,43.
Ascomys (synonym of (.eoiiiys), 109.
canadensis, 120,
niexirauus 2ul .
Basioccipital, 40.
Basisphenoid, 43.
Brain case, floor, 35.
Callosity, nasal, 16.
Canal, alispbenoid, 43.

infraorbital, 39, 55.
Cheek pouches, 18.

muscles. 101.
Color phases, 19.
Cranial variations, 63-68.
Cranium. (-SVe Skull.)
I'ratogeonijs :

Genus, defined, 150-151.
Key to species, 151.
Cratogeoniy i'astanoi)s, 159, 160.

cranial m e a s u r e -
meiits, 211.
castanops goldmani, 160,161.

cranial iiieas-
uremeuts, 211.
estor, 155, 156.

cranial measurements, 210.
fulTeseens, 101,162.

cranial measure-
ments, 211.
merrlanii, 151-153.

cranial measure-
ments, 210.
creocetes, 156, 157.

cranial measurements,
211.
peregrinus, 158, 159.

cranial measurements,
211.
perotensis, 154, 155.

cranial measurements,
210.
Cribriform plate, 50.
Dental armature, 69.
Digastric muscle, 100.
Diplostoma (synonym of (ieonijs), 109.
fusca, 120.

Dynamics of cutting machine as a whole, 88-97.
incisors, 89, 90.
molars, 90-93. .
Ectoturbinal, 52.
Enamel in incisors, 70.
in molars, 78-83.
in premolars, 78.
Enamel cap in young teeth, 84-87.
Enamel organ, 87.
Eudoturbinal, 52.
Ethmoid, 50-52.
Exoccipital, 41.
Feet, 15.

Fissui-e, sphenoidal, 36.
Floor of brain case, 35.
Food, 19.

treatment of, 98.
Foramen ovale, 39, 45.

rotundum, 39,45.
Fossa, olfactory, 35.
pterygoid, 36.
sphenoid, 35.
spheno-pterygoid, 36.
Frontal, 49, 65.

descending plates, 35, 49.
Geographic distribution, 26.
Geoinys:

Genus defined, 109-112.
Key to species, 113.
Oeoniys arenarius, 139-141.

cranial measurements, 207.
breviceps, 129-133.

craiiial measurements. 205.
breviceps attwateri, 135-137.

cranial measure-
ments, 205.
breviceps sagittalis, 134.

cranial measure-
ments, 205.
bullori, 147.

cranial measurements, 214.
bursarius, 120-127.

cranial measurements, 204.
canadensis, 120.
castanops, 159.

cranial measurements, 211.
clierriei, 194.

cranial measurements, 215.

217

218

INDEX.

(■eoinys cinerea, 120.
clarkii, 159.
f^niosu8, 170.

cranial measurements, 2i;t.
grandis, 175.

cr.anial measurements, 214.
gymnurus, 164.

cranial measurements, 212.
heterortus, 186.

cranial measurements, 215.
hispidus, 181.

cranial measurements, 215.
lutescens, 127-129.

cranial measurements, 204.
nierriaiiii, 152.

cranial measurements, 210.
mexicanus, 201-202.

• ci'anial measurements, 202.
nelsoiii, 147.
oregoneiisis, 120.
per!«onatu8, 141-144.

crani.al measurements, 206.
fallax, 144-145.

cranial ineasurements, 206.
pinetis, 113.
scalops, 174.

cranial measurements, 214.
texensis, 137-139.

cranial mea-surements, 206.
tuza, 113-115.

cranial measuremeuts, 208.
tuza floridanus, 115-119.

cranial measurements, 208.
tuzaiiiobilensis, 119-120.

cranial measurements, 208.

Heterogeomys:

Genus defined, 179-180.
Key to species, 180.
Heterogeoniysi liispidus, 180-183.

cranial measureuu^nts. 215
torridus, 183-185.

cranial measurements, 215.
Incisor teeth, 70-72.

in young, 83.
Infraorbital canal, 39.
Interparietal, 41-43.
Jaw, 60.

movements, 102-103.
muscles, 98-101.
stroke, 97.
Jugal, 57.
Key to genera, 23.

Key to species of Cratogeoniys, 151.
fteomys, 113
Heterogeomys, 180.
Macrogeomys, 186.
Orthogeomys, 173.
Pappogeomys, 147.
Platygeomys, 164.
Lachrymal, 57.
Lambdoid crest, 39.
Macrogeomys :

Genus defined, 185-186.
Key to species, 186.

Macrogeomys clierriei, 193-195.

cranial measurements, 215.
costaricensis, 192-193.

cranial me asu re-
ments, 215.
dolichocephalus, 189-192.

cranial measure-
ments, 215.
tieterodns, 186-189.

cranial measurements,
215.
Mandible, 60.

Masseter muscle, 99-100.

influence on skull, 105-107.
influence on teeth, 107-108.
Mastoid bulla, 33, 46-60.

process, 60.
Maxilla, 54-56.
Maxillo-turbinal, 34.
Meatus auditorius, 33-59.
Mesethmoid, 50.
Molar teeth, 74-83.

in young, 85.
Morphology of skull, 33-63.
Month, division into two chambers, 17.
Mns biirsarins, 120.
ludoricianus, 120.
saccatus, 120.
tuza, 113.
Muscles, cleido-mastoid, 102.
digastric, 100.
external pterygoid, 100.
internal pterygoid, 100.
latissimus dorsi, 102.
masseter, 99-100.

influence on skull and teeth,
104-108.
rhomboideus, 102.
sterno-niastoid, 102.
temporal, 99.

transverse mandibular, 100-101.
trapezius, 102.
Muscles of cheek pouches, 101.
head and neck, 102.
jaw, 98-101.
Xarial passage, 39.
Kasal Itones, 57-58.
callosity, 16.
Nasoturbiual, 34.
Occiput, 05.
Orbitosphenoid, 48-49.
Orthogeomys :

Genus defined, 172-173.
Key to species, 173.
Orthogeomys grandis, 175-176.

cranial measurements, 214.
latifroiis, 178-179.
nelsoni, 176-178.

cranial measurements, 214.
scalops, 173-^75.

cranial measurements, 214.
Os planum, 50.
Osteodentine, 87.
Palate, 34.
Palatine bones, 53.

INDEX.

219

Palato-pterygoid jdate, 34.

Pappogeoiiiys:

fc. Genus defined, 145-147.

Key to species, 147.
Pappou:eoiiiys albiiinsus, 147.

cranial measurements,
214.
biiUeri, 147-149.

cranial mea.surements. 214.
Parietal, 46.

Paroccipital process, 3.5.
Periotic capsule, 58-60.
Phylogenetic tree, 24.
Platygeomys:

Genus defined, 102-164.
Key to species, 161.
Platygeoinys fumosus, 170-171.

cranial measurements. 213.
gyniniiruis, 164-160.

cranial measurements,
212.
plaiiioeps, 108-170.

cranial measurements,
212.
tylorliinus, 167-168.

cranial raeasnreraents,
212.
Postorbital process, 35.
Pouches, cheek, 18.

method of filling, 18.
muscles, 101.
Premaxilla, 56.
Premolars, 72-74.

deciduous, 83.
permanent, in yoiing, 84.
Presphenoid, 37, 47.
Process, angular, 67.
mastoid, 60.
paroccipital, 35.
postorbital, 35.
Progression backward, 16.
Psendostoma (synonym of Oeoniys), 109.
bursarius, 120.
rastaiiops, 159.
floridana, 115.
Pterygoid bones, 52-53.
muscles, 100.
plate, external, 36, 54.
Saccophorns (synonym of Geoiiiys), 109.

bnrsarius, 120.
Sagittal area, 39.
crest, 39.
Skull, alisphenoid bone, 43-45.
alisphenoid canal, 36, 43.
basioccipital l)one, 40.
basisphenoid bone, 43.
bony palate, 34.
changes with age, 61-63.
coossification of paired bones, 63.
cribriform plate, 50.
ectoturbinal bones, 52.
endoturbinal bones, 52.
ethmoid bone, 50-52.
exoccipital bone, 41.
external pterygoid plate, 54.
floor of brain case, 35.

Skull, foramen ovale, 39, 45.

foramen rotundum, 39, 45.

frontal bone, 49.

infraorbital canal. 39 55.

interparietal bone, 41-43.

jugal bone, 57.

laclirymal bone, 57.

lambdoid crest, 39.

list of bones, 40.

mandible, 60.

mastoid bulla, 31.

maxilla, 54-56.

mesethmoid, 50.

morphology, 33-63.

narial passage, 39.

nasal bones, 57-58.

orbitosphenoid bone, 48-49.

OS planum, 50.

palatine bones, 53.

palatopterygoid plate, 34.

parietal bones, 46.

paroccipital process, 35.

periotic capsule, 58-60.

postorbital process, 35.

premaxilla, 56.

presphenoid bone, 47.

pterygoid bone, 52-53.

pterygoid fossa, 36.

sagittal crest, 39.

.sphenoid fossa, 36.

sphenoidal fissure, 36.

spheno-pterygoid fossa, 36.

squamosal bone, 45.

supraoccipital bone, 41 .

tentorium (absent), 39.

turbinals, 34, 57.

turbinated bones, 34, .57.

tympanic bulla, 33.

tynipano-periotic capsule, 58-60.

vomer, 52.

vomerine sheath, 56.

zygomatic arch, 34.
Sorex mexicanus, 201.
Species, number and distribution, 27.
in Mexico, 30.

United States, 28.
Sphenoid bones :

Alisphenoid, 34, 43-45.
Basisphenoid, 33, 43.
Orbitosphenoid, 34, 48-49.
Presphenoid, 33, 47.
Sphenoul fossa, 35.
Sphenoidal fissure, 36.
Spheno-pterygoid fossa, 36.
Squamosal, 45.
Supraoccipital, 41.
Tail, an organ of touch, 16.
Teeth, 69-88.

changes from wear, 86-87.

dental formula, 69.

dynamics, 88-97.

of cutting machine as a whole,
88-97.
incisors, 89,
molariform teeth, 90-97.

enamel, 78-83.

220

INDEX.

Teeth, enaDiel, dynamics, 93-97.
incisors, 70-72.

dynamics, 89.
young, 83.
manner of attachment, 88-90.
molars, 74.

dynamics, 90-93.
enamel plates, 78-80.
young, 85- 87.
last upper molar, 76-77.
enamel, 78-83.
premolars, 72-74.

deciduou.s, 83.
enamel plates, 78.
permanent young, 84.
Temporal impression-s, 39, 42.

muscle, 99.
Tentorium (absent), 39.

Thoiiioniys, 198-199.

Tongue, 18.

Turbinals, 57.

Tympanic bulla, 33, 58-59.

Tympano-periotic capsule, 58-60.

Typo localities, 25.

Variation :

Cranial, 03-68.

Individual, 21.

Seasonal, 20.

Sexual, 20.
Vomer, 52.

Vomerine sheath, 56.
Zygogeomys, genus defined, 195-196.
Zygogeomys trichopus, 196-197.

cranial measurements, 209.
Zygomatic arch, 34.

222

NORTH AMERICAN FAUNA.

platj: 1.

(All natural size.)

Geoviyshur sari IIS (Shaw). Knoxville, Iowa.
(No. 2772 (? ad. Merriam collection.)

North American Fauna, No. 8.

Plate I

Senj. Mortimer, del

Geomys bursarius (Shaw)
Knoxville, Iowa.

No.2772. tfad.

B Meisel, pf^tc kch

224 NORTH AMERICAN FAUNA.

PLATE 2.

(All natural size.)

Cratogeomijs merriami (Thomas). Lerma, Mexico.
(No. 50110 S ad. U. S. Nat. Mus.)

North American Fauna, No. 8.

Plate 2.

¥at .size.

Geomys merriami Thomas
Lerma, Mexico.

No. 50110. o- ad.

B Metsel. pAete. HA.

226 NORTH AMERICAN FAUNA. [no. 8.

PLATE 3.

(All natural size.)

Platiifjeoinys fiymnurus Merriam. Zapotlaii, Jalisco, Mexico,
(,No. 45611 ^ ad. U. S. Nat. Mus.}

North American Fauna, No. 8.

Plate 3

B Mflzssl. photo liin

Geomys gymnurus Merriam

Zapotlan, Mexico.

No.^SSII.a" ad.

228 NORTH AMEKICAN FAUNA. [no.8.

PLATE 4.

(All natural size.)

Heterogeomys hispidus (LeCoute). Jico, Vera Cruz, Mexico.
(No. 55343 U. S. Nat. Mus.)

North American Fauna, No. 8.

Plate 4

Ifa^t.i

B-M'ftset, photo litk

Geomys hispidus. LeConte
Jico.Vera Cruz. Mexico.

No, 55343. <? ad.

230 NORTH AMERICAN FAUNA. [no. 8.

PLATE 5.
(All natural size.)

Macrogeomys dolichocephahis sp. nov. San Jose, Costa Rica.
(No. 36295 (? ad,, U. S. Nat. Mus.)

North American Fauna, No. 8.

Plate 5

BMezseLpka'aMA

GEOMYS D0LICH0CEPHALU5 sp.nov.
San Jose', Costa Rica.

No 36295.

232 NORTH AMERICAN FAUNA. [no. 8.

PLATE 6.

(All natural size.)

Zygof/eomys irlcliopus sp. nov. Nalmatziu, Micboacaii, Mexico.
(No. 50107 i ad., U. S. Nat. Mus.)

North American Fauna, No. 8.

Plate 6

Geomys trichopus Merriam
Nahuazin.Michoacan, Mexico.

B Affzsei. pho^o Hch

234 NORTH AMERICAN FT^ANA. [no.8.

PLATE 7.
(All iKitnral size.)

1. (ieomjia tma (Ord) ^ ad. Augusta, Ga. (Type locality).

(No. 58639 U. S. Nat. Mus.)

2, 5.6. (i. Iiiza mohihnsis ^ ad. Mobile Bay, Alabama. (Tyjte locality

(No. 46024 U. S. Nat. Mus.)
3 and 4. G. tuza fioridaintu ^ ad. 8an Mateo, Fla.

(No. 6512 c? ad. aud 6514 ^ old, Merriam collection.)

North American Fauna, No. 8.

Plate 7

I.GEOMYS TUZAo- 2,5&6.G.MOBILE:NSISo- 3&4.G.TUZA FLORIDANUSo-

236 NORTH AMEHICAN FAUNA.

platp: 8.

(All iiaturul size)

1 &■ 2. Cralogeomys oreocefes sp. nov. 9 iul. Mount I^opocatapetl, Mexico. Ti/jje.

(No. 57963 U. S. Nat. Mus.)
3. C. pcretjrinu.s HI), iiov. 9 ad. Mount Iztaccilmatl, Mexico. Tii2)e.

(No. 57964 U. S. Nat. Mus.)
4 & 5. C. estor sp. uov. Las Yigas, YGVix i'vnz, Mexico.

(4 = No. 54306 9 ad. and 5 =54308 <? ad. U. S. Nat. Mus.)
6. C. perotens'is sp. uov. 9 ad. Cofre de Perotc, Vera Cruz, Mexico.

(No. 54299 II. S. Nat. Mus.)

North American Fauna, No. 8.

Plate 8

B Meisel. phtih Uch

I 8:2 I ad. GEOMYS OREOCETES
4S&5c'G. ESTOR

3 ? ad. G. PEREGRINUS
6 ? ad. G. PER0TEN5I5

238 NORTH AMERICAN FAUNA. [m.8.

PLATE 9.
(All uiitiiral size.)

1. Geoniys arenariiis ^ ad. El Paso, Texas.

(No. 58339 U. S. Nat. Mus.)

2. G. texensis $ Mason, Texas.

(No. 4161 Merriam collection.)

3. G. atttvateri ^ ad. Rockport, Aransas County, Texas.

(No. 51382 U. S. Nat. Mus.)

4. Gr. sagittalis ^ ad. Galveston Bay, Texas.

(No. 44957 U. S. Nat. Mus.)
5 & 7. G. lutescens ^ ad. Cherry County, Nebraska.

(5 = 25640 <? yg. ad.; 7=25471 <? old, U. S. Nat. Mns.)
6. G . breviceps $ ad. Mcr Rouge. Louisiana.

(No. 46673 U. S. Nat. Mus. )

8. G. biirsarhis 9 3,d. Knoxville, Iowa.

(No. 2024 Merriam collection.)

9. <T, bitrsarius $ ad. Knoxville, Iowa.

(No. 2625 Merriam collection.)

North American Fauna, No. 8.

Plate 9

BMfZsei. phatc lith

I.GEOMYS ARENARlUSo-ad. 2.G.TEXENSISo-ad. 3.G. ATTWATERIo- ad. 4.G.SAGnTALISo-ad.
5&7.G. LUTESCENSo-ad 6,G. BREVICEPSo'ad. 8 5 ad.9o-ad. G. BURSARIU5

240 NORTH AMP:RICAN FAUIn^A. lNo.8|

PLATE 10.
Under side of mandible.

(All natural size.)

1. Geomys iuza flnridauus (Baelnnaii). San Mateo, Florida.

(No. 6511 (? Merriam collection.)

2. G. tuzamobilcnsis HT[>. nov. Mobile Bay, Alabama.

(No. 46023 (? U. S. Nat. Mns.)

3. Cralof/eomys oreocetes sp, nov. Mount Popocata]ietl, Mexico.

(No. 57963 5 U. S. I\at. Mus.)

4. C. peregrinits sp. nov. Mount Iztaccilmatl, Mexico.

(No. 57964 $ U. S. Nat. Mus.)

5. C. merriami (Thomas). Amecameca, Mexico.

(No. 57970 <? U. S. Nat. Mus.)-

6. Geomys btirsariiis (Shaw). Knoxville, Iowa.

(No. 2772 (J Merriam collection.)

7. Macroyeomys dolichoccphalus sp. nov. Sau .lose, Costa Rica.

(No. 36295 ^ U. S. Nat. Mus.)

8. Platyyeomys yymnuriis Merriava. Zapotlan, .Jalisco, Mexico.

(No. 45611 J U. S. Nat. Mus.)

North American Fauna, No

Plate 10

IGE0MY5TUZAFL0RIDANU5/ 2 G. M0BILENS15 • 3.G.0RE0CETES i 4.G.PEREGRINUS ^
5.G.MERRIAM[o' 6.G. BURSARlUSo- 7. G.DOUCHOCEPHALUSn- 8.G.GYMNURU5 •

242 NOltTU AMERICAN FAUNA. [no.8.

PLATE 11.

(All inilnral size.)

1. VappoaeonuiH hnUeri (Thomas), ^^ierra Nevada de Coliiiia, .Jalisco, Mexico.

(No. 4.5(>22 i U. S. Nat. Mus.)

2. MacrofjeomijHheterodus (VateTH). Costa Kica, Mexico.

(No. c? U. S. Nat. Mus.)

3. HeterogeoiiiijH costaricenHis sp. nov. Pacuare, Costa Rica.

(No. 225.51, sex ?, U. S. Nat. Mus.) Tyj)'!.

4. Plaiyf/eomnHfumosus Merriam. Coliuia City, Mexico.

(No. 4.5211 <? U. S. Nat. Mus.)

5. Ortliof/eomijs latifrons fill. nov. Guatemala.

(No. , sex ?, U. S. Nat. Mus.) Tt/pe.

6. O. latifrons (uuder side of maiulil>le of sauie skull as 5.)

North American Fauna, No. 8.

Plate 11

l^Mu3ler,cl€i,

B.ifeiseU ph^to Int^

1.GE0MY5 BULLERI - 2. G. HETER0DU5 '/ 3. G . C05TARICEN 51 5
4. G. FUMOSUS- 5 & 6. G. LATIFRONS

244 NORTH AMERICAN FAUNA. Ino.8.

PLATE 12.

(All natural size.)

Cratogeomys castanops (Baird). Las Auiiuas, Colorado. (Type locality.)

(No. 47368 ^ U. S. Nat. Mus.)

1". Basioccipital of same specimen.
Cratogeomys fulvescens sp. nov. Chalcliicomula, Mexico. (Type locality.)

(No. 53498 <? U. S. Nat. Mus.)

2". Basioccipital of same specimen.
Geomys persovai us fallax suhs]). nov. Corpus Christi, Texas. Ty2}e.

(No. 43845 ^ ad. U. S. Nat. Mus.)

3". Left audital bulla of same skull.
Geomys personatus Tvne. Padre Island, Texas. (Type locality.)

(No. 43294 ^ U. S. Nat. Mus.)

4*. Left audital bulla of same skull.

North American Fauna, No. 8.

Plate 12

I. GEOMYS CA5TAN0P5
3. G. PERSONATUS FALLAX

2. G. FULVE5CEN5
4. G. PER5 0 NATUS

246 NORTH AMERICAN FAUNA. [no. 8.

PLATE 13.
Left zygoma, showing variations in jugal bone.

(All natural size.)

1. riatygeomys tylorhinm sp. nov. Patzcuaro, Mexico.

(No. 47183 S U. S.Nat. Mus.)

2. r. gymnnrus Merriam. Zapotlan, Mexico.

(No. 45611 (? U.S. Nat. Mus.)

3. P. plan iceps s]^. nov. Tula, Hidalgo, Mexico.

(N0.559U6 <? U.S.Nat. Mus.)

4. Cratogeomys merriami (Thomas). Lerma, Mexico.

(No. 50110 <? U.S.Nat. Mus.)

5. C. perotensis sp. nov. Cofre de Perote, Mexico.

(No. 54295 9 U.S. Nat. Mus.)

6. C. estor sp. nov. Las Vigas, Mexico.

(No. 54308 <? U.S. Nat. Mus.)

7. C. estor sp. nov. Las Vigas, Mexico.

(No. 54306 9 U. S. Nat. Mus. )

8. C. oreoceteH sp. nov. Mount Popocatapetl, Mexico.

(No. 57963 9 U. S. Nat. Mus.)

9. Geomys fnza (Ord). Augusta, Georgia.

(No. 63842 S U. S.Nat. Mus.)

10. G. tuza floridanus (Aud. and Bach.). San Mateo, Florida.

(No. 6514 ^ Merriam collection.)

11. G. buraarius (Shaw). Knoxville, Iowa.

(No. 2624 (? Merriam collection.)

12. G. texensis sp. nov. Mason, Texas.

(No. 4161 ^ Merriam collection.)

13. G. arcnarhi8 sp. nov. El Paso, Texas.

(No. 25015 J U. S. Nat. Mus.)

14. G. personatusTrwd. Padre Island., Texas.

(No. 43294 ^ U. S. Nat. Mus.)

15. Papjmgeomys hulleri (Thomas). Sierra Nevada de Colima, Mexico.

(No. 45618 9 U. S. Nat. Mus.)

16. Orthogeomys latlfrons sp. nov. Guatemala. Type.

(No. U. S. Nat. Mus )

17. Cratogeomyscastanops {Y^a^x^). Las Animas, Colorado.

(No. 47368 <? U. S. Nat. Mus.)

18. Macrogeomysheterodtis (Peters). Costa Rica.

(No. U. S. Nat. Mus.)

19. Macrogeomys dolichocephalus sp. nov. San Jose, Costa Rica.

(No. 36295 <? U. S. Nat. Mus.)

20. Heterogeomys Mspidua (LeConte). Jico, Vera Cruz, Mexico.

(No. 55343 $ U. S. Nat. Mus.)

21. Heterogeomys torridus sp. nov. Guatemala.

(No.. — S U. S. Nat. Mus.)

22. Macrogeomys eherriei (Allen). Santa Clara, Costa Rica.

(No. 664 im. Costa Rica Nat. Museum.)

23. Macrogeomys costaricensis sp. nov. Pacuare, Costa Rica.

(No. 22551 im. U. S.Nat. Mus.)

24. Zygogeomys Irichopua sp. nov. Nahuatzin, Michoacan, Mexico.

(No. 50107 c? U. S. Nat. Mus.)

North American Fauna, No. 8.

Plate 13

v^^^^^

FMuOer,

dil

Nat size

B Meisel phah bh

I.GeOMYS TYL0RHINU5 o"

9.

TUZA d"

17.

CA5TAN0P5 .^•

2.

GYMNURU5 /

10.

TUZA FL0R1DANU5 -f

18.

HETERODUS c

3.

PLAN1CEP5 S

11.

BUR5AR1US /

19.

D0LICH0CEPHALU5 /

4.

MERRIAMl d"

12.

TEXEN5IS d-

20.

HISPIDU5 ..•

5.

PER0TEN51S J

13.

ARENARIU5 c^

21.

HISPIDUS .(form)

6.

E5T0R >/

14.

PERSONATUSo'

22.

CHERRIEl

7.

EST OR ?

15.

BULLERI V

23.

C05TARICENSI5

8,

OREOCETES ?

16.

LATIFR0N5

24.

TRICH0PU5 d"

248 NORTH AMERICAN FAUNA.. [no. 8.

PLATE 14.
Posterior molars and palatopterygoids.

(All double natural size.)

1. Zi/f/of/eomys trichopus sp. nov. Nahuatziu, Michoacan. Mexico.

(No. 50107 c? U. S. Nat. Mus. )

2. Geouii/s hi(rsariiis (Shaw). Knoxville, Iowa.

(No. 2624 cJ Merriain collection. )

3. Macrof/eojnys helerodas (Peters). Costa Rica.

(No. U. S. Nat. Mus. )

4. Gcomys pemonatns True. Padre Island, Texas.

(No. 43294 (? U. S. Nat. Mus. )

5. Geinnijii peraonatun fallar subsp. uov. Corpus Cliristi, Texas.

(No. 43292 9 U. S. Nat. Mus. )

6. Cratogeomiis castdiiopn (I5aird). Las Animas, Colorado,

(No. 47368 c? U. S. Nat. Mus. j

7. Cralogeomijs mvrriami (Thomas). Lerma, Mexico.

(No. 50110c? U. S. Nat. Mus. )

8. Flatyfjeomiis fumosus Merriam. Colima, Mexico.

(No. 45213 c? U. S. Nat. Mus. )

9. Plat\igcomy>i planicepn sp. nov. Volcan Toluca, Mexico,

(No. 55906 i U. S. Nat. Mus. )

10. Macrogeomi/8 cosfaricciD^iii sp. nov. Costa Rica. Type.

(No. 22551 U. S. Nat. Mus.)

11. rappogeoniys httlleri (Thoiuas). Sierra Nevada de Colima, Jalisco, Mexico,

(No. 45618 5 U. S. Nat. Mus. )

12. Ileteroyeomyn hispidus (LeConte). .lico. Vera Cruz, Mexico,

(No. 55017 9 U. S. Nat. Mus. )

13. Geomys texeiisis sp. nov. Mason, Texas.

(No. 4168 9 Merriam Collection. )

14. Geomys liitmccns Merriam. Woodward, Oklahoma.

(No. 48566 (? U. S. Nat. Mus. )

15. Geomys tn~a inobilensis sp. nov. Mobile Bay, Alabama.

(No. 46025 J U. S. Nat. Mus. )

16. Geomys tuza jloridanus (And. and Bach.). San Mateo, Florida.

(No. 6511 1? Merriam Collection. )

North American Fauna, No. 8.

Plate 14

i ^^

GeOMYS TR1CH0PU5 ITOb.

BUHSARius (Shaw)
HETER0DU5 Peters

PER50NATU5 Tfue
PER50NATU5 FALLAX nOb.
CA5TAN0P5 Bait'd

MERRiAMi Thomas
FUM05US Merriam

B Afeisel. pho to U(h

9. G. PLANICEP5 nob.

10. C05TARICEN5I5 nOb.

11. BULLERi Thomas

12. HI5P1DU5 LeConte

13. TEXEN5I5 nob.

14. LUTESCENS MeiTiam

15. M0B1LEN5I5 nOb.

16. TUZA FLORlDANUS(BaCl"llTian)

2r)0 NORTH AMERICAN FAUNA. [no. 8. i

\

PLATE 15.
(All n.itnral size.)

1. Macrof/eonujs cherriei (Allen). Santa Clara, Costa Rica.

(No. 664 iiii Museo Nacional de Costa Rica). Type.

2. netcrogeomys torridus sp. iiov. Cliichicaxtle, Vera Cruz. Mexico,

(No. 63629 9 ad. U. S. Nat. Mus.). Type.

3. Occiput of Macrogeomys dolichocephalus sp. nov. San Jose, Costa Rica,

(No. 36295 (? ad. U. S. Nat. Mus.). Type.

4. Occiput of Heierogeomys liispidus (LeCoute). .Tico, Vera Cruz, Mexico.

(No. 55343 <? ad. U. S. Nat. Mus.)

5. Occiputof Pappogcomys hidleri (Thomas). Sierra Nevada- de Col inia. .Jalisco, Mex-

ico. (No. 45618 9 yg. ad. U. S. Nat. Mus.)

6. Occiput of CVrt/or/eowu/s werrirtmi (Thomas). Lcrma, Mexico.

(No. 50110 c? ad. U. S. Nat. Mus.)

7. Occiput of I'latygeomys gymnuriis Merriam. Zapotlau, Jalisco, Mexico.

(No. 4.5611 S ad. U. Nat. Mus.)

8. Upper incisors of Macrogeomys dolichocephalus.

9. Upper incisors of Crafogeomys merriaml.

10. Ujiper incisors of Zygogeomys frichopus.

11. Upper incisors of Geomys hursarius.

12. Upper incisors of Geomys tuza.

North American Fauna, No. 8.

Plate 15

Mezsel, pho(c iith

I. GeOIMYS CHERRIEl 2.G.T0RR1DU5 3 & 8. G. D0LICH0CEPHALU5
4.G.HI5PIDU5 5.G.BULLER1 6 & 9. G. MERRIAM I 7. G. GYMNURUS
10. G. TRICH0PU5 II G. BUR5ARIU5 12, G. TUZA.

252 NORTH AMERICAN FAUNA, [no, 8.

PLATE 16.
1 and 2. Hetcrogeomys iorrUhis jiiv. Motzorongo, Mexico (No. 636-43 IT. S. National
Museum).
Molariforin teeth, showiiij;' deciduous premolars in situ; also unworn m 3, and
immature pattern of crowns in m 1 and 2.

1. Left upper series.

2. Left lower series.

Ix. Permanent upper jjremolar, uncovered to show unworn enamel

crown.
a, Permanent premolar not yet in place; h, deciduous premolar; c,
third u])per molar; d, third lower molar.
3, 4, and 9. Geomi/i^ hiirsdriiis ]nv. Elk River, Minn. (No. 4909 Merriam coll.)

Molariform teeth, showing- deciduous premolars in situ; also unworn m 3, and
immature pattern of crowns in m 1 and 2.

3. Left upper series.

4. Left lower series.

4x. Transverse section of m; about three-fourths down, showing that
the tooth is a single prism below, and that the enamel is confined
to its posterior border.

9. Left lower series from outer side, showing relations of permanent and

deciduous premolar, bilophiodont crown of mj, and forms of mi
and lUo (which show the manner in which the change occurs from
the double prism above to the single prism below).
a, Permanent premolar not yet in place; b, deciduous premolar; c,
third upper molar; d, third lower molar.
5, 6, and 7. Heierocjeomys torridus juv, (same specimen as in fig. 1).

Right upper premolar, showing unworn enamel cap and relations of enamel
and cement. The cement bands are shaded.

5. Outer side of tooth.

6. Inner side.

7. Posterior face.

a, Outer cement band of anterior prism; h, postero-external cement
band of posterior prism; c, inner cement band of posterior prism;
d, inner cement band of anterior prism ; e, lower end of enamel,
showing position of enamel organ.
8. Mncrof/eomi/s lieterodus ad.

Right upper premolar, s^^owini;' relation of cement bauds (unshaded) to enamel

(shaded) ir atur" tooth after the enamel cap [shown in figs. 5,

6, and 7] has w" . ,ft'.

10 and 11. Zygogeomijs trichopns ^vw . Nahuatzin, Mexico (No. 50104 U. S.Nat. Mus.).

Crowns of molariform series showing permanent enamel pattern and 'osteo-

dentine' islands.

10. Left ui)per series.

11. Left lower series.

12 and 13. Jhterogeoiinjs liisjjidus ad. Motzorongo, Mexico.

Right upper premolar, after the enamel cap of the young tooth has worn off,
showing permanent enamel pattern.

12. Outer side of the tooth (should be compared with 5, whichshows same
side of same tooth before the wearing down of the enamel cap begiiib).

13. Crown of same tooth.

a. Outer cement band of anterior prism.
h. Postero-external cement band of posterior prism.
Shaded bauds show the enamel.
14-17. Vruiogeomys cantanops i\w. Las Animas, Colorado.

14 and 15. A very young individual, but older than Nos. 1 and 4. The
deciduous premolars have been shed, but the enamel caps of the
permanent premolars (a) and the last true molars (m^c and mjd)
have not yet worn down far enough to show the enamel pattern of
the adult tooth (which may be seen in figs. 16 and 17). The crown
of the last lower molar (d) is still a double prism.
16 and 17. Another immature individual of the same species, but enough
older than 14 and 15 to show the permanent form and enamel pat-
tern of the adult teeth.
18 and 19. Geomys hiirsnrius ini. Elk River, Minnesota.

Crowns of molariform series showing permanent enamel pattern.

18. Left upper series.

19. Left lower series.

20 and 21. Macvogeomys cherriei im. Santa Clara, Costa Rica. Type.

Crowns of molariform series showing permanent enamel pattern.

20. Left upper series.

21. Left lower series.

North American Fauna, No. 8.

Plate 16

.. .r:-i^--ofa B Mersel. phalo Ink

1.2.5,6,7, IE & 13 Heterogeomys hispidus 3,4.9,18 & 19. Geomys bursarius

8. HeTEROGEOMYS HETER0DU5 lO&ll. ZYGOGEOMYS TRICHOPUS

I4,I5.I6&I7 Cratogeomys CASTANOPS ZOSc ZI.Keterogeomys cherriei

254

NORTH AMERICAN FAUNA.

[no.

PLATE 17.

(All natural size.)

Skulls seen from above ; vault of cranium cut away, showing door o

brain case.

1. Hetcrogeomtia torridus. Motzorongo, Vera Cruz, Mexico
2- Ziiyogeomijs Irichopus. Nahuatzin, Michoaeau, Mexico.

3. Geomys hitrsarius. Portland, North Dakota.

4. riati/geoviiis t/yinuunis. Zapotlan, .Jalisco, Mexico.

5. Cratof/eomys merriami. Amecauieca, Valley of Mexico.

Key to pi. 17.

tV Hva

Fia. 9. — Young skull of Cratogeomys merriami, vault of cranium cut to show floor of brain cage,

ae Anterior opening of alispbenoid can

as Alisphenoid bone.

bo Basioccipital.

bs Basispheuoid.

f Condyle of exoccipital.

cr Cribriform plate of ethmoid.

em External auditory meatus.

ex Exoccipital.

// Floccnlar fossa.

fr Frontal.

fro Descending or orbital plate of frontal (the
animal is so young that theplate.s of the
two sides have not yet united below).
■;' Jugal.

I Lachrymal.

ma Meatus auditorius internus,

mb Mastoid bulla.

n Nasal.

of Optic foramen.

OS Orbitosphenoid.

pet Petrous part of periotic.

pm,z Ascending arm of premaxilla.

ps Presphenoid.

ptf Spheno-pterygoid fossa.

sf Apex of sphenoidal fissure.

SO Snpraoccipital.

sq Squamosal.

^6 Superior face of tympanic or audital bull

zmx Zygomatic root of maxilla.

n

North American Fauna, No. 8.

Plate 17

BM^zseLpho'c 'M

'• HeTEROGEOMYS T0RR1DU5 Sp, nOV. 2.ZYG0GE0MY5 TRICH0PU5 Sp. QOV.

3. Geomys BUR5ARIU5 (Shaw) 4. Platygeomys gymnurus (Merriam)
5 Cratogeomys merriam I (Thomas)

b'56

NORTH AMERICAN FAUNA.

I NO. 8.

PLATE 18.
(All natural size.)

Vertical median longitiidiiial section of skull (mesethmoid and right
half of vomer in place).

1. Geomys hiirsariits $ . Kuoxville, Iowa.

2. Zijiiocicomy^ trichopus 9. Naluiatzlu, Mirhoacan, Mexico.

3. Heieroyeonn/.s torridus ^. yg. ad. Motzorongo, Vera Cruz, Mexico

4. Cndof/eomys merriami $ . Tlalpam, Valley of Mexico.

5. riatygeomys (jymnurus ^ . Zapotlau, Jalisco, Mexico,

Key to pi. 18.

Fig. 7. — Longitudiual vertical median section ot skull of Crator/eomys merriami, showing interior
of brain case and nasal chamber. Vomer and mesethmoid in place.

pa Parietal.

pet Petrous part of periotic capsule
2)1 Palatine.
pinx Premaxilla.
p.i Prcsphenoid.
Pterygoid.
Supraoccipital.
Squamosal.

Tympanic bulla, (antero-superior part,
■wliich alone appears within the brain
case.)
Vomer.

Vomerine sheath of maxilla.
First endoturbinal (Below and somewhat
behind it the anterior ends of the sec-
ond, tliird, and fourth cndotiu'bjnals
may be seen.)

1

Anterior palatine foramen.

2

Incisive foramen.

3

Meatus auditorius internus.

4

Floccular fossa.

5

Upper part of sphenoidal tissure.

as

Alisphenoid.

bo

Basioccipital.

bs

Basisphenoid.

c

Condyle of e.xoccipital.

fr

Frontal.

h

Haraular process of pterygoid.

ip

Interparietal.

me

Mesethmoid plate.

mt

Maxillo-turbinal.

vix

Maxilla.

n

Nasal.

nt

Naso-turbinal.

op

Lower border of os planum.

pt

It

North American Fauna, No.8.

Plate 18

^MuIUr.iel

B Mfi^eU pfutp luh

I-GEOMYS BUR5AR1U5 2. ZYGOGEOMYS TRICH0PU5 S.HETEROGEOMYS TORRIDUS
4. GRATOGEOMYS MERRIAMI 5. PLATYGEOMYS GYMNURUS

258

NORTH AMERICAN FAUNA.

PLATE 19.

(All natural size.)

1. Orthogeomys scalops 9 ad. Oaxaca, Mexico (skull from above).

2. Orthogeomijs scalops 9 a-d. Same specimen (l»ase of cranium).

3-7. Median longitudinal section of nasal chamber (vomer and mesetbmoid removed)
showing turbinated Ijones.

3. Geomys bm'sariiis ^ . Knoxville, Iowa.

4. Zygogeomi/s trichopus 9 • Nahuatzin, Michoacau, Mexico.

5. Heterogeomys forridiis J . Motzorongo, Mexico.

6. Cratogeomys mcrnami ^ . Tlalpaia, Valley of Mexico.

7. Platygeomys gymnitrus J. Zapotlan, Jalisco, Mexico.

Key topi. 19.

n\ It ;i^t

Fig. 10. — Longitudinal vertical median section of front part of skull of Oeomys bursarius.
moid and. vomer removed to show turbinated bones.

Meseth-

1 Anterior palatine-foramen.
3 Incisive foramen.

3 Vacuity in front of pre.spheDoid (present in Oeomys bursarius and tuza only. It is partly over-

lapped posteriorly:by the ascending wing of the vertical plate of the palatine, ap.).

4 Presphenoid fenestrum. Present in all species.

5 Upper part of sphenoidal fissure.
It First or superior endoturhiual.

2t Second endotiirbinal.

3t Third endoturbinal.

it Fourth endoturbinal.

ap Ascending wing of vertical plate of palatine.

fr Frontal.

mt Maxillo-turbinal.

mic Maxilla (the upper pointer rests on the maxillary .surface of the narial passage, the lower on the

sawed body of the bone).
n Nasal.
nt Naso-turbinal.
op Os planum.

pi Palatine (the upper pointer rests on the palatine face of the narial passage, the lower ou tlie
sawed horizontal body of the bone).
pmx Premaxilla.
ps Presphenoid.
vr Vomerine ridge of os planum (unites with the lateral wing of the vomer).

North American Fauna, No. 8.

Plate 19

FtTidUr, M. B Mfisel phot:, UiK

& 2 OrtHOGEOMYS SCAL0P5 ? 3. GE0MY5 BUR5ARIU5 4. ZYGOGEOMYS TRICH0PU5
5. HETEROGEOMYS T0RRIDU5 6. CRAT0GE0MY5 MERRIAMI
7. PLATYGEOMYS GYMNURUS

NoBTH Amebican Fauna, No

Maps 1 <fe 2.

MAP 1. -A DISTRIBUTION OF GENUS THOMOMYS.

B DISTRIBUTION OF GENUS OEOMTS (B = G. buraanua group ; B'= G.
tuza group.)
MAP 2.-DtSTRIBUTI0N OF GENUS CRAT0GE0MY8.

PLATE XII.

[All double natural size.]

Figs. 1- 3. Sorex {Atophyrax) hendirii palmeri. Oregon City, Oregon. Type.
(No. 56898, U. S. Nat. Mus.)
4- 5. Sorex {Microsorcx) hoyi. Elk River, Minn.

(No. 2520, Merriam collection.)
()- 7. Sorex calif ornicus. Walnut Creek, Contra Costa County, Calif.

(No. 44428, U.S. Nat. Mus.)
8- 9. Sorex ienelbis. Lone Pine, Owens Valley, California. Type.

(No. 32495, U. S. Nat. Mus.)
10-11. Sorex merriami. Fort Custer, Mont. Type.

(No. 4861, 2 , Merriam collection.)
12-13. Sorex macrodov . Orizaba, Vera Cruz, Mexico. Type.
(No. 58272, <?, U. S. Nat. Mus.)

124

Noilh American Fauna, No 10

Plate XII

1-3. Sorex bendirii 2'alvier/. (1,7. 8. californicus. 10,11. S.merriami.

4.5. S.hoyi. H.O. S.tenellus. 12,13. S.macrodon.

NORTH AM. FAUNA, No, II

Frontispiece

Bridled Weasel, Putorius frenatus.
Valley of Moxico

r

Black-footed Ferret, Putorius nigripes

Western Kansas.

U. S. DEPARTMENT OE AGRICULTURE

DIVISION OF ORNITHOLOGY AND MAMMALOIiY

I^ORTH AMERICAN FAUNA

N^o. 11

[Actual (late ofpublicatiou June 30, 1896]

%.

SYNOPSIS OF THE WEASELS OF NORTH AMERICA

C. HART MERRIAM

WASHINGTO>^^

GOVERNMENT PRINTING OFFICE
18 90

LETTER OF TRANSMITTAL.

U. S. Department of Agriculture,

Washington, D. C, ilfay 9, 1896.
Sir: I have the lioiior to traiisinit herewith for publication, as Ko.
11 of North American Fauna, a Synopsis of the Weasels of North
America.

Respectfully, C. Hart IMerriam,

Chief of Division of Ornithology and MammaUxjy.
Dr. Chas. W. Daeney, Jr.,

Acting Secretari) of Agriculture.

CONTENTS.

Page.

Iiitioductiou 5-7

Siibjifeints Putoriiis (the ferrets) 7-9

Subgenus Ictis (the weasels) 9

List of North American weasels 10

Descriptions of species - , 10-32

Table of cranial measurements '63

ILLUSTRATIONS.

(All natural size.)

PLATES.

Frontispiece. Heads of Black-footed Ferret and Bridled Weasel.

1. Skulls of Putoriiis nigripes and F. putorius.

2. Skulls of rutoriiis arciicus, alascensis, c'lvoytnmi, streatori, and rixosiis.

3. Skulls of Putorhts frcnaius, lonnicauda, and tropiculis.

4. Skulls of Puiorius noi'eboraveiisis, rvashingtoni, iwul jieninanhi.

5. Skulls of Putoriiis longicauda, cicognani, iiocchoracensifi, rixosiis, j^eninsuhr, and

arcticiis.

TEXT FIGURES.

1. Puiorius nigrijics, ,} old. Trego Count j^, Ivans.

2, 3. Putorius cicognani, ^ ad. Elk liiver, Minnesota.

4-6. Putorius vuvchoracensis, $ ad. Adirondacks, New York.
7-9. Putorius longicauda, ^ ad. Fort Sisseton, S. Dakota.
10, 11. Putorius longicauda sjyadix, J . Elk River, Minnesota.
12-14. Putorius arizoncnsis, $ ad. Boulder County, Colo.

15. Putorius freiiaf us, 9 ud. Cofre de Perote, Vera Cruz. Mexico.

16. Putorius tropicalis, $ ad. Jico, Vera Cruz, Mexico.

4

fl

No. IL NORTH AMEEIGAN FAUNA. June, 1896.

SYNOPSIS OF THE WEASELS OF NORTH AMERICA.

By C. Hart Merriaaf.

The preseut sj-nopsis includes the one ferret and all of tlie weasels
yet d iscovered in jSTorth America north of Panama. Of the true weasels
(subgenus Ictis ) no less thau 22 species and subspecies are here recog-
nized, 11 of which are described for the first time.

Until very recently the group has been in a state of chaos, but now,
thanks to Outram Bangs's excellent paper entitled ^A review of the
weasels of eastern Nortli America," the obscurity that has so long-
surrounded our eastern species has been cleared away and the task of
revising the whole group is rendered comiiaratively easy. Additional
material is needed from certain parts of the West, particularly from
southeastern Alaska and the middle and northern parts of the Great
Basin, and much remains to be learned respecting the extent to which
intergradation exists between allied forms having contiguous ranges.

Excepting the circumpolar type, represented in America by the weasel
of the barren grounds {ruforixs arcUcus nob.), and in Eurasia by the
closely related P. erminea, the weasels of North America fall naturally
into two groups, characterized by important cranial differences, and
having com])lementary geographic ranges. The first is a boreal group
comprising five forms: richordsoni, alascensis, cicognani, streatori, and
rixosKS, the southernmost of which {cicognani) reaches only the northern
United States. The other is an austral group comprising tlwfrcnatiia
and longicauda series and including P. peninsula', of Florida. Of this
series only a single species (P. arizonen.ns) reaches the lowermost of the
boreal zones, and this only in the mountains.

Between these two groups are two very interesting species, novehora-
censis and irojjicalis — the former inhabiting the eastern United States,
the latter the tropical belt of Mexico. Mr. Bangs has already shown
that the female of P. 7ioreboracensis resembles P. cicognani, while the
male resembles P. longicauda. The case of P. iropicalis is exactly
parallel, the female resembling cfcognani, while the male resembles
frenatus.

' Proc. P.iol. Soc. Washington, X, pp. 1-24, Feb. 25, 1896.

6 NORTH AMERICAN FAUNA. [Noll,

Among mammals the female is often less specialized than the male
and consequently bears more resemblance to the ancestral stock, thus
giving a clew to the line of descent when this can not be determined
from the iiuile alone. In the present instance the females of novehora-
censis and tropieaJis have small, smoothly rounded skulls without sagit
tal crests and with narrow audital bulhe and inflated squamosals, as
in the cicognani series, wiiile the males have large angular skulls with
well-developed sagittal crests, relatively broad audital bulhe, and iiat
squamosals, as in the loiif/icauda-frenatus series. The inference is that
the austral longivmuhi-frenatus series was derived from the boreal
cieognani stock, and that the differentiation took place in the South.
P. norehoraeensis occupies middle ground geographically, and may have
become differentiated from cicognani under existing conditions in the
area it now inhabits; but P. fropicalix, which inhabits tropical Mexico,
must either have originated from the cicognani stock when the latter
was driven southward by the cold of the Glacial epoch, or must have;
accomplished a very remarkable migration.

Turning now to the weasel of the tundras {F. arcticus), the female is;
also found to resemble the cicognani type, indicating — at least so far
as the American species go — that the whole group (subgenus Ictis) has
si^rung from an ancestral type related to 7*. cicognani.

Probably cicognani itself is a strongly specialized ty])e, although the
specialization took place a long time ago and seems to have been in
the direction of greater simplicity. The tendency has been toward a
narrowing of the skull as a whole and the obliteration of its promi-
nences and angles. The zygomata have been reduced and drawn in
close to the sides of the cranium, and the brain case has been nar-
rowed, elongated, and smoothly rounded off, as if to enable the head to
pass through small openings. The body as a whole has undergone
parallel modification, presenting the extreme degree of slenderness
known among the mammalia. This type of weasel seems to have been
developed for tlie express purpose of preying ui)on field mice or voles,
its narrow skull and cylindrical body enabling it to enter and follow
their runways and subterranean galleries. The extreme development
of the type is presented in P. rixosus and P. streatori^ whose exceed-
ingly small size and almost serpentine form make it possible for them
to traverse the burrows of even the smaller mice.

It is an interesting fact that the geographic range of the cicognani
group is almost coincident with that of the field mice of the subgenus
Microtuft. Farther south, where these mice occur sparingly or not at
all, the cicognani series of weasels is replaced by the larger and more
powerful longicauda-frcnatuH series. Where the ranges of the twO'
overlap, as on the northern plains, the large weasel (P. longicanda)
preys chiefiy on pocket gophers [Thomomys and Geomys) and ground
squirrels {Spcrniopliilus franlUni and S. 13-lincatufi), while the smaller
species {cicognani and rixosus) prey chiefly on mice.

June, 1896.] SYNOPSIS OF THE WEASELS OF NORTH AMERICA. 7

Similarly in the far Nortli, where the frozen tundras are inhabited by-
lemmings as well as voles, two weasels are present: the tiny, narrow-
skulled rixosus, which feeds mainly on mice, and the large, broad-skulled
arcticus, which feeds chiefly on lemmings and rabbits.

It seems clear, therefore, that the different types of weasels have been
developed by adaptation to particular kinds of food.

It is much to be regretted that sj)ecimens of the South American
weasels are not available for study in connection with the North Amer-
ican species. The only one I have seen is P. offinis Gray, which ranges
from Costa Rica to northern South America. While differing specif-
ically from frenat lis it clearly belongs to the same group.

Except in winter, weasels are usually so difficult to procure in any-
thing like satisfactory series that but few are available from most of
the localities represented in collections. As a rule, the number is too
small to afford reliable average measurements; hence the averages here
given are subject to correction.

The skull drawings in PI. I and those in the text (except figs. 10,
11, 15, and IG) were made by Benjamin Mortimer. Those in Pis. II to
V, inclusive, were drawn by Dr. James C. McOonnell under the super-
visi(m of the author. About half of the skulls shown in the latter
plates were used by Mr. Bangs in his paper already referred to.

Except where the contrary is distinctly stated, all the measurements
ill this paper were taken in the flesh by the collector. It is hardly
necessary to add that all measurements are in millimeters.

Genus PUTORIUS Cuvier, 1817.

Key to subgenera (for American forms only) :

Size large, about equaling tlie mink (Luh-eola); facial bar black; legs and feet
abruptly darker than upper parts subgenus Putorius.

Size medium or small, never more than lialf as large as the mink (Liitreola);
facial bar white or absent; legs and feet concolor with or paler than upper
parts subgenus Ictis.

Subgenus PUTORIUS Cuvier, 1817.

P«7on«s Cuvier: Rt-gne Animal, I, 147-149, 1817.
Cynonijionax Coues : Fur-Bearing Animals, 99, 147-148, 1877.

PUTORIUS NIGRIPES Aud. & Bach. Black-footed Ferret.

(PI. I, figs. 1, la, lb.)

1851. Putorius nUjrlpes Aud. & Bach. : Quadrupeds N. Am., Vol. II, pp. 297-299, pl.

93, 1851.
1877. Coues: Fur- Bearing Animals, 149-153, 1877.

Type locality. — Plains of the Platte River, in Nebraska.

GeograpMc range. — Great Plains, from western North Dakota and
northern Montana to Texas ; not known west of eastern base of Rocky
Mountains.

Characters. — Size of the mink; ears rather large; color buffy, with a

8

NORTH AMERICAN FAUNA.

[Xo.ll

(lark area in middle* of hack; fore and liind feet, end of tail, and baud
across fa'ce (inelnding- eyes) black.

Color. — Ground color pale yellowisli or buli'y above and below,
clouded on top of head (and sometiiiies on neck also) by dark-tipped
liairs; face crossed by a broad baiid of sooty black, which includes the
eyes; feet, lower part of legs, terminal third of tail, and preputial
region, sooty black; back, about midway between fore and hind legs,
marked by a large patch of dark umber-brown, which fades insensibl}'
into the buffy of surrounding parts; muzzle, lips, chin, a small spot
over each eye, a narrow band behind black facial bar, and sides of
head to and including ears, soiled wliite; anterioi- margin of ear near
base clouded with dusky.

Cranial characters. — Skull large and massive, very broad between
orbits, and deeply constricted behind postorbital processes,' whicb are
strongly develo])ed; zygomata strongly bowed outward; audital bulhe
obliquely flattened on outer side; a jn'ominent bead over lachrymal
opening.

Compared with our American weasels, the skull of l*ittorins nigripes

may be told at a glance
by its great size, the
basilar length in adult
males averaging about
05 mm., and in females
about 02 mm. Com])ared
with P. crersmanni of
s^ southern Siberia, it may
be distinguished by the
greater postmolar pro-
duction of the palate,
and by other minor cra-
nial characters. From
the common polecat of Europe {Tutor i no putorins) it differs in several
important characters, as may be seen by reference to PI. I. In P. imto-
rivs the postorbital region is very broad, the postmolar part of the
palate exceedingly long, and the anterior part of the audital bulLne, very
different.

Remarl\s. — The black-footed ferret bears no resemblance whatever to
any other American mammal, but is very closely related to the Sibe-
rian Pntorins crersmanni. It differs from the latter in having much
shorter and coarser fur, larger ears, and longer postmolar extension
of the palate.

In some specimens of Ptitorius oiifiripcs the ]>ale buffy of the under
parts is clouded across the breast between the fore legs, suggesting the
dark breast of P. eversmanni. The dark facial mask encircles the eyes

'This coustrictioD deepens -witli age, as in all the Aveasels. It is verj'^ deep iu tlie
slinll slio^vn in the accompanying text figure (fig. 1), which is that of an old indi-
vidual; much less deep in the younger specimen shown on PL I, fig. 1.

'Fm.l — I'liforiiis nigripps ^ ad. Trego Connty, Kans.

JUNE,1S00.] SYNOPSIS OF THE WEASELS OF NORTH: AMERICA. 9

(incliuling- the wliitisli suprnorbital spot) and dips slightly forMavd
before passing- transversely across tlie face, so that its ])osterior border
is in front of the plane of the outer angles of the eyes. Its anterior
border sometimes extends forward almost to the nasal pad, but this is
unusual. The black of the feet "reaches up and covers the fore ]eg to
the elbow, except along- the outer side, and the hind leg- to near the
knee, excei)t ])osteriorly.

,}[casKr('m(')ifs.^ — Average of 3 males: Total length, 570; tail A^erte-
br;c, 133; hind foot, (JO. Average of 2 females: Total length, 500; tail
vertebra^, 1-0; hind foot, 55.

Craiiidl nicasnrenioifs. — Average of 4 skulls of adult males: Basal
length, 04; basilar length of I iensel, 02.5; zygomatic breadth, 43; mas-
toid breadth, 37; breadth across postorbital processes, 22.5; iiiteror
bital breadth, 18; breadth of constriction, 12.5; palatal length, 33;
postpalatal length, 31.5. Average of 2 skulls of adult females:
Basal length, G0.5; l)asi]ar length of Heusel, 58.5; zygomatic breadth,
39; mastoid breadth, 34.5; breadth across postorbital processes, 20;
interorbital breadth, 1G.5; breadth of ccnstriction, 12; palatal length,
31; postpalatal length, 20.

Subgenus ICTIS Kaup, 1829.

Icth Kanp: Entwickeluiigs-Ciescliiclite mul Naturliches System der EuropJiiscben

TLierwelt, ])p. 40-41, 1829. (Contaius only a single species, Mitsiela nil.'/arin.)

ScLulze: Faunsp Saxonica>, Jlauimalia, p. 170, 1893.
ArctogaU Kaup : Entwickcluugs-CJescliicbte und Naturliches System der Euroi)ai-

sclien TLierwelt, p. 30, 1829. (Contains two species, cvmlnea and hoccamela.)
firt/e Wagner: Sujiplement Sclireber's Saugtliiere, II, p. 234, 1841. (Contains four

species, /iV'/u(fy(s, crmiiiea. hoccamela, and luthjaris.)

The names Icti.'; and Arefof/alc were proposed simultaneously in the
same publication. Each is accompanied by a diagnosis and included spe-
cies. The two names, therefore, according to Canon 18 of the A. O. U.
Code of i^omenclature, are e(|ually pertinent. In se(iuenc(; of pagiim-
tion Arctogalc conies 10 pages ahead of letis. let is contaius a single
species {rulg((ri.s = niralis Linn.), while ArctogaU' has two {erminea
and hoccaDicla). The reasons for choosing Trtis instead of .1 rctogaJe are :

(1) The type of Ictis is fixed beforehand, since it contained only a single
species, while in Arctogale the type must be established arbitrarily;

(2) Arctogale is now in current use for another genus of small carniv-
ora;2 to transfer it to a different group would lead to much confusion,
and would be a, great and seemingly unnecessary calamity. Hence,
since there is no rule to the contrary, the better course seems to be to
adopt Ictis and allow Arctogale to t\ill into synonymy.

'The number of specimens of which reliable flesh measurements are available is
too small to afford satisfactory averages.

'•'Jrcfojra/ta'eters, 1864, a genus of Viverrida-; (Jray, Proc. Zool. Soc. London, 18()4,
pp. 508, r,42-543 ; Blanford, Fauna P.ritish India, .Alanimalia, p. 114, 18S8; Flower and
Lydekker, Introduction to Study of Mammals, p. 533. 1891; Lydekker, Royal Nat.
Hist,,!, p. 401,1893-94.

10 NORTH AMERICAN FAUNA. [No.lt.

rurtliermore, Ictis has been already revived by Schulze (Faunae
Saxonicjie, Mammalia, 170, 1893), thou.yh used by bim iu a much more
comprebensive sense tban tbat originally intended.'

List of North American IVeaiiels with type localities.

No.

Name.

Type locality.

1

Northeastern North America (north of lat. 4P)

2
3
4

Fori Franklin, Great Bear Lake.

Skajiit Valley, Wasliington.
Osier, Saskatchewan.
Point Barrow, Alaska.
Kafliak Lsland, Ala.ska.

5
6
7
8
9

rixosvs

arcticus

areticus kadiaccnsls

Trout Lake, Mount Adams, "Washington.
Tarpon Springs, Florida.
Carlton House, Saskatchewan.
Fort Snelling, Minn.

10
11
12
13
14
15
16
17
18
19
20
21
22

peninsvlce

longicauda

longicauda spadix

Fla'^stalf, Arizona.

Black Hills, South Dakota.

Southern California.

Rogue Eiver Vallev, Oregon.

Valley of Mexico.
Pinabete, Chiapas, Mexico.

frenatus leucoparia

tropicalis

affinis

Patzcuaro, Michoacan, Mexico.
Jico, Vera Cruz, Mexico.
Colombia, South America.

PUTORIUS CICOGNANI Bonap. Bonaparte's Weasel.

(PI. II, figs. 3, 3ft, 4,4a.)

1829. Mustela (Putoritis) vulgaris Richardson: Fauna Boreali-Americana, Mammalia,
pp. 45-46, 1829.

1838. Mustela cirognanii Bonaparte: Iconografla Fauna Italica, I, fasc. XXII, p. 4,

1838; Charlesworth's Mag. Nat. Hist., II, p. 37, Jan., 1838.

1839. Fatorins cicoguanii Ricbanlson : Zoology Beecliey's Voyage, p. 10*, 1839.
1857. Baird : Mammals North America, pp. 161-163, 1857.

1891. Mearns: Bull. Am. Mus. Nat. Hist., N. Y., Ill, p. 235, May, 1891.

1896. Pidorius riehardsoni cicognani Bangs : Proc, Biol. Soc.AVash., X, pp. 18-21, Feb. 25,

1896.
1877. Putoriiis vulgaris Cones: Fur- Bearing Animals, pp. 102-109, 1877. Merriam:

Mammals Adirondacks, pp. 54-56, 1882 (habits) ; and most recent authors.

Type locality. — Nortbeastern Nortb America.

Geograpliic (UsirihuUon. — Boreal forest covered parts of North Amer-
ica from New England and Labrador to coast of sontheastern Alaska
(Jnneau, Wrangel, and Loring), and soutli in the Rocky Mountains to
Colorado (Silverton). It occurs in the interior of British Columbia (at
Sicamous), but in the Puget Sound region is replaced by a smaller and

1 Schnlze included in Ictis the two European weasels, vulgaris and erminea, and
also the mink, lutrcola, and polecat, imtoria.

'juNE,l89G.] SYNOPSIS OF THE WEASELS OF NORTH AMERICA.

11

darker form, P. fitreatori. In the United States it is common in New
England and New York, and in the forest-eovered parts of Minnesota.
It probably oocnrs also in northern Michigan and Wisconsin.

Gciirral characters. — Sixe small; tail slender and rather short; color
of under parts covering- toes and inner sides of botli fore and hind feet;:
color of npi)er ])arts never encroaching on belly, ])nt ending- along- a.
straight line.

Color. — Upper T)arts in ftummer pelage : uniform dark brown, hardly
darker on head; end of tail blackish; no dark spot behind corners of
mouth; under i)arts, usually including ui)per lip, white, more or less
tinged with yellow. In winter pelage: i^ure white with a strong yellow-
ish tinge on rump, tail, and under parts; end of tail black.

Cranial characters. — Skull sraall.^ light, narrow, and elongated witlr-
out marked postorbital processes, and only a slight postorbital constric-
tion; zygomata narrow, and jiot bowed outward; brain case elongate
and subcylindric; audital bulliB small, narrow, and siibcylindric, almost
continuous anteriorly (except in old age)
with the greatly inflated sipuimosals;
palate narrow; the tooth rows more
nearly parallel than in the other spe-
cies; skull of female similar to that
of male, but smaller. Contrasted with
riehanhoniy the skull of cicognani is
smaller, the audital bullae decidedly
smaller, and the dentition lighter. In
nearly every series of eii-ognani there are
one or two old males whose skulls are
abnormally large and closely resemble
skulls of riehardsoni, except that the
audital bulhe are always smaller.

Measurements. — Average of 5 males from Ossipee, N. H. : Total
length, 278; tail vertebne, SO; hind foot, 30.5. Average of 3 females:
Total length, 230; tail vertebra?, 69; hind foot, 30.5.

PUTORIITS CICOGNANI RICHARDSONI (Boiiap.). RichcariLsoii's Weasel.

1829. Mnstela (_Putoriiis) erminea Ricliardson: Fauna Boreali-Amcricana, p]). 46-47,
1829. (lu part: specimen from Fort Franklin, Great Bear Lake. Not M.
erminea Linn.)

1838. Musteta richarilsoiii Bonap. : Charles worth's Mag. Nat. Hist., Vol. XI, p. 38,

1838. (l)aseil on Richardson's specimen from Great Bear T^ake).

1839. Piiioriiis richard.'ioni U'igIi. : Zool. Beechey's Voyage of Blossom, Mannniilia, 10*,

1839.
1896. Bangs: Proc. Biol. Soc. Washn., X, pp. 1-24, Feb. 2;-), 1896. (In part.)

Type locality. — Fort Franklin, Great Bear Lake.

Geographic (listrihut'ton. — Iludsoniau timber belt from Hudson Bay
to interior of Alaska and British Columbia.

General characters. — Similar to P. cicognani but larger; tail of
medium length, its terminal third black.

Figs.

i aud 3. — P. cicognani cf ad. Elk
Eiver, Minnesota.

12 NORTH AMERICAN FAUNA. [No.n.

Color. — TTppor parts dull clioeolate brown, tliis color reacliiiig down
on both fore and liind feet to base of toes; underparts whitish, more
or less suffused witli yellowish, the pale color extending out in a very
narrow aud sometimes interrujjted strip along inner side of hind feet
to toes; tail concolor all around except at tip, which is black for about
one-third the total length of tail. In n-intcr pelage: white all over
except termiual third of tail, which is black; ruraj) and belly more or
less tinged with yellowish.

Cranial characters. — Skull long, narrow, and .subcylindric like that of
cieognani, from which it differs chiefly in larger size, larger audita!
bulhe, and heavier dentition.

Remarls. — P. richardsoni, as pointed out by Mr.Bangs, is simply a more
northern form of eieoguani, with which it intergrades completely. It
inhabits the Hudsonian timber zone while cieognani inhabits the Cana-
dian. On the north, where the timber ends and the tundra begins, the
range of richardsoni meets that of arcficns. The two species differ
widely in both cranial and external characters. The light subcylindric
skulls of richardsoni J with the narrow frontals and appressed zygomata,
require no comparison with the broad massive skulls of arcticus with
their broadly flattened frontals and widely spreading zygomata. The
external differences are almost as marked. In richardsoni the under-
parts are nearly white or, at most, only tinged with pale yellowish; tlie
color of the upjier parts covers both fore and hind feet, reaching the
base of the toes; the tail is relatively long, coucolor except at the tip,
which is black for about one third its length. In arcticus the under
parts are deep yellow; the color of the upi)er parts stops short of tbe
fore feet and reaches only halfway down the hind feet; the tail is short,
yellow below on its basal half, and has a long, black pencil covering at
least half its entire length.^

Measurements. — (From dry skin of male from Fort Simpson): Total
length, 390; tail vertebra', 05; hind foot, 43 (probably 45).

PUTOKHTS KICHARDSONI ALASCENSIS siibsp. nov. .Tiuipaii "Weasel. .
(PI. II, figs. 2. 2a.)

Type from Jmiean, Alaska. Xo. 74423, ^ ad., IT. S. National Museum, Dept. Agric.
coll. Collected August 22, 1895, by Clark P. Streator. Original number 4806.

General characters. — Similar in size and general appearance to P.
richardsoni, but white tips of fore and hind feet more extensive and
interorbital region very much broader.

Color. — Upper parts dull chocolate brown, this color reaching down
on fore legs to wrists and on hind legs to middle of iipper side of feet;

1 It is not strange that Mr. Bangs failed to discriminate between arellcns and
rii;harchoui. Tiie available material is scanty and mostly of poor quality, aud most
of the skins had the skulls inside. Through the kindness of INIr. F. W. True, cura-
tor of mauHuals in the United States National Museum, tbe skulls have been removed
and placed at my disposal.

JUNE, 1896.] SYNOPSIS OF THE WEASELS OF NORTH AMERICA. 13

tei'iniual third of tail black; uuder i)arts, iuchiding^ upper li]), fore feet,
and distal half of hind feet, soiled white, tinged with yellowish. Winter
pelage probably white.

Cranial characters. — Skull similar to that of P. richardsoni, but very
much broader between orbits and across muzzle; postorbital processes
more strongly developed; constriction deeper.

EemarlxS. — Mr. Streator obtained two males of this new weasel at
Juneau in the latter part of August. He obtained also, at the same place
and time, three females, which in color aud markings agree with the
males, but are hardly half as large. Their skulls are as small as those
of true cicognani, which they closely resemble. If they are the females
of ulascensis, as seems probable, then this weasel exhibits as great
sexual difference in size as P. noveuoracensls, in which respect it stands
unique as a member of the cicognani group. The only alternate possi-
bility is that cicognani and alascensis occur together at Juneau, and that
of the 5 specimens collected there by Streator the 2 males are alascensis
and the 3 females cicognani.

Measurements. — Average of two males from Juneau, Alaska: Total
length, .335; tail vertebrse, 95; hind foot, 48, Average of three females
from same place: Total length, 270; tail vertebra;, 77; hind foot, 34.

PUTORIUS STREATORI sp. uov. Puget Sound Weasel.

(PI. II, figs. 5, 5(t, 6, 6a.)

Type from Mouut Vernou, Skagit Valley, Washington. No. 76646, ^^ a<l., U. S. Nat.
Mils., Dept. Agric. coll. Coll. Feb. 29, 1896, by D. R. Luckey. (Original number 3 )

Geographic (listribution. — Puget Souud and coast region of Washing-
ton and Oregon; south at least to Yaquina Bay (Newport), Oregon.
Confined to a narrow strip along the coast.

General characters. — Similar to Putorius cicognani., but smaller and
darker, with color of upper i)arts encroaching on belly.

Color. — Upper i^arts, including upper lip and fore and hind feet,
uniform dark chocolate brown, darkest on head, and encroaching far
on belly aud throat (often meeting along middle of belly); terminal
third of tail black; under parts narrowly and irregularly white, faintly
tinged with yellowish. In winter pelage at low altitudes the color of
the upper parts is paler (almost drab brown) and the toes may become
white; at higher altitudes the whole animal changes to white, ^ except
the end of the tail, which always remains black.

Cranial characters. — Sku,ll of male similar to that of male cicognani,
but smaller, slightly broader in terorbi tally, and with somewhat more

'Mr. R. E. Darrell, of Port Moody, British Columbia, writes me : " I have discovered
that, although the weasels do not change color down near salt water, they do chang'? to
the white winter coat in the mountains." Specimens in the Department collection
from Mount Adams, Washington, killed in February and March, are in the white
winter pelage. The type and a female from the same locality (Mount VernoUj,
Skagit Valley) are in the drab-brown wuiter pelage.

14 NORTH AMERICAN FAUNA. [No.ll.

prominent postorbital processes and smaller aiidital bulla', Sknll of
female very mucli smaller and more delicate tliau that of male,
resembling" female o£ cicognani, but snmller.

Rniwrks. — Futorins streatori is a dark Pacific; Coast form of cicognani^
with which it may be found to intergrade. It differs conspicuously
from cicogncmi in the color of the under parts, the dark chocolate brown
of the back and sides encroaching far on the throat and usually meet-
ing along the median line of the belly, thus reducing the white to a
narrow and irregular strip, which expands on the anterior part of the
throat, on the breast behind the fore legs, and immediately in front of
the hind legs, and stops abruptly on the under surface of the thighs.

Five winter specimens from Sumas, British Columbia, kindly loaned
by ]\[r. Outram Bangs, point toward intergradation with eicognani. In
three out of the five, the toes of both fore and hind feet are white, and
the color of the ujiper jjarts is much paler than in summer pelage.
Two of these specimens have the bellies broadly white, as in eicognani.
They are also much larger than streatori. Specimens from Sicamous,
in the interior of British Columbia, are fairly typical eicognani^ having
the nnder parts broadly white; the upper lip, a strip along the inner
border of the hind feet, and the toes of both fore and hind feet, while.
Specimens from southeastern Alaska (Juneau, Wrangel, and Loriug)
must also be referred to eicognani, and not streatori.

Measurements. — Unfortunately, no flesh measurements are available
from the type locality. Specimens from Trout Lake, near Mount Adams,
Washington, are slightly smaller than the Mount Vernon specimens,
and measure as follows: Average of two adult males: Total length,
270; tail vertebne, 83; hind foot, 33. An adult female: Total length,
210; tail vertebra', 51; hind foot, 24.

PUTORIUS RIXOSUS Bangs. Bang's Weasel.

(PI. II, tigs. 7, 7a.)

1857. rutorius pusillus Baird: Mammals N. Am., pp. 159-161, 1857. (In part: speci-
men from Pemhina.)
1896. rutorius rixosus Bangs: Proc. Biol. Soc. Wash., Vol. X, pp. 21-22, Feb., 1896.

Tyjw locality. — Osier, Saskatchewan, Canada.

Geographic distribution, — Boreal America from Hudson Bay to coast
of Alaska (St. Michaels); south to northern Minnesota (Pembina) and
Montana (Sun River).

General characters. — Smallest weasel known; tail short and without
black til) ; only American weasel lacking the black tip.

Color. — Summer pelage: Upper parts dark reddish brown; tip of tail
not darker ; under parts white. In ivinter pelage: Pure white all over,
including end of tail.

Cranial charaeters. — Skull (of type specimen, 9 ad.. No. 642 Bangs'
Coll.') very much smaller than the smallest female of any other known

• I am inclebted to Mr. Bangs for the privilege of examining this specimen. Unfor-
tunately, the hasioccipital is broken otf ; hence the basilar length is estimated.

jDNE.isne.] SYNOPSIS OF THE WEASELS OF NORTH AMERICA. 15

species (total length from occiput to front of preninxillic, 28.05 ^asal
length, liG.ii; zygomatic breadth, 14; length of palate, 11; interorbital
breadth, 5.5; breadth across postorbital processes, 7.5; length of aiidi-
tal bnlla^, 9.5). The skull is a miniature of P. cicognani except that tlie
postorbital processes are more i)romiuent, the braiu case more com-
pressed, and there is a distinct sagittal ridge.

MeaHuremenis. — Tjq^e specimen, female, measured in flesh: Total
length, 150; tail vertebra^, 31; hind foot in dry skins, 20-22.

PUTORIUS ARCTICUS sp. nov. Tuuclia Weasel.
^ (PI. II, tigs. 1, 1«; PI. V, figs. 6, 6«.)

Tijpc from Point Barrow, Alaska. No ^liiTn <? ^^- U- S. Nat. Mus. Collected July
16, 1883, by John Murdoch. Original number, 1672.

GeodrapMc distrihution. — Arctic coast and tundras. Specimens ex-
amined from Anderson Kiver, Franklin Bay, old Fort Good Hope, lower
Mackenzie River, Point Barrow, and St. Michaels.

General characters. — Size large; ears small; tail short but with very
long black pencil; underi)arts yellow (including underside of basal half
of tail).

Color — (Type specimen, male adult.) Upper parts, including upper
lip, dark yellowish brown ; chin white ; under parts deej> ochraceous yel-
low, broadly including inner and posterior sides of fore legs, whole of
fore feet, distal half and inner side of hind feet, and under side of tail
to or nearly to black tip; black tip very long, covering at least half of
tail (including long terminal hairs); color of upper parts not encroach-
ing on belly. In icinter 2iela(je, white all over except long black tip of
tail; the white tinged with yellow posteriorly.

Cranial characters. — Skull rather large, broad, and massive; frontal
very broad interorbitally; muzzle broad and blunt; postorbital proc-
esses moderately develoj^ed; postorbital constriction marked; zygo-
mata strongly bowed outward; brain case subtriangular and rather
short; audita! bulhe subcylindric; postglenoid space smaller than in
richardsoni and hardly iuHated except in female. Contrasted with
P. richardsoni, the skull of P. arcticus is somewhat larger, ]nuch broader,
and more massive; braiu case subtriangular instead of subcylindric;
zygomata bowed far outward instead of appressed; postorbital pro-
cesses more prominent; i)ostorbital constriction much deeper; frontal
much broader interorbitally; palate broader posteriorly; dentition
heavier. Adult male skulls of P. arcticus resemble certain old males
of washingtoni, but differ in much greater breadth of frontal between
orbits, broader muzzle, and blunter jiostorbital processes. P. arcticus
resembles true erminea of Sweden much more closely than it does any
American species.

Rcmarhs. — Putorius arcticus, which has been heretofore confounded
with erminea or richardsoni, is one of the most strongly characterized
species of the genus. It is a large animal with deep ochraceous yellow

16 NORTH AMERICAN FAUNA. [N'o.ll.

under i)arts aud a rather short tail which ends in a remarkably long
black pencil. The skull differs from all other American weasels in the
great breadth of the frontal region and the breadth aud bluntuess of
the muzzle, in both of which respects it resembles true erminca. The
only American species wliose skull approaches it at all is P. washhuf-
toni, as mentioned above. In exterjuil characters the differences are
too great to require comparison.

It is interesting to find in this country an Arctic circumpolar weasel
which, though S[)ecifically distinct, is strictly the American representa-
tive of the Old World erminea. The ])attern of coloration, as described
above (under color), is precisely as in enninea, but the tints differ
materially. The upper pai'ts in erminca lack the golden brown of
((rcticHs, and the under parts are very much paler aud of a different
tint, being 'i)ale sulphur yellow instead of ochraceous. Moreover,
arcilcuH lacks the whitish border to the ear which is present in erminea.
In winter pelage the two seem to be indistinguishable except by cranial
characters.

A small form of arcticns oc(;urs ou Kadiak Island, xVlaska. It has
smaller aud narrower audital bulhc, less spreading zygomata, less
divergent tooth rows, and decidedly shorter postmolar production of
l)alate. It is probably wortLy of recognition as subspecies l((diacensis.
An adult male (No. G52!)0) collected April 25, 1804, by B. J. Bretherton,
measured in the flesh: Total length, 318; tail vertebr;i', 80; hind foot,
41. It is in the white winter i)elage, just beginning to change, aud the
terminal half of the tail is black.

MeaHnrcmentti. — From dry skin of tyi)e, male adult. Point Barrow,
Alaska: Total leugth, 380; tail vertebrte, 75; pencil, 55; hind foot, 48
(at least 50 in the flesh).

PUTORIUS NOVEBORACENSIS De Kay. New York Weasel.

(PI. IV, figs. 1, la, 2, 2a; PI. V, tigs. 3,3a.

1810. J'lttorius iioveboracensisDe Kay: Catal. Mammalia New York, p. 18, 1810 {nomen

nudum); Zoology of New York, Mammalia, p. 36, 1842.
1840. Emmons: Rept. Quadrupeds Massachusetts, p. 45, 1840.
1857. Baird : Mammals N. Am., pp. 166-169, 1857.
1896. Bangs: Proc. Biol. Soc. Wa8li.,X, pp. 13-16, Feb. 25, 1896.
1877. Futorlas ( Gale) erminea Cones : Fur-Bearing Animals, pp. 109-136 ( in part), 1877.
Futo7-iHs erminca Thompson, Aud. & Bach, (part), Allen, Merriam, aud most recent
authors.

Type locality. — New York State.

Geographic distribution. — Eastern United States from southern Maine
to North Carolina, and west to Illinois,

General characters. — Male large; female small; tail long aud busby,
much longer than in cicoynani, but shorter than in lonyicaiula; the
black terminal part longer than in any other species except articus,
covering oue-third to one-half the tail and measuriug 50 to 75 mm.
Animal turns white in winter in northern part of range. Extraordinary
sexual difference in size aud crauial characters.

JtTNE,189G.] SYNOPSIS OF THE WEASELS OF NORTH AMERICA.

17

Color. — Smnmer peUuiv: Uiiper parts, including fore and liiud feet
and anal region, and often encroaching irregularly on belly, ricli dark
chocolate brown, sometimes suggesting sealbrown ; under]>arts (usually
including ujtper lip) white, more or less washed with yellowish; no
yellow on under side of tail or on hind feet, the color of under parts
stopping short of ankle. Wintir pelage: In southern jjart of range
similar to summer pelage, but upper parts paler, nearly drab brown.
Northern specimens white all over except terminal third of tail, which

is jet black; throat, belly, posterior
half of back and tail always suilused
Avith yellowish.

Cranial characters. — Skull of male
large, heavy, and elongate; sagittal
ridge present in adults; postorbital
l)rocesses and constriction mod-
erately developed; zygomata not
hoired oittirard; audital bulla' rather
narrowly oval, usually rounded an-
teriorly as well as posteriorly. Skull of female very small, light, and
narrow, Avith brain case elongate and subcylindric, much as in cicognani;
audital bulhe sjuall, narrow, and not rising abruptly anteriorly from
intlated squamosals, which latter are elongated and strongly inflated as
in cfcognani. Skulls of males may be distinguished from those of male
longicauda by shorter postorbital processes, less nmrked postorbital
constriction, less triangular brain case, lower sagittal ridge, very much
narrower zygomata, which are not bowed outicard, narrower palate, and
narrower audital bulhe, which are more rounded anteriorly. The resem-
blance to F. icashingtoni is very much closer, but male skulls oi noreho-

FlG. 4. — Putorius nuvcb'iracoisi'; cT ;id,
daulis, New York.

Adiron-

5 and li. — I'uturins tiiireboraceimis. Adirondacks, Kew York.

racensiH may be distinguished by larger size and much larger audital
bnllffi. The female skull, owing to the inflation of its squamosals
inferiorly, needs no comparison with either washingtoni or longicauda,
but is with difllculty separated from cicognanl in regions where the two
species overlap. The postorbital processes are longer and the car-
nassial and sectorial teeth larger in the females of norehoraceHsis than
in cicognanl from the same localities.

Bemarls. — Futorins uorehoraccnsis may usually be distinguished from
P. cicognanl l)y larger size and also by the longer and more bushy tail,
16932— No. 11 2

18 NORTH AMERICAN FAUNA. [No.U.

aiul greater length of the l)lack terminal part. Females of nonhora-
ccn.si.s, however, sometimes resemble males of cicognanl rather closely.
They may be distinguished not only by the greater length of the tail
but also, if in summer pelage, by the absence of yellow from the under
side of the tail and inner sides of the hind feet, which parts in cicognani
usually show more or less yellow.

Measurements. — Average of 10 males: Total length, 407; tail ver-
tebra>, 140; hind foot, 47. Average of 10 females: Total length, 324;
tail vertebra', 108; hind foot, 34.5.

PUTOKIUS WASHINGTON! sp. iiov. Wushiugton Wetisel.

(PI. IV, figs. 3, 3a, 4,4a.)

Tyi)e from Trout Lake, base of Mouut Adams, State of Wasliingtou. No. 7(5022, J^
ad., U. S. Nat. Miis., Dept. Agriculture collectiou. Collected December 15, 1895, by
D. N. Kaegi.

General characters. — Similar to P. novehoracensis in size and general
appearance, but with longer tail and shorter black tix). Female very
much smaller than male, as in novehoracensis.

Color. — Color in summer pelage unknown (probably dark chocolate
brown). There are two winter pelages, probably dependent on alti-
tude. In drah winter pelage: Upper parts uniform drab brown; end
of tail black; under parts white, more or less suffused with pale yel-
lowish. The color of the upper x)arts encroaches on the sides of the
belly as in novehoracensis, and a brown spot is present behind the cor-
ners of the mouth, which may or may not be confluent with the brown
of the cheeks. In the type and two other specimens the hind legs and
feet are the same color as the upper parts except that the toes are
tipped with whitish and the tips of the fore feet are white. In another
specimen, collected January 22, the white is more extensive, covering
all of the fore feet and about half of the hind feet. In summer pelage
the legs and feet are doubtless the same color as the upper parts, the
white of the belly stopping high up on the thighs. In white wintev
l^elage: White all over except black tip of tail; tail, rump, and belly
strongly suffused with yellow. In one specimen (No. 76004, male,
February 7, 1890) the yellow reaches forward over the back nearly to
the shoulders; in another (No, 76588, male, February 4, 1896) the whole
back is white.

Cranial characters. — The skulls of the two sexes differ greatly: that
of'the male resembles novehoracensis closely iu size and general char-
acters, but differs in having the audital bulla- much shorter and the
X)ostorbital processes less strongly developed. Thoi^ostorbital constric-
tion is equally marked. The skull of the female is very much smaller
than that of the male, averaging about 38 mm. in length, while the
male averages 45 mm. Contrasted with the female of novehoracensis
the brain case is broader»posteriorly and less cylindric. The audital
bulhe are more sharply separated from the squamosal inflation and the
latter is only slightly marked, not reaching tJie plane of the bullie. The

JrNE,l896.j SYNOPSIS OF THE WEASELS OF NORTH AMERICA. 19

resembliiiice therefore to P. cicognani is much less marked in the female
ica.shitujtoiii than in the female novehoracensin.

Eemarls. — This new species is represented in the collection by 14:
skulls and 6 skins, of which the greater number are males. The female
is darker than the males, and the top of the head is darker anteriorly
than the rest of the upper parts, while in the males it is concolor with
the back. These differences are probably seasonal, the female not
having comi)leted the change from summer to winter pelage, though
collected December 11. All are from the Mount Adams region.

MeasnremenU. — The skins, which are well made, afford the following
approximate measurements: Male, total length, 240; tail vertebra', 155;
hind foot, 44. Female, total length, 300 ; tail vertebra?, 120 ; hind foot, 37.

PUTORIUS PENINSULA Rhoads. Florida Weasel.

(PI . IV, figs. 5, 5a; PI. V, tig. 5.)

rutorius ptninsuhv Rhoads: Proc. Acad. Nat. Sci. Phila., June 1894, 152-155.
Bangs: Proc. Biol. .Soc. Wash., X, pp. 10-13, Feb. 25, 1896.

Type /oca^i^j/.—' Hudsons,' 14 miles north of Tarpon Sjmngs, Fla.

Geographic distribution. — Peninsula of Florida; limits of range
unknown.

General characters. — Size rather large, about equaling male of Puto-
rius noveboracensis ; skull similar to that of longicanda^ but with very
large audital bulla*.

Color. — Upper parts dnll chocolate brown, darkest on head; upper
li]) and chin whitish; rest of under i)arts, including fore feet and toes
of hind feet, yellowish; a brown spot behind corners of mouth ; a small
tuft of white hairs under anterior root of ear. The color of the under
parts covers the belly broadly and is not encroached ui^on by the color
of the upper jyarts. Irregular and inconstant white markings are some-
times present between and behind the eyes.

Cranial characters. — Skull rather massive, resembling that of longi-
canda, hwtvfith higher sagittal crest; less spreading zygomata; narrower,
higher, and more swollen audital bulhe, and less prominent postorbital
l)rocesses. Contrasted with P. norehoracensis the postorbital constric-
tion is deeper, the l)rain case higher and moresubtriangular, the audital
bnlla; higher and more swollen, the upper carnassial tooth decidedly
larger, and the molar smaller. The upi)er molar is peculiar : It is short,
hardly expanded at either end, and implanted at right angles to the
premolar series.

Measurements. — An adult female from Tarpon Springs, Fla. : Total
length, 374; tail vertebrje, 127; hind foot, 44.5.

PUTORIUS LONGICAUDA Bonaparte. Long-tailed Weasel.
(PI. Ill, figs. 3, 3a, 4, 4a; PI. V, figs. 1, la.)

1829. Musfehi (FiitoriHS) erminea Kichardson: Fauna Boreali-Americaua, pp. 46-47,

1829 (in i)ait: Spocinien from Carlton House).
1838. Miislehi lonyicauda Bonaparte: Charlesworth's Magazine Nat. Hist. N. S.,

II, p. 37-38, 1838 (based on Richardson's long-tailed variety of erminea from

Carlton House).

20

NORTH AMERICAN FAUNA.

[Xo. n.

Fl(i.7.

-Puturius longiraudu.
S. Dak.

Fort Sissetou,

1839. Putorius loiigicauda Rich. : Zool. Beecbey's Voyage of Blossom, p. 10," 18"0.

1857. Baird: Mammals N. Am., pp. 169-171, 1857.

1877. Cones: Fur- Bearing Animals, pp. 136-142, 1877.

1896. Bangs: Proc. Biol. Soc. Wash., X, pp. 7-8, Feb. 25, 1896.

Type locality. — Carlton Hoase, on Noitli Saskatcliewan River,
Canada.

Geognqjhic (lutribnUon. — Great Plains from Kansas nortli\vai<l.
General cliaracters. — Size large (adult males averaging al)out 4.">() mm.

in total length) ; tail veiy long (ver-
tebra* 155 mm. or more in males),
its bla(-k ti]) ratlier short; under
l)arts always strongly yellowish or
ochraceous.

Color. — Upper parts pale yel-
lowish brown, or pale raw-umber
brown, becoming darker on head;
terminal part of tail black; chin
and upper lip all the way round
white; rest of under parts varying
from strong buff'y yellow to ochraceous orange, the color extending from
throat jjosteriorly, including upper side of fore feet, inner side of hind
feet, and upiier side of hind toes; under side of tail more or less suffused
with yellowish; soles of hind feet brownish. In worn summer pelage
the color of upper parts is decidedly paler, and in some old specimens
the upper and lower surfaces are not sharply differentiated. The
orange tinge of the under
parts is strongest on the
throat.

Cran ia I c h a r a cter ,s . —
Skull large, broad, and
massive, with well-devel-
oped postorbital proc-
esses, strongly marked
postorbital constriction,
and a moderate sagittal
crest; zygomata bowed
strongly outward; brain
case subtriangular as seen
from above ; audital bulla;
rather broad and subrect-
angular; palate broad;
dentition heavy; audital bulla- anteriorly rising abrui)tly from s((ua-
mosal, which is not inflated in either sex; skull of female similar to
male, but smaller, and with only a slight sagittal ridge. Contrasted
with male skulls of novehoracensh and tcasJiingfoni, the male of lotif/i-
cauda is broader and relatively shorter, with more spreading zygomatic
arches, longer postorbital processes, deeper postorbital constriction,

Figs. 8 and 9.—/'. Inmiicaiifla rf ad. Fort Sisseton, S. Dak.

JiNE, 1896 1 SYNOPSIS OF THE WEASELS OF NORTH AMERICA.

21

1111(1 imicli broader and more rectangular audita! bulliv^, wbicli as a rule
are broadly truncate iustead of narrowly rounded anteriorly.

Mt'dsnrements. — Average of 4 males from jjlaius of Saskatcbewan aud
Alberta : Total lengtb, 450 ; tail vertebne, 1 65 : bind foot, 51. Average
of 3 females: Total lengtb, o87; tail vertebr*, 144: bind foot, 44.

Pl'TORIUS LONGICAFDA SPADIX Bangs.
J'ulurius loiniiciuida ■'<pudix Ban.irs: Proc. Piol. Soc. Wash., X, pp. 8-9, Feb. 25, 1896.

Ty]}e locaJitij. — Fort Snelling, near Minneapolis, Minn.

Geographic di.stribntio)i. — Edge of timber belt in Minnesota, along
boundary between Transition and Boreal zones.

General cliaracters. — Similar to P. lonr/icauda, but mucb darker.

Color. — Swrnner pelage : Upper parts chocolate browu, darkest on tbe
bead, but paler tlian in nove-
horacevsis ; cbin and upi)er lip
whitish all round; rest of under
parts, including upper surfaces
of fore feet and toes of hind
feet, butfy yellow ; terminal part
of tail black. Winter pelage:
Snow-white everywhere except
bbick tip of tail and a yellow-
ish suffusion on rest of tail, and
sometimes also on under side of
bind i'i^et.

Cranial charaeters. — As in P.
longieauda.

Measurements.^ — Average of G
males from Fort Snelling, iMinn. : Total length, 460; tail vertebrae, 166.5;
liiiid foot, 54.5. Average of 3 females: Total lengtb, 356; tail verte
bra-, 132; hind foot, 43.5.

PUTORIUS SATURATUS .sp. uov. Cascade Mountain Weasel.

Tjipe from Siskiyou, near soutlieru boundary of Oregon (altitude, about 4,000 feet).
No. 65930, cT ad., U. S. Nat. Mas., Department of Agricultnrc colli-ctiou. CoUei'ted
.June 6, 1894, by Clark P. Streator. Orig. No. 3905.

General charaeters. — Similar to P. arizonensis, but larger and darker,
with belly more ocliraceous, and with distinct spots behind the (!orners
of the mouth.

Color. — Color of upiier jiarts in summer pelage (June) dark raw
umber brown, becoming much darker on the top of the head and nose;
terminal part of tail black; a brown spot at corner of mouth which
may be confluent with brown of cheeks ; color of upper parts extending
over outer side of forearm to wrist, and over hind foot to toes; chin

Fjgp. lu aiul 11.

■ Putorius I. spaiiix ? ad. Elk Kiver,
Miiiuesota.

'These measnrenieuts were taken iu the liesh by Dr. E. A. Mearns, to whom I am
indebted for them.

22 NORTH AMERICAN FAUNA. rXo.ll.

white; rest of uiuler parts ocliraceons or orange yellow, including the
fore feet, and reaching narrowly do\A'n the under side of hind leg to
ankle, whence it may or may not extend in a nariow line along inner
side of foot to toes; under side of tail more or less suffused with goldeu
chestnut; anal region chestnut brown; in «'orn pelage the colors are
everywhere much paler.

Cranial characters. — Skull similar to tliat of P. arizonoisi.s but with
postorbital processes broader at base and less peg like.

RemarTcs. — This handsome weasel rei)laees lowjicauda on the Cascade
and Siskiyou mountains of Oregon and Washington, reaching a short
distance into British Columbia. The only specimens examined have
come from Siskiyou, Oregon, and Chilliwack, British Columbia (the
latter, No. 3553, collection of E. A. and O. Bangs).

MeasnremenU. — Average of 2 males from Siskiyou Mountains, Ore-
gon: Total length, 423; tail vertebrae, 164; hind foot, 48.

PUTORIUS ARIZONENSIS Mearns. Mountain Weasel.

rutoiius arisonensis Mearns: Bull. American Miiseniu Nat. Hist., Vol. Ill, No. 2, pp.

234-235, May, 1891.
Fiitorius loxgicauda Merriam : Mamnjals of Idaho, N. Am. Fauna, No. 5, pp. 83-84, Aug.

1891 (from mountains of Idaho).

Type locality. — San Francisco forest, Arizona (a few miles south of
Flagstaff).

Geoyraphic diHribution. — Broadly, the Sierra Kevada and Rocky

Mountain systems, reaching British
Columbia in the Itocky Mountain re-
gion, but not known north of the Sis-
kiyou Mountains in the Sierra-Cascade
system.

General characters. — Similar to Pnto-
rius longicauda in color and markings,

n.j. 12,-P. arizunensig d ad. Boulder |j„(- much Smaller in sizC.

Color. — Upper parts from occiput to
black tip of tail, raw umber brown; head decidedly darker; end of tail
black; chin and upper lip all round white; rest of under parts includ-
ing upi^er surfaces of fore feet and inner half of hind feet and ujjper
surfaces of hind toes ochrace<ms or ochraceous yellow, varying in tint.

Cranial characters. — Skull similar to tliat of lonyicauda but decidedly
smaller and less triangular; narrower across mastoids and more bulg-
ing in parietal s.

Eemarls. — Putorins arizonensis is a mountain form of longicauda^
wliich it closely resembles except in size. The type specimen, collected
by Dr. Mearns on the pine plateau of Arizona a few miles south of
Flagstaff, is an immature female and is of unusually small size. A
male obtained by him near the same place is of the normal size, as is
another male in the Department collection from Springerville, Ariz.,

JuxE,l8<)G.] SYNOPSIS OF THE WEASELS OF NORTH AMERICA.

23

collected by E. W. Nelson. Specimens from the iiortbern Eocky
Mountain region (St. Mary Lake, Montana, and Salmon River and
Palisimeroi Mountains, Idaho) dil'fer in color from the typical animal
from Arizona and Colorado, and agree with aUeni from the Black Hills
in having the upper parts strongly sntiused with golden brown, the
yellow of the under parts yellow^ rather than ochraceous, and the under
side of the tail strongly yellow on the basal half or two-thirds. The
skulls, however, lack the tiattened audital bulhe of alleni. Specimens
from the Sierra Nevada in California are hardly distinguishable from
the Kocky Mountain animal. The only apparent external differences
are that the yellow of the under jjarts reaches uj) farther under the
chin, the white of the upper lip is less extensive, and the under side of
the tail is more suffused with yellowish. But none of these characters is
constant. In one specimen from Donner, Calif. (No. 2650, female, Mer-
riam Coll.), even the white upper lip is as marked as in Eocky Mountain
specimens; it reaches all the
way round, fills the space under
the nasal pad to the nostrils,
and broadens strongly under
the eyes. In cranial charac-
ters also the differences are
slight and inconstant. The
jjostorbital processes are longer
and more slender, often becom-
ing peg like in old males. The
audital bulhe average smaller
and more convex anteriorly,
and in the female are decidedly
narrower and more subcylin-
dric. But in an adult fenmle
from Fort Klamath, Greg., the bulhe are nearly as broad as in Rocky
Mountain females. The three female skulls I have seen of the Sierra
form are decidedly smaller than females from the Eocky Mountains,

The Sierra specimens show a strong tendency to grade into, or at
least toward xantlwgenys. In nearly half the specimens examined white
hairs are present between the eyes, and in several they are sufidciently
numerous to form a conspicuous white spot, though the spot is not
large and rectangular as in true xanthogenys. The white cheek spots I
have not seen in Sierra specimens, but the brown spots behind the cor-
ners of the mouth are sometimes present (as in No. 30055, male, from
Upper Cottonwood Meadows, near Mount Whitney, Calif.).

A specimen from St. George, Utah, an old female, differs in some
respects from typical arizonensis. The skull is small and relatively
short, and the shortening is mainly in the palate and rostral part, which
measures 2 mm. less than the average of adult females of arizonensiH of

Figs. 13 and 14. — P. arizonensis cT ad.
Colo.

Boulder County,

24 NORTH AMERICAN FAUNA. [No. 11.

the same size. Moreovei', tlie ])ostorbital ]>rocesses are longer and more
slender than in any feniahvof arizoiwusi.s 1 have examined from either
the Rocky ]\rountain or Sierra systems. Externally the St. (leorge
specimen diflers from typical ariz<>ne)i.sis iu the following })articulars:
Yellow of nnderparts more strongly tinged with ochraceous; wliiteof
npi)er lip narrow and not reaching aronnd anteriorly; brown of npi^er
X)arts reaching down on outer side of arm to wrist; a small bi^»wu spot
bearing two bristles just behind each corner of mouth. In this respect,
and this only, it resembles xanthoffcnys ; there is no trace of white on
the cheeks or between the eyes.

Meusnrements. — Average of 5 nudes from the Rocky MountaiDs:
Total length, 385; tail vertebne, 144; hind foot, 44.5. Average of 4
females: Total length, 358; tail vertebra^, 130; hind foot, 40.

PUTORIUS ALLENI sp. nov. T.lack ] fills Weasel.

Type fi'om Custer, Black Hills, Soutli Dakota. No. ;}f3ft, <? ad., Merriain collection.
Collecteil July 12, 1888, hj Vernon Bailey. Original No. 90.

Geogra'phiv disfnhiition. — Black Hills, South Dakota.

Characters. — Similar to i*. arizonensis in size and general characters,
but upper parts nu)re suffused with yellowish and audital bulhe Hatter.

Color. — Upper parts from occi])ut to black tip of tail golden or yel-
lowish-brown, in some lights with an olivaceous tinge; head dark
brown, without yellowish tinge; upper lip and chin white; rest of
nnderparts, including inner sides of legs, whole of fore feet, toes of
hind feet and under side of basal part of tail, intense buffy yellow.

Cranial cJiaracters. — Skull similar to that of arizonensis, but audital
bulla^ much flatter and somewhat smaller; brain case slightly flatter
and bulging laterally immediately behind constriction; frontal some-
what broader interorbitally ; skull as a whole shorter. The skull of an
old female (No. 7441, Am. Mus. JSTat. Hist.) is much smaller than the
male, and the audital bulla? are narrow and not flattened. In both
sexes the postorl)ital processes are strongly develoj^ed.

Roitarls. — Putorius aUeni is an isolated and only slightly differen-
tiated form of F. arizoncnsis, from which it is completely cut off' geo-
gra])hically. It is surrounded on all sides by the large weasel of the
plains, P. Jnngicanda. In worn summer pelage the color differences
that distinguish it from arizone».sis are not apparent.

I take ])leasure in naming the species in honor of Dr. J. A. Allen,
of the American IMuseum of Natural History, New York, who has
recently published an important paper on the mammals of the Blaci<
Hills, and to whom I am indebted for the loan of three additional
Sjiecimens.

Measurements (of type specimen, male adult). — Total length, 372; tail
Aertebriie, 137; hind foot, 44.

June, 1800] SYNOPSIS OF TH.E WEASFXS OF NORTH AMERICA. 25

PUTOKIUS XANTHOCtENYS ((iray). California Wca.s.l.

1843. Musiela jioiihotjoiys Gray : Annals and Magazine Nat. Hist., XI, j)]!. 118, 1813.
1857. Putorius xanthogniys Baird: Mannnals N. Am., pp. 17(5-177, 1857.
1877. Fuforiiis (Gale) hi'asiliensix frenatiis Cones: Fur-Bearint;' Animals, ])]>. lJi!-146,
1877 (in part).

Type loealify. — ►Soiitlirrn ( 'alifornin, inobably vicinity of San Diego.

Geographic distrihiitton. — Sonoran and Tinnsition iannas of ( -alifor-
iiia, on both sides of the Sierra ISTevada.

(ieneral characters. — Size medium; tail long'; face conspicuously
marked with whitish, but vest of head not black; under i)arts
ochraceous.

Color. — Upper jiarts from back of head to terminal i)art of tail in
summer pehigc raw-umber brown, tinged with golden: in n-i liter pelaije.,
drab brown, without yellowish suffusion : head always darker, becom-
ing dusky over nose; a large rectangular s[)ot between ej'es, and a
broad oblique band between eye and eir, whitish; end of tail black;
a brown spot behind c orners of mouth ; chin white ; rest of under parts,
iiiclnding- fore feet all round and inner side and toes of hind feet, vary-
ing from l)uliy ochraceous to ochraceous orange. In some specimens
the ochraceous covers the greater part of the hind feet as w^ell as the
toes.

Cranial characters. — Skull of the longicauda type and practically
indistinguishable in size and characters from /'. arizonensis; skull as a
whole short and broad ; zygomata bowed outward; postorbital processes
strongly developed; sagittal ridge distinct; andital bulhe moderate,
usually truncate anteriorly; skull of female similar to that of male,
but smaller.

Remarls. — Putorias xanthogcnys inhabits the San Joaquin and Owens
valleys and the t\ hole of southern California except the liigiier moun-
tains. In ascending' the mountains it gradually loses the facial mark-
ings and seems to grade into P. arizonensis^ the weasel o,f the mountain
summits.

Measurements. — Average of 7 males from southern California: Total
length, 402; tail vertebra', 150; hind foot, 43.5. Average of 3 fenuiles:
Total length, 3G8; tail vertebras 135; hind foot, 40.5.

rUTORIU.S XANTHOGENYS OREGONENSIS snbsp. nov. Orejion AVeasel.

Tijpe from Grants Pass, Rogue River Valley, Oregon. No. f|"is, 9 ad., U. S. Nat. Mns.,
Dept. Agric. Coll. Collected December 19, 1891, by Clark P. Streator. Original
number 1404.

(leof/raphic distribution. — liogue Iviver Valley, Oregon; limits of
range unknown.

(ieneral characters. — Similar to F. xanthogenys but decidedly larger,
darker m color, and with face markings much restricted.

Color. — LTpi)er parts m winter pelage pale chocolate brown, slightly
darker on head; a small and ill-delined ])atch between eyes, and a imr-

26 NORTH AMERICAN FAUNA. [No.ll.

row veriical bar between eye and ear, -white; throat white; rest of
under parts, including fore feet and inner sid(^s and distal half of hind
leet, i)ale yellowish; terminal one-lifth oi' tail black; rest of tail above
and below concolor with back and without the yellowish tinge which
is characteristic of xanthogcnys.

Cranial characters. — Skull similar to that of .rantliogriii/s but larger
and decidedly broader. The slvull of the type, an adnlt female, com-
pared with skulls of xantliogenys of the same sex and age I'roni south-
ern California, differs in the following particulars: Skull everywhere
broader; muzzle, palate, interorbital breadth and constriction very
much broader; zygomata more si^readiug.

3feasuremcnts. — Type specimen, female adult: Total length, 41L*; tail
vertebra^, 155; hind foot, 44. ,■,

PUTOEIUS FEENATUS (LicliteBsteiu). Bridled Woasel.
(PL III, tigs. 1, la, lb, 2.)

1813. Mustela IjraslUensiti Sevastianoff : Mem. Acad. Imp. Sci. St. Petersburg, IV,
356-363, Table iv, 1813. (Name ou plate only; diagnosis in text.) Preoc-
cupied by Mustela ij-asiUensis [an otter] Gmeliu, 1788.

1832. Mustela frenaia Liclitenstein : Darstellung neuer oder wenig bekanuter Sau-
gethiere, PI. XLII and corresjionding text (unpaged), 1832.

18.57. rutorius frenatus Baird: Mammals N. Am., 173-176, 1857.

Tyj)e locality. — Valley of Mexico, near City of Mexico.

General characters. — Size large; tail long; its black tip relatively
short; head black, with conspicuous white markings.

Color. — Top of head blackish, interrupted between eye and ear by a
broad, whitish band, which is nearly confluent with a x)atch of same
color between the eyes; rest of upper parts brown; a dark spot behind
corners of mouth; chin and throat whitish; rest of under parts ochra-
ceous yellow; forefeet to or above wrists whitish or jjale bufly yelloAV-
ish, continuous with and shading into ochraceous of under parts; color-
of under parts extending down on inner side of hind legs and feet to
toes, which are whitish or yellowish white.

Cranial char<tcters. — Skull large and massive, with strongly devel-
oped postorbital processes, deep postorbital constriction, marked sagit-
tal crest, and peculiar audital bulhe, which are obliquely truncated
anteriorily (the inner side reaching farthest forward) and abruptly
highest on inner side, falling away suddenly on outer side so as to
form a rounded ridge along the inner side of the longitudinal axis of
the bulla. The skull oi frenatus resembles that of longicaucla, but is
considerably larger, and differs in the form of the audital bulla* just
described, and also in the extent of the postglenoid space, which is
much larger than in longicauda. The dentition is heavy and the
upper carnassial tooth relatively shorter than in longicauda. The
ramus of the under jaw is much more convex interiorly.

Jiemarl's. — Liclitenstein, in his original (lescrii)tion oY j\h(stelaj'rcnata,
states that the tail is about one-third longer than that ol' the European

Juml896.1 SYNOPSIS OF THE WEASELS OF NORTH AMERICA. 27

weasel {erminea) ; that only its extreme tip is black ; that tlieliead, ears,
aD(l crown are black, this (loloring' f^idiiiij into the reddish brown of the
upper iKirts on (he back of the head behind the ears; that the facial
markings, throat, and breast are white; the remainder of the under
parts ocher yellow. The white spot between the eyes is described as
heartshaped, and in the colored plate it is shown to be nearly, but not
quite, continent with the white i^atc^h between the eye and ear. Tlie
colors in the plate are not good, as the whole under parts are white
instead of ocher yellow, and the black tip of the tail is not shown. The
specimen seems to have been in worn pelage. Liclitenstein had two
specimens, both collected by Depi)e near the City of Mexico.

Fortunately, the Department collection contains two specimens col-
lected by E. W. Xelson at Tlalpam, in the Yalley of Mexico, which may
be considered topotyi)es of frcnatus, for they not only came from the
same locality as Lichten stein's types, but also agree essentially in every
detail with his excellent description. The only points in Avhicli the
description fails to agree absolutely with the specimens is that in the
hitter the white of the throat is less x)ure and the black tip of the tail
perhaps a tritic more extensive than one would infer from the descrip-
tion; but the throat is white in contrast with the strongly ochraceous
yellow of the rest of the under parts, and a si)ecimen in the United
States National Museum from the City of Mexico (No, lOGO, 9 ad.,
J. Potts) has both throat and breast white, as in the original description.

The statement that only the extreme tip of the tail is black was made
in comparison with the European weasel {erminea), in which nearly half
of the tail is black. Hence the description agrees entirely with the
S])ecimens in hand. One point not mentioned in the description is
shown in the plate, namely, that the hind feet and toes are in large
part whitish or yellowish white. The quantity of white is variable.
In a young male from Tlalpam (No. 508U7) it is restricted to the inner
side of the foot, hardly reaching the toes, while in an adult male from
the same locality (No. 50826) it includes the toes. The whitish sjiot
between the eyes is also variable, both in form and extent. Liclitenstein
described it as lieart-sha]>ed, and his figure shows that it is narrow
where it approaches closest to the stripe between the eye and ear, with
which it is nearly, but not quite, contluent. This is precisely its coii-
■ dition m the adult male from Tlalpam, which may be considered a
duplicate type of the species. In this specimen the median white S])ot
is almost divided by the dark color of the forehead, which pushes down
between the eyes, so that the whitish spot might be described as a
narrow stripe over each eye, the two becoming confluent below. In
the young specimen the white spot is subrcctangular and not divided
by tlie black of the forehead.

Note on Putorivs hrasiliejisis. — In 1813 a Ilussian naturalist, Sevas-
tiauoff, gave the name ^MuHteJa bra.silien.sis' to a weasel brought to
St. Petersburg by Capt. A. J. Krusensteru oji his return from a voyage

28 NORTH AMERICAN FAUNA. INo.lL

arouud the world. The aiiiuuil Avas said to liave come from Brazil, but
110 definite locality was ijiveii. In the iininerous ])nl)licatioiis that have
since api)eared relating to the mammals of Brazil and adjacent terri-
tory, no weasels are mentioned as inhabiting that country, and. the
species described from the mountains to the westward differ so widely
from Sevastianoflf's hrasilien.sis that it is almost certain his animal did
not come from Brazil, The original description (including measure-
ments) agrees in every respect with P. frenaivH of Lichtenstien from
the Valley of Mexico, indicating that tlie two animals are identical.
On this assumption the well-known and appropriate name frenafus
would have to fall before the earlier and inappropriate ^hrasilie)isu.^
Fortunately, however, Sevastianoff placed his animal in the genus
Mustela, and the name ilf».S'^e/rt hrasiliotsis is preoccupied by Gmeliu
for a South American otter. (Syst. ISTat., ed. 13, ]). 93, 1788.) Hence,
unless some earlier name is found, frenatus will stand for the Mexican
bridled weasel.

Measurements. — An adult male from Tialpam, Valley of Mexico (type
locality): Total length, 505; tail vertebra^ 203 ; hind foot, 53. Average
of G males from Brownsville, Tex. : Total length, 188 ; tail vertebra', 192;
hind foot, 51. Average of 3 females from Brownsville: Total length,
438; tail vertebra?, 187; hind foot, 41.5.

PUTOKIUS FKENATU8 (tOLDMANI sul.sp. nov.

Tijpe from Pinahete, Cliiapas, Mexico. No. 77519, ^ ad.. V . .S. Nat. Mus., Dept. Agric.
coll. CoHei-ted Feb. 10, 1896, by E. A. (4ol(linan. Altitude about 8,200 feet ( = 2,500
meters). Original number 9279.

Geof/raphh' (lisfrihution. — Mountains of southeastern Chiapas; limits
of range unknown.

General characterfi. — Similar to P./renaUis in size and general char-
acters, but tail and hind feet longer; light markings more restricted;
black of head reaching much farther back on neck; color of upper parts
darker and more extensive, encroaching on sides of belly and covering
fore and hind feet; black tip of tail longer.

Color. — Upper parts, including whole of fore and hind feet, dull, dark
chestnut brown, washed with black on the neck from shoulders forward,
and becoming pure black on the head; face marked by a whitish i)atch
between the eyes, and a narrow, oblique band between eye and ear; a
blackish spot behind angle of mouth; color of under i)arts salmon
ochraceous, reaching wrists interiorly, but not reaching heels; terminal
third of tail black.

Cranial characters. — Skull rather large; zygomata moderately spread-
ing; squamosal inflation moderate, but large for a member of the/re-
natns series; audital bulla' small, steep on inner side, and only slightly
elevated anteriorly above squamosal inflation. The skull as a whole
resembles that of frenatus, but differs conspicuously in the greater
length and inflation of the postglenoid part of the squamosal, greater
breadth of the basioccipital, and in the size and form of the audita!

Jlne,1896,] synopsis of the WEASELS OF NORTH AMEIilCA. 2!)

bill lie. The latter are very narrow, low anteriorly where they meet the
iiitlated squamosal without an abrupt step, and high along the inner
side.

Kemarloi. — Mr. E. W. Nelson writes me that this tine weasel is found
sparingly in the forest about Pinabete, Chiapas, at an altitude of 7,0(10
to 8,000 feet (2,100 to 2,500 meters). The type specimen was shot in
the afternoon while hunting on a lieavily wooded hill slope. It was
heard making long, slow leaps over the dry, crisp leaves. Coming to a
log, it stood up and rested its fore feet on the log, in which position it
was shot by Mr. (ioldman.

A specimen from Cerro San Felii)c, Oaxaca, is intcrniediiitc, both
ill coloration and cranial characters, between typical freiKitKn and
(johlmdni ; hence there is little room for doubt that complete inter-
gradation exists between the two.

Measurements. — Type specimen, male adult: Total length, 50f; tail
vertebrie, 201 ; hind foot, 58.

PUTORIUS FRENATUS LEUCOPARIA sulisp. nov.

Type from Patzcuaro, JSIiohoacau, Mexico. No. 'm\i, ^ ad., U. S. Nat. Miis., Dept.
Agric. coll. Collected July 27, 1892, l)y E. W. Nelson. Original number 2i»G0.

General characters. — Similar to Putorius frenatus, hut slightly larger;
black of head extending posteriorly over neck; white face markings
much more extensive; the spot between the eyes very much larger and
broadly confluent on both sides with whitish area between eye and ear,
which area also is much more extensive in all directions than in
frenatus.

Color. — Upper parts from shoulders to black tip of tail, dark brown;
neck, crown of head, nose, ears, and sides of face to a little behind the
eye, black ; black of head between eyes and ears divided by a broad
band of bufty white which is broadly confluent with butty yellow of
throat and chin; a narrow border of whitish on upper lip; rest of
under parts ochraceous yellow (including whole of forefeet, inner sides
of hind legs and feet, and terminal half or nearly half of upper surfaces
of hind feet, where the color becomes paler, being butty ochraceous, as
on the throat).

Cranial characters. — Skull similar to that of fniKd us, Imt larger;
audital bulhe much narrower; postorbital processes less strongly
developed.

Remarlcs. — This handsome weasel presents the maximum of black
and white markings known in the froiatus group, the black of the head
reaching back over the neck and the white face markings covering a
large area. In the type specimen a white stripe 50 mm. in length
extends down the middle of the nape from a point between the ears
more than halfway to the shoulders. This, liowever, is probably ab-
normal, though a trace of it exists in a female from the same locality.
This form is the poorest subspecies described in the present paper.

30 NORTH AMERICAN FAUNA. [Xo.li.

Me<(siircments. — Average of 2 males from Patzcuaro (type locality):
Total length, ~iU); tail vertebra", 201; liiiid foot, 53. An adult female
from same place; Total length, 400; tail vertebra-, 150; bind foot, 42.

rUTORIUS TROPIC ALIS sp. uov. Tropiuul Bridled Weasel.

(PI. Ill, tigs., 5, rm, 6, (irt.)

Type iVom Jico, Vera Cruz, Mexico No. oliW, <^ ad., U. S. Nat. Mu.s., Dept. Agric.
coll. Collected July 9, 1893, by E. W. Nelson. Altitude 6,000 feet ( = 1,800 meters).
Original number 5195.

Geographic <lisfribution. — Tlie tropical coast belt of southern Mexico
and Guatemala from Vera Cruz southward.

General characters. — Similar to FutoriuH frenaiasj but much smaller
and darker, with the white face markings less extensive, the belly pale
orange instead of ochraceous, and under side of tail very much darker.

Color. — Upper parts deep umber brown with a fulvous tone; head,
ears, and neck, black, passing gradually into brown of back just iu
front of the shoulders; terminal one-fourth (or a little more) of tail,
black; face markings as in froiatus, but less extensive and whiter;
under parts ochraceous buff on throat and fore feet, becoming rich
orange buff on belly and inner side of thighs, whence (becoming paler)
the color reaches out in a narrow interrupted stripe along the inner
side of the hind feet to the toes, which are irregularly bufiy.

Cranial characters. — Skull of male similar in general to that of frc-
natiis, but smaller, relatively longer, with less spreading zygomata, less
strongly developed postorbital processes, and probably broader postor-
bital constriction (the type skull was infested with parasites) ; audital
bullae smaller and very much narrower; carnassial teeth and upper
molar smaller. The skull of the female is very much smaller than that
of the male, and has the smoothly rounded brain case of the cicognani
group, without trace of a sagittal ridge. The squamosals are strongly
inflated, resembling thosG of cicognani and the female of novehoracensis.
It differs from the female frenatus in much smaller size, very much
smaller audital bulhe, more inflated squamosals, smoothly rounded
brain case without trace of sagittal crest, and broader interorbital
constriction, which is immediately behind x)ostorbital processes instead
of one-fifth the distance from the processes to the occipital crest (fig. 15).

Remarls. — On first examining the skins of this weasel sent home by
Mr. Nelson, I supposed it to be merely a tropical subspecies of frenatm;
but on comparing the skulls I am forced to accord it full specific rank.
The difference is greatest in the females, and is really very remarkable,
as may be seen from the accompanying figures (figs. 15 and 16). The
female of frenatus (fig. 1(>) resembles the male of the same species (pi. Ill,
fig. 1), while the fennile of trojricalis (fig. 15) resembles the cicognani
group — representing another section of the genus. The case is parallel
to that of r. noveljoracensis already described. The female of trojncalis,
like that of novehoracensis, shows arrested development or absence of

Ju.NE,lS96.] SYNOPSIS OF THE WEASELS OF NORTH AMERICA.

31

Fig. 15 — P. frenatus

Fin. 10. — 1'. Iriqncalis ? .

the specialization that characterizes the male, while the females of
icashin(/toni and frenaUis have advanced further and are moie like
the male. In the case of the female skulls of frenatus and tropicalis
here figured, it is interesting to know that they were taken within
a few miles of one another — frenatus on Cofre de Perote. at an
altitude of about 12,500 feet;
tropicalis at Jico on the plain
below, at an altitude of 5,000 or
G,000 feet.'

The Department collecti<ui
contains four specimens of this
weasel, all collected by Mr, Nel-
son in A'era Cruz. Three of
t-liem, two adult males and one
old female, are from Jico; the
fourth, an immature female, is
from Catemaco, and presents the
extreme of differentiation in in-
tensity of color. The hind feet
are dark tliroughout and the color of the upper parts is peculiarly dark
and rich, as in P. affinis.

Measurements. — Average of two adult males from Jico, Vera Cruz
(type locality): Total length, 442; tail vertebrie, 175; hind foot, 50.
An old female from same place: Total length, 333; tail vertebrt^, 121;
hind foot, 37.

PUTORIUS AFFINIS (Gray).

Musfehi affinis Gray: Aimals & Mag. Nat. Hist., -ith ser., XIV, p. 375, Nov.

Type locality. — "New Granada" [= Colombia].

General characters. — Size large; tail long; color very dark
black anteriorly; facial markings obsolete or nearly so.

Color. — Upper jiarts nearly pure black on head and neck, fading
imi)erceptibly to rich blackish brown on back, rump, and tail; black
tip of tail long, but not strongly contrasted with dark color of rest of
tail; under parts narrowly ochraceous orange, narrowest behind angle
of mouth, where it is encroached on by the blackish of the cheeks. Face
usually unmarked, but a whitish streak sometimes present in front
ot ear.

Cranial characters. — The only skull of this weasel I ha\e seen is from
a skin (No. 13770, U. S. Nat. Mus.) collected by Dr. Van Patten, at San
Jose, Costa Rica. It is immature, but differs strikingly from frenatus
in the greater breadth of the frontal region and the flatness of the
audital bulla'. The constriction is little marked, which may be due to

1S74.

almost

'The ditiereuce in size of the two species is well shown by the iie.sh measurements
of these two specimens. Femaile frenaf us, Cofre cle Perote: Total length, 418; tail
vcrtebrre, 100; hind foot, 45. Female trojncalis, Jico: Total length, 333; tad verte-
bra-, 121 ; hind foot, 37,

32 NORTH AMERICAN FAUNA. [No.il.

parasites in the frontal sinuses. The young skull af!'ords the, following
measurenieuts: Basal length, 50: zygomatic breadth, 29', postjjalatal
length, 20; i>alatal length, 24 ; iuterorbital breadth, 12; breadth across
postorbital i)rocesses, 15; breadth of constriction, 14.

General ronarls. — There are several specimens from Oosta Kica iu
the National Museum collection which ajiparently belong to this
species. In these specimens the color of the upper parts is exceed-
ingly dark from the color of the tips of the hairs; but the color imme-
diately underlying the black tips is deep fulvous brown, giving a very
rich tone to the pelage. The orange of the under parts is narrow and
does not reach the feet; on the hind legs it stops on the thighs, and on
the forelegs it stoi)s short of the wrists.

McdsxrciiK'itts (from dry skins in U. S. Nat. Mus.). — Total length,
about 510; tail vertebra\ about 180; hind foot, about 52.

JUKE,189C.] SYNOPSIS OF THE WEASELS OF NORTH AMERICA. 33

Tahh of averarje cranial meanHrementu of North American WcaseU.

F. cicognani

Locality.

P. richardsoni.
P. alascensis . . .
P. ttreatori

P. rixosux

P.arcticvs

P.kadiace7isis

P. novehoracetuis

P.xvashiiiijtoni .. ..

P.peninndce

P. longicaxida

P. spadix

P. saturattis

P. arizonensis

P. alleni

P.xanthorienys...

Pfrenatus

/'. tropicaUs

Ossipee, N. H

Elkliiver, Minn

Do...

Mount Forest, Ontario. .

Great Slave Lake

Juneau, Alaska —

Skagit Talley, "SVash . . .

Do

Trout Lake, Wash

Do

Osier, Saskatchewan . . .
Point B:!rrow, Alaska . .
Franklin Bay, Arctic

Coast.
St. Michaels, Alaska . .

Do

Kailiak Island, Alaska.
Adirondacks, X. T ,

Do

Trout Lake, "Wash

Do

Tarpon Springs, Fla...
Carlton House, Sas-
katchewan.

Do

ElkEiver, Minn

Do

Siskiyou Mountains,
Oregon.

Springei-ville, Ariz

Boulder County, Colo..
Sierra Kevada, Cal

Do

Black Hills, S. Dak....

Do

Southern California

Do

Tlalpam, Mexico

Cofre de Perote, Mexico
.Tico, Tera Cruz, Mexico

Do..

M

^K

pa

38. 5^ 37. 5
40. 2 39
33.5, 32.5
32.5^ 31.5

40
42
34
29.5

33.5 32.5

28. sj 28
2G.5126

44.5 43

43.5' 42

43
38
'42

47

42
37
'41
45.5

38.5 37.5
44.2 43
38. 3 37. 5
45.5 44
48 47

17.5

24

24.5

20

16.5

18

15.5

14.2

29.5

27.5

26.5

22.5

24

27

20

26

2L5

27

30.5

^

§"2

18.5 10.5
19.5 n
16 ' 10
16 i 9
20.5 11.5
21 I 14

f^ -<

13 i

'' I
IG. 5

13.51

13.5

23 I

22. 5 j

22. 5

19

20.5

23.5

18. 5I

23 I
I
20

24

26

8.7 25.5
26.5

7.8 22

7 j 21.5
9.7 27

11 i 28.5

8.5j 23

7.5 20

8 22
6.5 19
5.5, 17.5

11

10

9.8|
8.5'
7.5
14.5' 12.5'

16.5

17
14
14
18
19
15
12
14.5

22

22.5

19. 5

18

23

24

20

18

19.5

B

4

4

13

13. 5|

12

12.5

14.5'

11

12.5

10.5

14

15.5

11 28.5

12 28
10 24. 5
10. 5 > 27
11.3 30

25.3

10.2

27.5

24

29

42.5 26 23 12 10.5 26.5

46.5

43

44

29.5 26 ! 14.5 11.5

26 I 23.5 13
29 25 14

41
43

44. 5' 43. 5,
39.5 38
42 I 40.8
38.5 37.5

42.5
41

43.5
47.5

26

28.5

28

22.5

27

23

27.5

24

33.5

25.5

28

22.5

23

23

23

20.5

22

23.5

..;

27.5.
23

24.5

19.5;

12.5

13

14.5

1L3

13.2

12

13.5

12

15.5

13

15

12

10.5
11

10.5
10.5
10.5

9
11

9

9.5

9.5
12
10
10.5

9

28
28.5

26.5
27
28
24

12.5 16.5
11 '15
20.5 24

19.5 24
19. 5I 24
16. 5' 21
17. 5' '24
21.5 25.5
16 ' 22.5'
21 23 !
17.8 20.5
21 [ 24.5
23 25

20.5 22.5
23.5 24
20.5' 23.5
21 24

20 22
20.5 23.5'

21
18

26.2 20

24.5' 18-

27.5

26

33.5

29

32

24.5

23 I

21 !

22 I
20.5

20.5 23.2
19.5' 22 j
24.5 27.5

19.5

22

16

25

27
21.5

' Estimated.

10932— No. 11-

IIsTDEX.

[Synonyms iu italics.]

Arctogale, 9.

('yuoinyonax (synonym of Piitoriias), 7.

Gale (synonym of Ictis), 9.

Ictis, subgenus, 9.

list of species, 10.
iluatela hrasiliensis, 20.
tieognani, 10.
erniinea, 9.
enninea, 11.
frenata, 26.
longicauda, 19.
ricliardsoni, 11.
vulgaris, 9.
rulgaris, 10.
xanthogenys, 25.
Putorius, genus, 7.

key to subgenera, 7.
list of species with type localities, 10.
subgenu.s, 7.

table of cranial measurements, 33.
T'utdiius atlinis, 31-32.

alascensis, 12-13.
alleni, 24.
arctic.us. 1.5-16.
ari/.onensis, 22-24.

Putorius boccamela, 9.

cicognani, 10-11.
erminea, 15, 16.
erminea, 16.
eversmanni, 8.
frenatus, 26-28.
goldmanui, 28-29.
kadiacensis, 16.
leucoparia, 29.
longicauda, 19-21.
nigripes, 7-9.
noveboracensis, 16-18.
oregonensis, 25-26.
peninsula^, 19.
piigilhts, 14.
putorius, 8.
richardsoni, 11-12.
rixosns, 14-15.
saturatus, 21-22.
spadix, 21.
streatori, 13-14.
tropicalis, 30-31.
vulgaris, 10.
washingtoui, 18-19.
xanthogenys, 25.

35

PLATE I.

Fig. 1. Putorius nigripts, $ ad., Trego County, Kaus.
(Xo. 4143, Merriam coll.)

1. Upper side of skull,
la. Under side of skull.
1?). Side view of skull.

2. Putorius 2)utorius, $ ad., Brunswick, Germany,
(No. 4661, Merriam coll.)

2. Upper side of skull.
2a. Under side of skull

36

North American Fauna, No. 11.

Plate I.

1. Putorius nigripes cf ad. Trego County, Kansas.

2. Putorius 2)utoi-ius (f ad. Brunswick. Germany.

PLATE II.

Fig. 1. I'liforius arcticus. Poiut Barrow, Altvska (type).
S ad., No, 23010, U. S. Nat. Mus.
2. Piiiorius alascensis. Juueaii, Alaska (type).

^ ad., No. 74423, U. S. Nat. Mus., Dept. Agric. coll.
3 aud 4. Puiorius cicotinani.

3. ^ ad., Biicksport, Me., No. 4247, Bangs coll.

4. 5 ad., Mount Forest, Ontario, No. 789, Bangs coll.
5 aud 6. Puiorius strcatori. Mount Vernon, Skagit Valley, Wash.

5. ^ ad., No. 76646, U. S. Nat. Mus., Dept. Agric. coll. (type).

6. 5 ad., No. 76623, U. S. Nat. Mus., Dept. Agric. coll.
7. Puiorius rixosus. Osier, Saskatchewan.

5 ad., No. 642, Bangs coll. (type).
38

North American Fauna, No. 11.

Plate II.

1. Putorius arcticus.

2. P. alascensis.

7. P. rixosus.

3, 4. P. cicognaui.
5, 6. P. streatori.

PLATE III.

Figs. 1 and 2. Putorius frenatus.

1. (? ad., Tlalpam, Mexico, No. 50826, U. S. Nat. Mus., Dept.

Agric. coll.

2. 9 ad., Cofre dc Perote, Vera Cruz, Mexico, No. 54278, U. S.

Nat. Mus., Dept. Agric. coll.
3 and 4. Putorius longicauda. Carlton House, Saskatchewan (type locality),

3. <? ad., No. 73183, U. S. Nat. Mus., Dept. Agric. coll.

4. 9 ad.. No. 75483, U. S. Nat. Mus., Dept. Agric. coll.
5 and 6. Putorius tropicalis. Jico, Vera Cruz, Mexico.

5. S ad., No. 54994, U. S. Nat. Mus., Dept. Agric. coll. (type).

6. 9 ad., No. 54993, U. S. Nat. Mus., Dept. Agric. coll.
40

North American Fauna, No. 1 1

Plate III

1, 2. Futorius J'renati

3, 4. P, loiKjicauda.

G. P. tropicaUs

PLATE IV.

Figs. 1 and 2. ritiorius noveboracensis. Adiroudacks, New York.

1. S ad., No. 3843, Merriam coll.

2. 9 ad., No. 5598, Merriam coll.

3 and 4. Putoriu8 tvashingtoni. Trout Lake, Washington.

3. <? ad., No. 76322, U. S. Nat. Mus.. Dept. Agric. coll. (type),

4. 5 ad. No. 67321, U. S. Nat. Mus., Dept. Agric. coU.
5. Puforiiis peninsula'. Tarpon Springs, Fla.

9 ad.. No. 2.379., Ehoads coll.
42

North American Fauna, No. 1 1 .

Plate IV.

1, 2. Jr'uturius )Lovcbonict')isis. y, 4. f, wushiiKjtoni. 5. F, peninsula:.

PLATE V.

Fig. 1. Putorius longicaiida (Bonap.)-

1. S 'iil-j Cai'ltou House, SaskatclieTvan, No. 73183, U. S. Nat. Mus.,

Dept. Agric. coll.
la. Q ad., Carltou House, Saskatchewan, No. 75483, U. S. Nat. Mus.,
Dept. Agric. coll.

2. Putorius cicofjuani (Rouap.).

2. J, Bucksport, Me. No, 4247, Bangs coll.

2a. 9, Mount Forest, Ontario No. 789, Bangs coll.

3. Putorius noveboracensis De Kay.

3. S ad., Adirondacks, New York No. 3843, Merriam coll.
3a. 9 ad., Adirondacks, New York No. 5598, Merriam coll.

4. Putorius rixosHs nob.

9 ad. (type), Osier, Saskatchewan, No. 642, Bangs coll.

5. Putorius 2>eniH8nlw l»hoads.

9 old, Tarpon Springs, Fla. No. 2379, Khoads coll.

6. Putorius aicficus sp. nov,

6. <? , St, Michaels, Alaska No, 313213, U. S, Nat. Mus.
6rt, 9, St. Michaels, Alaska No. 36246, U, S. Nat. Mus.
44

k-

North Ameiican Fauna, No. 11.

Plate V.

U. S. DEPARTMENT OF AGRICULTURE
DIVISION OF ORNITHOLOGY AND MAMMALOGY

NORTH AMERICAN FAUNA

INTo. 1

o

[Actiuil tUite of publication, July 23, 18116.]

GENERA AND SUBGENERA OF VOLES AND LEMMINGS

]".Y

GERRIT S. MILLER, Jr.

Prepared under the directiuu of

Dr. C. HART MERRIAM
CHIEF OF DIVISION OF ORNITHOLOGY AND MAMMALOGY

WASHINGTOi^

GOVERNMENT PRINTINO OFFICE
189G

LETTER OF TRANSMITTAL.

United States Depart3Ient of Agriculture,

Division of Ornithology and Mammalogy,

Washington, D. C, ^laij 12, 189G.
Sir : I have the bouor to trausmit herewith, and recommend for pub-
lication, the manuscript of l!^o. 12 of North American Fauna, treating
of the Genera and Subgenera of Voles and Lemmings, and comprising
results of investigations carried on in the Division of Ornithology and
Mammalogy by Gerrit S. Miller, jr.

Eespectfully,

C. Hart Merriam,

Clilef of Division.
Dr. Chas. W. Dabney, Jr.,

Acting Secretary of Agriculture.

3

CONTENTS.

PagR

Introduction.... 7

The subfamily Microtincr and its main divisions 8

List of genera and subgenera of Microthup 9

Geographic distribution 9

Habits 10

Nomenclature 11

History of former classifications 19

Characters on •svhicli the jiresent classification of the sul)geuera oi Mi trot us is

based 24

Keys 28

Descriptions of living genera and snl)geuera 32

Descrijitions of extinct genera and suijgenera 73

Note on 'Arvicohi ' iniermedius Newton 75

ILLUSTRATIONS.

PLATES.

1. Skulls of Microtus macropiiH, 3/. pinetorum, M. arvalis, M. curtatiis, M. oregoni, M.

tvrresiris, M. albicauda, 21. fertilis, Evotoinys (lapperi, Pheuacomys oramonds,
Lemmus nigrijjes, Sijnaptonujs u'ranrjeli, S. helaletcs, Dicrostonyx torquatus.

2. Hony palates of Phenacomys. Mlcrotns, Laguru-s, Pitymys, Arvicola, Alticola,

Anteliomys, Eothenomys, Evotomys, Xeofiiei; Dicrostonyx, Lemmus, Fiber.

3. Mandil)les of Sy'naptomy.<<. Phenacomys, Microtus. Evotomys.

TEXT FIGURES.

1. First upper molar of six specimens of Microtus pennsylvanicus.

2. Second upper molar of six specimens of Microtus pennsylvanicus.

3. Third upper molar of eighteen specimens of Microtus pennsylvanicus.

4. First lower molar of eighteen specimens of Mierotus pennsylvanicus.

5. Second lower molar of four specimens of Microtiis i)ennsylvanicus.

6. Third lower molar of four specimens of Microtus pennsylvanicus.

7. Palatal view of skull of Microtus arvalis and Evotomys gapperi.

8. Enamel pattern of molar teeth of Synaptomys cooperi.

9. Palatal view of skulls of Synajjtomys aud Mictomys.

10. Enamel pattern of molar teeth of Synaptomys innuitus.

11. Enamel pattern of molar teeth of Lemmus lemmus.

12. Left front foot of Lemmus lemmus.

6 ILLUSTRATIONS.

13. Enamel pattern of Dicrostonyx from Ungava.

14. Ear of Dkrostonjix and Lemmus.

15. Left front foot of three sjjecimens of iJicrostonyx from Alatska, showing seasonal

change in form of middle claws.

16. Side view of molars of adult and young Phenacomya. \

17. Enamel pattern of molar teeth of Fhevacomys celatua.

18. Side view of molars of adult and youug Evoiomys.

19. Enamel pattern of molar teeth oi Evotomys gapperi.

20. Side view of molars of adult Microtus.

21. Left front foot of Microtus terrestris.

22. Enamel pattern of molar teeth of Microtus {Eothenomys) melanofjaster.

23. Enamel pattern of molar teeth of Microtus (Anteliomys) chiiiensis.

24. Audital bulla' of Microtus (Microtus) arralis and M. (Layurus) 2}allidus.

25. Enamel iiattern of molar teeth of Microtus (Layurus) luteus, M.{L.) lagurus, and

M. (L.) palUdus.

26. Enamel pattern of molar teeth oi Microtus (Jliicola) alhicanda.

27. Audital bullfe of Microtus (Alticola) albicauda and M. {Hyperacrius) fertilis.

28. Enamel pattern of molar teeth of Microtus {Hyperacrius) fertilis.

29. Enamel pattern of molar teeth of Microtus (Pedomys) austerus,

30. Enamel pattern of molar teeth of Microtus (Phaiomys) strauchi.

31. Enamel iiatteru of molar teeth of Microtus {Pitymys) pinetorum and M. (P.) savii.

32. Enamel pattern of molar teeth of Microtus (Chilotus) oregoni.

33. Enamel pattern of molar teeth of Microtus (Microtus) arvalis, M. (M.) nivalis, M.

(M.) penufsylvanicus, and J/. (M.) ratticeps.

34. Enamel pattern of molar teeth of Microtus (Arvicola) terrestris and M. (A.)

macropus.

35. Enamel pattern of front lower molar of type of Microtus arvicoloides.

36. Enamel pattern of molar teeth of Microtus (Xeofiber) alleni.

37. Dorsal view of skull of Fiber.

38. Side view of molars of adult Fiber.

39. Enamel pattern of molar teeth of Fiber zibethicus.

40. Enamel pattern of molar teeth of '^/'luco/a' interrnedius.

No. 12. NORTH AMERICAN FAUNA. July, 1896.

THE GENERA AND SUBGENERA OF VOLES AND LEMMINGS.

By Gekrit S. Miller, Jr.

The followiug revision of the genera and subgenera of voles and lem-
luiugs is chiefly the result of a study made in the Division of Ornithol-
ogy and Mammalogy of the collections belonging to the United States
Department of Agriculture. This material has been supplemented by
specimens from my own i^rivate collection and those of Mr. Outram
Baugs, Mr. S. N. lihoads, and Dr. C. Hart Merriam. I have also had
access to the voles and lemmings in the American Museum of Natural
History, the United States iSlational Museum, and the British Museum.
Thanks are due to all who have placed material at my disposal, and
especially to Mr. Oldlield Thomas, curator of mammals in the British
Museum.

Hitherto no attempt has been made to com])are in detail the voles
and lemmings of the Old and New Worlds. This is the necessary result
of the poor quality and small number of specimens from the opposite
side of the Atlantic to be found in museums and private collections in
both Europe and America. In consequence of this lack of material,
writers who have been thoroughly acquainted with indigenous voles
and lemmings have either made no comparison of these with exotic
forms, or have reached faulty or at least incomplete conclusions with
regard to groups occupying widely separated geographic regions.

For determining the relationships of the different voles and lemmings
the collection in the British Museum offers exceptional facilities. It
contains representatives of all the recent genera and subgenera found
in the Old World, and lacks only one of those peculiar to America.
The collection is, moreover, especially rich in specimens identified by
the more prominent writers on the subject — a circumstance of the
utmost importance.

The drawings for the illustrations in this paper, except fig. 9 and
Pis. I and II, were made under my constant supervision by Mr. F.
Miiller. Pis. I and II were prepared by Dr. James 0. McOonnell.
Figs. 4, 5, 8, and 10 of PI. II were drawn in ink by Dr. McConnell from
pencil drawings made at the British Museum by Mr. Hollick. Fig. 7
of the same plate is by Dr. ]McConnell from a pencil drawing by Mr. A.

7

8 NORTH AMERICAN FAUNA. [X0.12

Westergreii. The tracings of the enamel pattern of Microtus htteiis
and ]\L lagnrns are enlarged from flgs. 10, 11, 15, and 16 of PI. XIII of
Biicbuer's 'Wissensclmftliclie Eesnltate der von N. M. Przewalski nacb
Central-Asien nnternommenen Eeisen.' In fig. 22 the enamel patterns
of the front lower molar and middle and back upper molars are enlarged
from Mr. Hollick's pencil drawing of a specimen from Fokien, China
(British Mnsenm Register 92. 10. 12. 52), the other teeth from fig. 1, PI.
XLYI of Milne-Edwards's '■ Recherches jiour servir a 1' Histoire Natnrelle
des Mammiferes.' Fig. 23 is compounded in the same way from Mr.
Hollick's drawing and the original figure published by Thomas.

THE SUBFAMILY MICROTINyE AND ITS MAIN DIVISIONS.

The subfamily Microtince^ is a group of murine rodents closely related
to the Keotomincc, Cricefince, and MyotaJpincv.'^ It is distinguished from
the first and second by cranial and dental characters; from the last
chiefly by peculiarities in external form.^ While it is not the purpose
of the present paper to discuss the relationships of the Microtina' to
any of these, it is important to consider in some detail the larger divisions
of the subfamily itself before taking up the genera and subgenera.

The members of the subfamily Microtina' fall naturally into two
supergeneric groups, the Lemmi and Microti, or lemmings and voles.
The former includes the genera Synaptomys, Lemmus, and Dicrostonyx,
the latter the genei'a Phenacomys, JEvotomys, Microtus, and Fiber.

Lemmi. — Skull generally broad and massive; lower incisors short,
with roots ending on inner side of molars (PI. Ill, fig. 1); crowns of
maxillary teeth scarcely, if at all, narrower posteriorly than anteriorly
(figs. 10, 11, and 12); tail usually shorter than hind foot (in Synaptomys
slightly longer); palms and soles usually without distinct tubercles.

Microti. — Skull comparatively slender and lightly built: lower
incisors long, with roots ending on outer side of molars (PI. Ill, figs. 2
and 3); crowns of maxillary teeth distinctly narrower posteriorly than
anteriorly (figs. 17, 19, 21-35); tail usuallj' much longer than hind foot
(in the Asiatic species of Lngurus distinctly shorter) ; palms and soles
always with distinct tubercles.

In external appearance the lemmings and voles differ considerably.
The former are mostly thick-set animals, with powerful fossorial feet,
long, dense fur and very short tails, while the latter are more slender,
with longer tails and with the fur and feet not so highly modified.

^ =^ ArrieoUna' Auct. This name, however, must be abaudoned, together with the
generic name Arvicola (see p. 14).

'^ = Sij)hneinct3 Auct. As SipJineus (Brants, 1827) must give place to Myotalpa (Kerr,
1792) (see Allen, Bull. Am. Mus. Nat. Hist., New York, VII, p. 183, 1895) it is neces-
sary to make a corresponding change in the name of the subfamily.

•The characters separating the Miiotalpinw from the Microtiniv are of much less im-
portance than those separating the latter from auy of its other allies. So close, indeed,
is the resemblance between the two that it may eventually prove necessary to unite
them under one name. Lack of material jjrevents auy final conclusion at present.

Jl-LY, 1896.1

LIST OF GENERA AND SUBGENERA.

9

Altbougb the voles and leuimiiigs may usually be distinguished at a
glauce, there are certain genera and subgenera the exact position of
which is not at first apparent. Thus the species of Lagnrns, although
volCvS, so closely resemble lemmings in exteriml appearauce that their
true relationships have been only very recently detected. On the
other hand, Synaptomys, a true lemming, has much the superficial
appearance of certain forms of Microtus.

LIST OF GENERA AND SUBGENERA OF MICROTIN^.

Genera.
Synaptomys.

Lemmus.

Dicrostonyx.

Phenacomys.

Evotomys.

Microtus.

Subgenera.
Mictomys.

Eotbenoruys.

Anteliomys.

Lagurus.

Alticola.

Hyperacrius.

Pliaiomys.

Pedomys.

Pityrays.

Chilotus.

Microtus.

Arvicola.

Neofiber.

Fiber.

Tyi)es.
Synaptomys cooperi.
Synaptomys inuuitus.
Lemmus lemmus.
Dicrostonyx torquatus.
Pbenacomys intermedins.
Evotomys rutilus.
Microtus arvalis.
Microtus melanogaster.
Microtus cbinensis.
Microtus lagurus.
Microtus stoliczkanus.
Microtus fertilis.
Microtiis blythii.
Microtus austerus.
Microtus pinetorum.
Microtus oregoni.
Microtus arvalis.
Microtus terrestris.
Microtus alleni.
Fiber zibetbicus.

The following groups are known to occur in both hemispheres:

Lemmus. Microtus (gouns and subgenus).

Dicrostonyx. Lagurus.

Phenacomys? Pitymys.

Evotomys. Arvicola.

The following groups have been found in the Old \Torld only:
Eothenomys. Alticola.

Anteliomys. Hyperacrius.

Phaiomys.

The following groups have been found in America only:
Synaptomys. Cbilotus.

Mictomys. Neofiber.

Pedomys. Fiber.

GEOGRAPHIC DISTRIBUTION.

The subfamily Microfinw is distributed throughout the extratropical
region of the Northern Hemisphere. In the north some members of
the group approach the extreme limit of mammalian life, while in the
south a few si)ecies enter the northernmost edge of the tropics. The
subfamily, which is clearly boreal in origin, reaches its highest develop-

10 NORTH AMEKICAN FAUNA. [No. 12.

ment in temperate Europe, Asia, and North America. Altliougli it is
probable that no species are common to both continents, five f?enera
and four subgenera of the genus MlcrotKs have a circumpohxr distribu-
tion. On the other hand, no genera are peculiar to the Old World, and
only two are confined to America. Asia has five subgenera of Microtm
not found in America, and America has three not known to occur in the
Old World.

HABITS.

The voles and lemmings occur in great abundance throughout the
region which they occupy. They live in an endless variety of situa-
tions, from sea beaches to marshes and Alpine mountain tops, and from
open ijlains to the densest forests. They are, perhaps, most numerous
in Avell-watered grass lands. In localities where they are abundant
most of the species make their presence Icnown by trails or runways
traced through the vegetation near their barrows. Occasionally, how-
ever, they occupy hollows in decaying logs or among loose rocks, and
use natural crevices instead of beaten paths. While the great majority
of species spend much of their time on the surface, protected by the
overhanging vegetation, a few live almost exclusively underground,
and in consequence of this habit have ac(iuired numerous modifications
which fit them for the needs of a subterranean life. Others are
amphibious and never occur at any great distance from water. At
least one member of the subfamily^ is said to live among the branches
of trees. The food is chiefly vegetable, though most species occasion-
ally eat animal food. The vegetable food consists principally of grass
stems, though roots, bark, leaves, seeds, and fruit are at times eaten in
varying (quantities. xVs voles are readily caught in traps baited with
meat, it is probable that flesh forms part of their normal food. Mollusks
are eaten freely when they can be obtained.

The voles and lemmiugs breed very rapidly during the warmer part of
the year. The number of young in a litter varies from one or two to ten.
Five is, perhaps, the average number in the majority of species, though
it is probably less in those in which the females have only four mamma?.

' Phenacomys longicauda True, from Oregon. In the original description of the
species (Proc. U. S. Nat. Mu.s., XIII, pp. 303-304, Nov. 15, 1890) Mr. True quotes as
follows from a letter from Mr. Anrelius Todd, who collected the type specimen: "It
lives exclusively, as far as I have been able to ascertain, among the boughs and
branches of the Oregon pine trees (Abies douglasi), making a nest of a size smaller
than a robin's ne.st. It is usually situated on the ui:)per side of a medium-sized
branch, perhaps 6 inches in diameter, and is composed of the leaves of the tree
deftly split in two from one end to the other and dried. The nest is neatly and
rather ingeniously made, and the sameness of the material is a novelty. * * *
The mouse is almost exclusively arboreal in its habits, but I think that I have reason
to believe that they sometimes come to the ground for food, as I have seen tracks iu
the snow around the trees which I think were made by these little animals. They
could be tracked up and down the tree, but to no great distance from it, and were
most likelv in search of food."

July, 1896.] NOMENCLATURE. 11

The 5'oung- are boru in nests made of soft vegetable libers. The nests
are usually placed iu a burrow or beneath shelter of some kind and
vary witli the size of the animals, but are usually about 200 mm. in
diameter. The species of Fiber make nests containing several bushels
of material. These are conspicuous objects in the marshes where the
animals live. Under conditions the nature of which is not understood
the rate of increase in certain species is occasionally so enormously
accelerated that an area becomes overcrowded and the animals wander
into the surrounding country in search of food. So far as known, such
'lemming mig'ratious' and 'vole plagues' are phenomena peculiar to the
Old World.'

NOMENCLATURE.

Before considering the characters of the genera and subgenera of
Microtintc it is necessary to examine a considerable jDart of the mass of
technical literature to which, during the past hundred and forty years,
the animals in question have given rise. Since Linmieus published the
tenth edition of the Systema ]Satura) more than fifty names have been
used for the less than two dozen namable superspecific groups recog-
nizable in the subfamily. In considering their claims to recognition
the names may be best taken up chronologically.

Mns Linnajus, 1758 (Syst. Xat., Ed. 10, p. 59), contained the following
species: Porcellus, leporinus, lemmus, marmota, monax^ crieetiis, terres-
tris, ampMMus^ rattus, musculus, aveUanarius, sylvaticus, striatusj lon-
gi2)€s,jaculus, volans. Since two'^ of these {lemmus and terrestris) are

■An account of the migrations of Lemmus hmmus in Norway is given by Prof. R.
C'ollett in Christiania Videnskabs-Selskabs Forbandlinger, 1895, No. 3.

For description of a vole plague in Scotland, see Report of the Departmental
( omiiiittce appointed by the Board of Agriculture to inquire into a Plague of Field
Voles iu Scotland. London, 1893.

-Appareutly three, but terrestris and ampliihius are, as Lataste has already shown,
the same animal. The Mus amphiiius of Linnams is nothing more than a hgment of
the imagination based on Ray's misconception that there is a large aquatic vole with
webbed feet.

Since the matter is of importance as determining the validity of the current name
of one of the most common European mammals, I ([uote Linn;eus's descriptions in full :

"[Mus] terrestris, 7. M. cauda mediocri subpilosa, palmis subtetradactylis,
idautis pentadactylis, auriculis vellere brevioribus.

"Mus cauda longissima pilosa, auribus subrotundis vellere brevioribus. Fn. svec.
29. Syst, Nat., 10, n. 5.

"Mus agrestis, capite graudi, brachiuros. Raj. quadr. 218.

"Habitat iu Earop;i' terra et aqua.

"Corpus fuscum subtus pallidum, at non albicans. Caput crassius, ore gibbo.
Cauda magis pilosa, quam iu Ratto, sed corpore dimido brevior, a pedibiis fere longior.

"Hortos Talpje instar misere effodit palmis licet jiarvis; natat in fossis et urinatur
plantis licet fissis ; Radices arborum decorticat, ])lantaruui consumit s. aufert ; Pullos
anatum in pisciuis occidit.

" [Mus] amphibius, 8. M. cauda elongata pilosa, plantis palmatis.

"Mus major aquaticus s. Rattus aquaticus. Raj. quadr. 217.

"Mus aquaticus. Beil. aquat. 3.5. t. 36.

12 NORTH AMERICAN FAUNA. [No. 12.

Microtiiies, it is necessary to see whether the name can be applied to
any genus of the subfamily. Linnaeus of course designated no type,
but subsequent usage has fixed the n;mie on the congeners of 31}is mns-
cuIks. As no sound principle of nomenclature is thus violated, the
name Mus should be kept in its present signification.

Castor Linn.Teus, 1758 (Syst. ^N^at., Ed. 10, p. 58), was originally pro-
posed for the species fiber and moschatus, but in the twelfth edition
of the Systema others were included, among them the muskrat. The
name, however, could by no process of subsequent elimination be applied
to the latter.

GJis Brisson, 17G2 (Regn. Anim., pp. 13, 113), is clearly l)ased on
the dormice,' although the genus includes 'la Marmotte de Bahama,'
'laMarmotte d'Amerique,' 'la Marmotte de Pologne,' 'la Marmotte des
Alpes,' and 'la Marmotte de Strassbourg,' in addition to 'le Loir,' 'le
Lerot,' and 'le Croquenoix.' The name must, therefore, take the place
of Myoxns Schreber, 1781, commonly used for the dormice.^ As none of
the species of Brisson's Glis are Microtines, the name would not be men-
tioned here were it not for its bearing on Glis Erxleben, 1777.^ (See p. 13.)

CunicuJns Brisson, 1762 (Eegn. Anim., p. 13), must also be consid-
ered, because it invalidates the use of Cuniculus Wagler as the generic
name of a lemming (see page 16).* The genus contained an assemblage
of forms which are now put in six genera distributed among five fami-
lies. Dr. C. Hart Merriam has recently shown (Science, n. s., I, p.

[Continuation of note from page 11.]

"Habitat in Europ.-p, Africne fossis, ripis, piscinis, hortis.

" Species mihi iion rite cognita.

"Fodit ad fossas et radicis arborum, natat, urinatiir, consumit radices, Hortis et
satis infestus, capitur Nassis e virgulis confectis sub aqua demersis."

The description of Mus terrestris is extended and applies to the water rat in every
particular, while the diagnosis of M. ampldh'ms is very brief and contains a glaring
error in the assertion that the animal has webbed feet. That the common water rat
was the animal which Linnivus had in mind when he described Mus ierresiris is
shown by the length and accuracy of the description and by his choice of the sjiecific
name {Mus terrestris is the Latin e([uivalent of the Swedish jordratta). That he
never saw 'Mus amphibius' is clearly indicated by the statement: "Species mihinou
rite cognita." It is thus evident that there is no excuse for retaining the specific
name amphibius, even though the error through which it is now generally ;ised has
passed current for nearly a century.

'In the Tabula Synoptica Quadrupedum secundum Ordines Sectiones et Genera,
on pages 12 and 13, the name is introduced as follows:

Cauda longa, vestita pilis ita dispositis at caudum iilanum efflciant Sciurus

Cauda longa, vestita pilis ita dispositis at caudum rotundam efificiaut GU$

-See Merriam, Science, n. s., I, p. 376, April 5, 1895.

'^Glis Brisson also antedates Glis Storr (Prodr. Meth. Mamm. 1780, p. 39), proposed
for Mus tamaricinus, M. longipes, M. cafer, M. sagitta, M. jaculus, M. nitidula, M.
avellayrarius, and M. gJis.

•"In the synoptic table (pp. 12, 13) the name is introduced as follows:
Cauda brevissima vel nulla:

Auriculis longis Lepus

Auriculis brevibus vel nullis Cutiicuhts

July, 1806] NOMENCLATUKE. 13

376, 1895) that by elimination Gimiculus cauda longissima Brissou
[ = I)i})us (tiactaga Olivier; becomes the type. The name is thus unten-
able for any of the 2Iurlda\ although Lemmm lemmus is one of the spe-
cies included by Brisson in the genus.

GUs Erxleben, 1777 (Syst. Eegn. Anim., p. 358), contained marmota,
monaXj canadensis, tscherkessicus, zemnii, lemmus, migratorius, barahensis,
arenarius, lagurus, and ceconomicus [—Mus songarus Pall.]. Although
this genus contains two lemmings, the name need not be considered,
since it is preoccuijied by GUs Brisson, 1702.

Arctomys^ Schreber, 1780 (Plates to Schreber's Siiugth., CCVII-
CCIX, 1780), contained the following species: marmota, moiiax, hohac,
empetra, and citiUus. Of these the first four belong to the genus A rc/oMiys
as now understood, and the last to Spermoijliilus. The latter genus was
defined in 1823 by F. Cuvier (Dents des Mammiferes, 1823, 1G0-1G2,
255), who restricted the nauie Aretomys to the group to which it is now
applied. Aretomys Schreber is mentioned here only on account of:

Lagomys Storr, 1780 (Prodromus Methodi Mammalium, p. 39).
Although Storr and Schreber bear the same apparent date, it appears
safe to take Schreber as the earlier, since Storr alludes to the genus
Aretomys, aud refers directly to the ^^[us glareolus Schreberi,' a species
published at the same time.^ Storr evidently proposed Lagomys merely
as a substitute for Aretomys, a name which he considered inappropri-
ate, because the animals to which it was applied resemble hares rather
than bears.^ It is thus a synonym of Aretomys and requires no further
consideration.^

Myoeastor Kerr, 1792 (Animal Kingdom, 1, Mamm., Syst. Cat. Nos.
458-521), included the coypu and muskrat. Xo type was designated,
but subsequent elimination fixed the name on the coypu. (See i>. 14.)

Ondatra, Link, 1795, (Zool. Beytrage, Vol. I, Pt. II, p. 76), contained
the same species as Myoeastor Kerr, of which the name is thus a
synonym.

Lemmus Link, 1795 (Zool. Beytrage, Vol. I, Pt. II, p. 75), has escaped
the notice of recent writers. Vague references to it occur in works

'This name is apparently antedated by Marmota Blumenbach ("Haudb. d. Natur-
gesch., 1779," fide Agassiz). I have been unable to verify the reference, and do not
know what species were included by Blumenbach iu the genus.

-On the dates of the parts of Schreber's S.'iugthiere, see Sherborn, Proc. Zool. Soc,
London, 1891, 587.

■"Sequuntur in eundeni finem nomina specierum, laudato Pallas pariter ad mures
tractaruni, quae luihi genus constituerunt, Lagomys, nee Aretomys dictum, nam
Lepori aptius quam Urso, comparari jiosse videantur. Dicendie species nomini-
bus 111. Pallas ieque adhibitis, ha>c suut; J/, arenarius, M. songarus, AT. furaii-
culus, J/, cricitus, M. accedula, M. phwiis, 21. lagurus, 21. gregaVis, 21. socialis, 21.
oeconomus, 2f. rutilus, 21. glareolus Schreberi, J/, monax, 21. marmofa, 21. empetra, 21.
arctovvjs, J/, citillus, 21. lemmus, 21. torquatvs, 2T. hudsonius, 21. taljnnus, 2f. capensis,
21. aspalax, 21. typclus" (sic).

^Lagomys Storr of course antedates Lagomys Cuvier, 1800, the current name for the
pikas.

14 NORTH AMERICAN FAUNA. [No. 12.

of the early part of the preseut century, but of late all traces have dis-
appeared, Lataste (Le Naturaliste, Tome II, p. 47.'3, 1882). after a long
and fruitless search, concluded tliat the name had probably never been
published, and that the references of the older authors were merely to
Link's manuscript. Mr, Oldfield Thomas has discovered Link's book
and linds that the genus Lemmus contained the species socialis, hif/urus,
Jemmus, torqi<ati(s, f/lareolxs, and hndso^iiKS,^ representing the modern
genera Lemrnvs, Dk-rosiony.i\ Microfns, and Evotnmys. As the name
Lemmus has been restricted by subsequent authors to the species
lemmus and its near allies, a group to -which no other generic name has
been specially applied, it must be retained in this sense,-

Microtus Schrank, 1798 (Fauna Boica, p. 72), included M. terrcstris,
M. amphihius [=M. terrestris Linn.), and ilf. '(/regarius.'' The Microtus
terrestris of Schrank is not the Mus terrestris of Linnanis, but the com-
mon field mouse of Central Europe, Microtus arralis (Pallas). J/, gre-
gurius Schrank, apparently based en one specimen from Bettbrnnn, is
probably a young 21. arralis. The third species, M. amphihius, is the
water rat, Microtus terrestris (Linnaeus). Thus the genus Microtus
originally contained two species, arralis and terrcstris. As the latter
was made the type of Arvicola by Lacepede in 1801. arralis nmst be
taken as the type of Microtus.

Fiber Cuvier, described in 1 798 but not named until 1800 (Tabl. f^lem.
de I'Hist. ^i\t. d. Anim, 111, 1798; Lecons d'Anat. Comp. I, Tabl. I,
1800), is the first and only generic name based exclusively on the musk-
rat, Cuvier, in establishing this genus, eliminated Fiber zibethicus from
Myocastor, and thus fixed the latter name on ^1/. coypu. (See page l-").)

Arvicola Lacepede, 1801 (Mem. de I'lnst., III., Paris, 1801, 489-^), was
based on Arvicola amphihius { = 2[us terrestris Linn.) alone, and not on
the European voles in general, as often supposed.^ Although the name
Arvicola can not be used in a generic sense, it is available fur the sub-
genus of which Microtus terrestris is the type.

Hypurhcus Illiger, 1811 (Prodr. Syst. Mamm. et Avium, ]i. 87), con-
tained the species lemmus. ampliibius {=terrestris), and arr((lis, or the
modern genera Lemmus and Microtus. As no type was designated, and

' Mr. Thomas has kiudly seut me a co^jy of the original diagnosis. It is as follows:
"Gen. 8 Lemmus, Lemming. Die Thiere dieses Gesr-hlechts kommen mit den vorigen
[Mils'] sehr ueberein, aber die Ohren sind viel kleiner nnd abgerundet, der Korper*
gedrungener, die Beiue verhiiltnissmiissig kiirzer, der Schwanz sehr kurz. AucBj-
weichen sie in der Lebensart von den vorigen ab. Sie niiheru sich Arctomys. Hieher
gehitren: Mus socialis, Ingurus, lemmus, torqudfus, (jlareoJus, liudsoiiius."

-See note on the names llraclnjurus, Mi/odes, Hijpudaus, and Lemmus, in Actes de la
Societc Scientitiqne du Chili, Tome V, iip. XX. XXI, 1895.

3 This is sometimes quoted: "Tableau des divisions, etc., de la class des mamnii-
feres, 1799." The paper was "lu le 21 prairial an. 7,'' though not published untill801.

■•Lacepi'de's description is as follows: "44 Cnmpagnol. Deux iucisives superieurs
non comprimees; deux incisives inferieurs tranchantes; molaires sillonn<^es; jjoint
d'abajoues; queue velue. Camjiagnol aciuatiqae — Arvicola amiihibius."

JcL\M896.] NOMENCLATURE. 15

as both Lemmus and Microtus were included in the then undivided
geuus Ijcmmn.s Liuk, the uame Hypmhvus juust lapse into sjniouymy.

Myodcs Pallas, 1811 (Zoog. Eosso -As., I, p, 172), embraced ten spe-
cies, now placed in four geiiera. The species are: Lemmus, forquatvs,
lagums, a^conomns, arvaJis, saxatUi.s, gra/aUs, sociali.s, alliarivs, and
rntiJus: the genera : Lemmus {lemmus), Dicrosfonyx {torquatus), Microtus
{a'cono7nus, arralis, saxatiUs, grecjaUs, sociaUs, alUarius, Jagurus), and
Evotomys ( rut this). Since Myodes contained species of exactly the same
modern genera as Lemmus Link and no groups not included in the lat-
ter, the name is a synonym of Lemmus.

Braehyiirus Fischer, ISlo (Zoognosia, I, 3d ed , pp. 14, iil; JU^ 1814, p.
5.5), contained the species: arralis, rutilus, am2)hihius, lemmus, torquatus,
alUarius, hJumenbachii, fulvus Geoffroy, niloticus Geoffroy: also the
' species dubia- ' : zemni, gregar'ms, sociaUs, lagurus, wcouomus. The uame
is a pure synonym of Lemmus Link, uidess it may be applied to some of
the exotic or dubious species.'

Ahnceola Blainville, 1817 (Xouv. Bict, d'Hist. Xat., IX, p. 287), i:)ro-
posed for ' le Genre Campaguol ' is probably an erratic misprint for
Arvicola. Xo type is mentioned.

Mynomes Rafinesque, 1817 (American Monthly ^Magazine, II, p. 15)
was based on Wilson's figure of the common meadow mouse of the east-
ern United States. The name is thus a synonym of Microtus Schrank
as Microtus arralis and M. 2)cnnsylranicus can not be separated sub-
generically.

Psammomys LeConte, 1830 (Ann. Lye. Xat. Hist., :N". Y., Ill, p. 132)
is tlie first name proposed for the subgenus containing Microtus pine-
tor nm. It is, however, preoccupied by Psammomys Cretzschmar, 1828
(Atlas zu der Eeise im Xordl. Afrika. Iste Abtli., Zool. (182G), Heft XI
1828, p. 50. Type Psammomys ohesus Cretzschmar) and so can not be
used here. The date of Psammomys LeConte is usually quoted as 1829,
but the paper on this genus, although read on December 21, 1829, was
probably not published until after the end of January, 1830, since papers
read January 11-25, 1830, are included with it in one signature.

Pitymys McMurtrie and Ammomys Bonaparte both appeared in 1831.

McMurtrie (American ed. Cuvier's Eegne Animal, I, p, 434) pointed
out that Psamiuomys LeConte is preoccupied, and for this name substi-
• tuted Pitymys. Bonaparte (Saggio Distrib. Metod. degli Anim. Vert.,
p. 20, footnote) after showing that LeConte's name Psammomys is not
tenable, proposed to change it to Ammomys, thus preserving the original
meaning of the word.^ It is impossible to tell which name is the earlier,

j 'This name has been supposed to be preoccupied bv ISrachi/unis Spix (Lataste
i Ann.Mus.CiY.St.Nat. diGenova, XX, p. 264; Biiclmer, Wissenscb. Result, der von
j N. M. Przewalski unternomm. Reisen, I, p. 127). Spix's uame, however, dates from

1823 and woukl in no way invalidate Jirachiptrus Fischer, were the latter on other

grounds tenable.

Preudiamo la liberta d' iutrodurre una piccola mutazioue ortografica uel noma
dato al nuovo genere dal 8ig. LeConte, la quale nou ne canibia per., il siguificato."

1() NORTH AMERICAN FAUNA. [X0.12.

but in the imcertainty Pitymys slioiikl be retaiued as tlie one adopted
by all subsequent writers.

Cuniculus Wagier, 1830 (Nat. Syst. d. Anipliibien, p. 31), included
three species {G. lemnius, C. torquatus, aud ('. <(.sp((Iax) now referred to
three genera and two subfamilies. The name has been commonly
applied to torquatus and its congeners, but its use is invalidated by
Cuniculus Brisson, published fifty-eight years before.

Hemiotomys DeSelys-Longchamiis, 1830 (Essai monograph, sur les
Campagnols des environs de Liege, p. 7), was i)roposed as a section of
Arvicola (=Microtus) to include the species fulvus {=arvalis) and
ainpMhius { = terrestris). As each of these had already received a ten-
able subgeneric name, Hemiotomys lapses into synonymy.

Pinemys Lesson, 1830 (Hist. i!^at. d. Mamm. et Ois. decouv. depuis
1788, Compl. ffiuvres de Buffon, V, p. 130), based on Psammomys pine-
torum LeConte, is a synonym of Pitymys McMurtrie.

Lagurus Gloger, 1811 (Gemeinniit/. Hand- u. Hilfsbuch d. Natuige-
schichte, 1, pp. XXXI, 97), is the earliest available name for the sub-
genus of which Mus lagurus Pallas is the type.^ (See footnote, p. 1!>.)

Dicrostonyx Gloger, 1811 (1. c, pp. XXXI, 97), is the tenable name
for the genus usually known as Cuniculus Wagler.^ This name has
escaped notice until very recently.^

Neodon Hodgson, 1819 (Ann. & Mag. Xat. Hist., 2d ser., Ill, p. 203),
is a synonym of Micnrtus, as its type, N. .siTclci^nensis Hodgson, can not
be separated subgenerically from Microtus arvalis.

'■'■ Myolemmus Pomel, 1851 (Ann. Sci. Soc. Auv'ergne),-' is a synonym
of Dicrostonyx Gloger. This statement is made on the authority of
Trouessart (Cat. Mamm. viv. et foss., Rodentia, Ft. II, p. 150, 1881), as
I have had no opportunity to verify the reference.

Misothermus Hensel, 1855 (Zeitschr. der Deutsch. geolog. Gesellsch.,
VII, p. 492), is stated by the author to be based on Myodcs torquatus
Pall. It is thus antedated by Myolemmus Pomel and Dicrostonyx
Gloger.

Pedomys, CMlotus, and Synaptomys are three names proposed by
Baird in 1857 (Mamm. X. Am., pp. 510, 517, 558). All are tenable for
the groups to which they were applied. Pedomys and Chilotus are sub-
genera of Microtus. Their types are Microtus austcrus and 2[. orcgonus,
respectively. ^Synaptomys is a genus, with aS'. cooper! as the type.

'Gloger'e description is as follows: " Theils aaf dem Ural und anderen Gebirgen,
tlieils audi in tieferen Gegeudeu Sibiriens, giebt es, drei oder vier andere Artcn niit
kleiueu, rundlicbeu oder spitzigen Daumniigelu uud von einfacherer F.'irbnng
(Laf/urus), die ziim Tbeile nicbt weuiger zum Wauderu geneigt scheinen. Z. B. L.
migratorhts/'

•Gloger says: "Von den uordamerikaniscben Lemiuiugeu zeicbuen sicb niauclie
durch eiu Paar hocbst sonderbare (gleicbsam doppelte) Vorderkralleu aiis, die 2 od»T
gar 3 Spitzeu iiber einander zu habeu sebeinen, weil sie uuter den Nageln grosse,
barte Ballenbervorragungen besitzen. Sie kuunen daber Gabelkraller (Dicrostonyx)
beissen."

^For a paper on Gloger's generic names for maunuals, see Tbomas, Ann. A. Mag-
Nat. Hist., 6tb ser., XV, Feb. 1, 1895.

July, 1890.] NOMENCLATURE. 17

Falu(Ucola Bhisius, 1857 (Fauna der Wirbelth. Deutscbl., Bd. I,
Siiugetbiere, p. 333), a subgenus o£ Arricola { = 2Ucrotus), contained tbe
species: amphibii(s { — terrestr is), nivalis, and ratticeps. As tbe first is
a member of tbe subgenus ArincoJa and tbe otbers eacb a true Micro-
tiis, tbe name can not be used. Moreover, it is preoccupied by PalKdi-
co/« Wagler, 1830 (Xat. Syst. d. Ampbibien, p. 200, type Bufo alhifrons
Spix).

Agricola Bbisius, 1857 (L c, p. 331), was proposed as a subgeneric
name for 2[icrotus (ig rest is. Tbe differences between tbis sjiecies and
the albes of M. arvaJis are too sligbt to entitle the groups to rank as
distinct subgenera; but assuming that it Mere desirable to separate
them tbe name Agricola would be antedated by Mynomes Kafinesque,
1817, based on 2Fivrotns pcnnsylninicus, a ibrm whose supersi)ecific
characters are exactly similar to those of M. agrestis.

Phaiomys Blyth, 1803 (Journ. Asiat. Soc. Bengal, XXXII, p. 89), is
the first and only tenable name proposed for tbe subgenus having
Microtiis hJythi as the type.

Ochetomys Fitzinger, 1807 (Sitzungsb. K. Akad. Wiss. Wien, LVI,
June, 1807, p. 47), included the water rats of Euro[)e. It is thus
equivalent to ArvicoJa Lacepede.

Fraticola Fatio, 1807 (Les Campagnols du Bassin du Leman, p. 30),
is a subgenus of Arvicola {=jMicrofus) containing: amphihius { = terres-
tris), nivalis, arvalis, ratticeps, and campestris {=arvalisF). As all of
these are species either of AHcrottis Schrank, or Arvicola Lacepede,
the name Fraticola can not stand. Fraticola is, moreover, preoccupied
in ornithology.

Sylvicola Yiitio, 1807 (1. c, p. 03), based on Microtus agrestis is exactly
equivalent to Agricola Blasius, 1857. The name is preoccupied in ornith-
ology, entomology, and conchology.

Terricola Fatio, 1807 (1. c, p. 73), contained Microtus suhterraneus and
M. savli. The name is, however, preoccupied in conchology by Terricola
Fleming, 1828.

Isodelta and Anaptogonia Cope, 1873 (Proc. Am. Philos. Soc, XII, p.
87), are the tenable names for two extinct subgenera found iu the Post-
pliocene cave deposits of Pennsylvania. Their types are Microtus
. speothen and M. hiatidens, respectively.

Evotomys Coues, 1874 (Proc. Acad. Xat. Sci. Phila., p. 180), is the
tenable name for the genus of which Mus rutilas is the type.

Micrurns Forsyth Major, 1870 (Atti della Societa Toscana di Sci.
Natural!, Ill, fasc. I, p. 120), founded on Mina Palumbo's description of
Arvicola nehrodensis (a Fityniys), is preoccupied by Micrnra Ehrenberg,
1831, a genus of Vermes.

Alticola Blanford, 1881 (Journ. Asiat. Soc. Bengal, L, pt. 2, jj. 93), is
the only name proposed for the Asiatic subgenus with Microtus stolicz-
' kanus as type.

! Eremiomys and Borioilon Polyakoff, 1881 (Mem. Acad. Imp. Sci. St.
10933— Xo. 12 2

18 NORTH AMERICAN FAUNA. [No. 12.

Peterribourg, XXXIX su})pl., p. 34), based, respectively, on Mns lagurus
Pallas and Mus torquatns Pallas, are synonyms of Lai/urus Gloger and
Dicrostonyx Gloger.

Keofiher True, 1884 (Science, IV, p. 34), was described as a genus
with iV". alloii, the only known species, as type. Recently it has been
shown that the characters of the animal are not enough to separate it
generically from Microtu.s, of which, however, Neofihcr forms a well-
marked subgenus.^

LasioiJodomys Lataste, 1887 (Annali del Mus. Civ. di Storia iSTatuvale
di Genova, ser. 2a, Vol. IV, p. 208), is a synonym of Phaiomys Blyth,
1803, the species on which the two names were based, Microtus hrandtl
Radde and Microtus hJythl Blanford ( = .!/. leucnrus Blyth nee Arvicola
leucuriis Gerbe), respectively, being in no way separable subgenerically.^

Phenacomys Merriam, 1889 (Xorth Am. Fauna, No. 2, p. 28), is the
tenable name for the genus of which Phenacomys intermedius is the type.

CampicoJa Schulze, 1890 (Schriften Xaturwiss. Vereins d. llarzes in
Wernigerode, V, p. 24), is a subgenus formed for the reception of the
species Microtus arralis, M. suhterrancus, and 31. camjycstris. It is
thus a compound of two subgenera, Microtus {ari-aUs and campestris)
and Pitymys {suhtcrraneus), each of which has previously received a
teuable name. Campicola is, moreover, preoccupied in ornithology
(Swainson, 1827).

Bramns Pomel, 1892 (Comptes Eeudus, Paris, CXIV, p. 1159), is
based on a mandible and the teeth of both jaws of a rodent from the
Quaternary phosphorites of Trara de Xedroma near Ain-Mefta, Tunis.
Although the author compares this fossil with the bones and teeth of
the water rat, he points out such striking differences between the two
that it is very doubtful whether Bramus can be considered a member
of the subfamily Microtincc. (See p. 73.)

Aulacomys Eboads, 1894 (American Naturalist, XXVIII, p. 182),
although based on an abnormal specimen, is the tenable name for a
group of American water rats, should tbe latter be considered sub-
generically distinct from Arvicola. The peculiarities of the original
si>ecimen of Microtus arvicoloides, the type of Atilacomys, are such that
the group was originally given full generic rank.

Mictomys True, 1894 (Proc. U. S. Xat. Museum, XVII, Xo. 999,
p. 242, Advance Sheet, April 20), was proposed as a full genus with
Mictomys innuitus True for tlie tyi)e and only known species. The name
is tenable, but the group is only a subgenus of 8yna})tomys.'^

Tetramerodon Rhoads, 1894 (Proc. Acad. Xat. Sci. Phila., p. 282), is
the most recent synonym of Microtus. The author, as Blasius had

iTrue, Report of the Smithsoniaa Institution for 1884, Part II, pp. 325-330. PI. II.
Merriam, North American Fauua, No. 5, p. 60, 1890. Chapman, Bull. Am. Mns. Nat.
Hist., New York, VI, p. 334, 1894.

2 See Actes de la Societe Scientifique du Chili, IV, p. CLXXXYIII, 1894.

3 See Merriam, Proc. Biol. Soc. Washington, X, p. .57, 1896.

July, 1896.]

HISTOKY OF CLASSIFICATIONS.

19

aheatly done nearly forty years before, divides the subgenus Arvicola
( = Mierotus) into two groups, based on tlie structure of the middle
upper molar. To the species with this tooth formed of five prisms he
restricts the name Mynomes, while to those with the same tooth made
up of only four prisms he applies the new name Tetramerodon. The
character in question is far too trivial to serve alone as the basis for a
subgenus. If, however, the advisability of subdividing the genus along
such narrow lines be admitted, the name Tetramerodon still becomes a
synonym of Microtus, since ^[. arvalis, the type of the latter, is itself
a species with the middle upper molar four parted.

HISTORY OF FORMER CLASSIFICATIONS.

The most important studies of the various groups of Microtina', but
more especially of the subgenera of Microtis, are those of De Selj^s
Lougchamps (183G to 1862), Blasius (1857), Baird (1857), Fatio (1867),
Coues (1874), Blanford (1881), and Lataste (1887). The names used by
these authors for the subdivisions of Microius adopted iu the present
classification are shown in the accompanying table:

Table of Xames used hij ^Infhors for the Suhijenera of Microtus.

Names used I De Selys | jjlasius,
111 the i)res- i Longchamps, ,0='-
ent paper. I 1836 to 1862. | ^''^'•

Baii'il.

1857.

Arvicola Memiotomys Palitdicola

I A rvicola Arvicola

Micrntus '\ i (part).

My names .. Agricola ..

Pltymys

Microtus .. .. Arvicola
I (part).

Pedo)iiys. .
Phaioinys.

Fatio, Coues,

1867. ! 1874.

Praticola
(part).

j Praticola

(part).
Hemioto- \ Sylvicola..
inys.

Blaiiforil,
1881.

Myonovies Xeodon,
(part).

Pitymys .. Terricola.A Pityviys

Pedomys.

Cliilutiis I ' Chilotus

Lagnrvs '

A Iticola

Ilyiieracrius ' '

Eofhenomyx.

Antelioinyi:
y>'0 fiber

Pedomys.

Paludicola

ChilotKS

Lataste,
1887.

Arvicola.

2Iicrotus.
Pitymys.

Lasiopodo-
mys.

Allicola . ..
A Iticola

part .-.
X e 0 don

(part).

De Selys Longchamps imblished two extended papers on the Euro-
pean Microtinw, and later a note supplementary to the first of these.
The first paper appeared in 1836 under the title 'Essai Monographique
sur les Campagnols des environs de Liege.' In this the author showed
that hitherto the voles had been divided into two groups, according
to their habits, the aquatic species being separated from those that are

20 NORTH AMERICAN FAUNA. [Ko. 12.

strictly terrestial. This proved nusatisfactory because tlie two were
found to intergrade imperceptibly. Heuce he proposed to rearrauge
tlie si)ecies according to the length of the ears. The first division, or
that in which the ears are extremely short or apparently absent, he
named Hemiotomijs, This the author subdivided into two sections,
neither of wliich he named. The first contained one species, Arvicola
fulvus {^=Microtns arvalis), distinguished by its short tail and by the
supposed absence of external ears. The second contained the water
rat. To Arvicola {=Microtus) proper were referred the three species,
arvalisj suhferraneus, and rufescens { = Ei'otomys glareolns). Six years
later, in liis Etudes de Micromammalogie, De Selys Longchamps fol-
lowed the same system of classification, but considerably extended it
and included species from Asia and iSTortli America. This later scheme
is as follows:

The genus is first divided into two sections, one of which consists of
species with ears shorter than the fur and with very small eyes, the
other of species with the ears as long as tlie fur and with tlie eyes well
developed. The first section contains two groups, (1) Hcmioiomys with
the European water rats and the American Arvicola riparius {=Microtus
pemisylvanicus), and (2) Microtus with the species/wZyws, savU, ceconomus,
and certain American forms not mentioned by name. The second sec-
tion is divided into three groups: (1) Arvicola with the species suhter-
ranens, arvalis, gregalis, alliarius, duodecimcostatus, and socialis; (2)
2[yodes with the two species ruhidus { = Evoiomys (/lareolus) and rntilits
[z=Evotomys rutilus) ; (3) Mynomes with the species 2}ratrnsis {=:Microti(s
pennsylimnicus). These groups and sections the author considers in no
way entitled to rank as genera or subgenera. He names them merely
for convenience. ^ In a postscript published at the time of distribution
of the last copies of the Essai Monographique, twenty-six years after
its appearance, the author makes a few corrections in the classification
previously adopted. He points out that his Arvicola fulvus is merely
a young specimen of ^rl. arvalis that by accident had lost its external
ears, and, furthermore, that the species suhterraucus should be trans-
ferred to the section Microtus.

The classification as finally perfected is as follows:

Genus Arvicola:

Group Hemiotoimjs (water rats).
Group Microtus {suhierranens and savii).
Group Arvicola (typical voles).
Group Mrjodes (glareolus).
Group Mynomes (2)eiiusijlca)ticiis).

' Je dois pr6venir que je m'opposerais entierement a relcvation d'aueune de ces
sections au raug de genre ou de sous-genre. Toutes passent de I'une a I'autre par
des nuances inseusibles dans la longueur de la queue et des oreilles; et, quant au
caractere tiro de la racine des dents, il est probable qu'il existe a uu degrd plus ou
moins fort chez d'autres espoces. Si je me suis perniis d'imposer a ces gioupes des
nonis latins pris parmi les synonvmes du genre, ce n'est nullement pour qu'ils puissent
etre iutroduits dans la nomenclature binaire, mais pour donner anx strangers I'id^e
des divers noms que j'ai employes en franvais. (Micromammalogie, p. 87.)

JrLY,lS9G.] HISTORY OF CLASSIFICATIONS. 21

The groups Hemiotomys, 3licroti(S, aiul Arvicola of De Si'lys Loug-
cliainps are exactly ecjuivalent respectively to tiie subgenera Arvicohi,
ritymi/s, aud Microtus of the present j)aper, while Myodes is the same
as the genus Evofomys. The grouj) Mynomes based on Eafiugsque's
description of Mijiiovics 2}>'f<t('>'''^is { = Microfiis peuHsylvcmicus) sliould be
united with Arricola (Microtus, as now understood), a course whicli the
author no doubt would have followed had he been acquainted with the
type species,

Blasius published in 1857, in his 'Fauna der AVirbelthiere Deutsche
lands,' a classitication of the voles based primarily on the pattern of
enamel folding in the first and second molars of the lower jaw and the
second molar of the upper jaw. This system differs in many ways from
that of De Selys Lougchamps, and is as follows:

Genus Arvicola:

Subgeims Hypudcvus {{/larcoJiis).

Subgenus raludicola {amphihius l—terreNfris'\, iiivali'^, ratliceps)

Subgenus Agricohi {agrestis).

Subgenus Arvicola :

A. Arvicola (campestris, arvalis).

B. Microtus De Selys j)art (subterraneiis, savii).

The subgenus Arvicola Blasius subdivides into two sections, A.
Arvicola and B. Microtus De Selys (part). The former includes the
species campestris and arvalis, the latter suhterraneus and savii. The
subgenus Hypudwus and the section Microtus are equivalent, resj^ec-
tively, to the genus Evotomys and the subgenus Titymys of the present
paper. Of the other groups, the restricted Arvicola contains the tyi)ical
species of the subgenus Microtus, Agricolay a slightly aberrant form of
the same, and Paludicola, the subgenus Arvicola and two aberrant
members of the subgenus Microtus. Blasius's subgenera Faludicola and
Arvicola are excellent illustrations of the unnatural results of a system
of classification based on one set of characters. While there is a general
similarity between the enamel pattern of the three species associated in
the former, Microtus tcrrestris differs from M, rattice^JS and j\[. nivalis
in the form of the skull, the number of plantar tubercles, the quality of
the fur, and in the presence of large musk glands on the sides. In the
subgenus Arricola Blasius associates two of the most distinct subgenera
of the genus Microtus {Microtus and Fitymys), and treats the differences
in the number of mamma' and footpads, form of skull, and size of eyes
as matters of trifiing importance in comparison with the general simi-
larity of the enamel pattern. On the other hand, the author recognizes
Agricola as a full subgenus, when the chief character on which the group
is based is the presence of a minute supplemental postero-internal prism
on the middle upper molar.

The classification adopted by Baird (Mamm, K. Am., 1857) is based
OH a combination of characters, and is thus much more satisfactory
than the artificial arrangement published almost simultaneously by

22 NORTH AMERICAN FAUNA. [No. 12.

Blasius. His classification of tlie subdivisious of Microtus is as
follows:

GenusylrricoZrt ;

Subgenus Hiipnilcvus (t/appcri).
Subgeiins Arricola (typical voles).

Section Memiotomys {most of the American species and the Euroj)can

ag7-esiis).
Section Chilotus (oregoni).
Section Pedomijs (aimterus).
Section Pifymys (piuetonim).

Baird's subgenera Hyjjudwus and Arricola are equivalent to the
genera Eroiomys and Microtus of tlie present paper, while his sections
Chilotus, Pedomys, and Pityniys are equal to the subgenera of the same
names. The section Ilemiotomys of Baird is the Arvicola of De Selys
Lougcliamps, and the subgenus Microtus of the present paper.

In 1867 ratio published a classification of the European voles in a
paper entitled 'Los Cam])agiiols du ]>assin du Leman.' This arrange-
ment is essentially the same as that of Blasius. Fatio, however, recog-
nizes Hyimdcvus {■=Erotomys) as a full genus, and raises the second of
Blasius's two sections of the subgenus Arricola to the rank of a sub-
genus, while the first he unites with Microtus terrestris, M. nivalis,
and M. ratticeps to form the subgenus J*raticola. He also arbitrarily
changes the names of certain groups. His classification is as follows:

Genus Hypudtvus (ylareolus).
Genus Arvicohi.

Subgenus rraticoJa (' nmjiltihins,' nivalis, arralis, ratticeps, campesiris).

Subgenus Sylricola {ayrestiis).

Subgenus Tcrricola {siihterrantus, savii).

The subgenus Tcrricola and the genus Hypudcvus are equal, respec-
tively, to the subgenus Vitymys and the genus Evotomys of the present
paper. The subgenus Sylricola is equivalent to the subgenus yl^r/co/fl
of Blasius, like it containing the x)entamerodont species of the subgenus
Microtus. The subgenus I'raticola includes the type species of both
Arvicola and ^licrotns, together with three other tetramerodont species
of the latter.

In 1874 Dr. Coues published, in the Proceedings of the Academy of
Natural Sciences of Philadelphia, an abstract of his monograph of
the North American Murida^, which appeared in full in Volume XI
of the Eeport of the United States Geological Survey of the Terri-
tories (Monographs of North American Eodentia). Here he presented
a classification of the xVmerioan Microtinw based primarily on Baird's
review of the group. The differences between the arrangements adopted
by Baird and Coues are so slight that a few words only are necessary
in regard to the latter. Dr. Coues recognizes the red-backed mice as a
distinct genus, which he calls Erotomys, after showing that the name
Hypuda'us generally used for the group is untenable. The subgenera
of Jlicrotns adopted by Dr. Coues are exactly equivalent to Baird's

JuLv,iso6.] HISTORY OF CLASSIFICATIONS. 23

sections of liis typical subgeuus Arvicola. Dr. Cones points out
Baird's eri-or in the application of tbe name Eeiniotomys De Sclys
Longcliamps, and substitutes for tbe latter tbe equally untenable
Mynomes Eafinesque.

In 1881 Bbmford proposed, in tbe Journal of tbe Asiatic Society of
Ben,£;al (Vol. L, Pt. II, pp. 88-117), a classification of tbe voles of tbe
Hiiiiidayas, Tibet, and Afghanistan. Tbe species occurring- in tbis
region be arranges in three sections, thus:^

Genus Arvicola:

Section I'ah((HcoIa, {hhjilii, mandrianus).

Section AUicola (sloliczkaniis, strachci/i, roylei, hlanfordi, ici/nnei}.

Section Xeodou {.nkkimensis, mdanoyaster).

Blanford's 'sections' P<//Hf?/cr)/« and Xeodou are excellent instances
of unnatural classifications based on single characters. Microtus hlythi
and ^r. mandrianus are species of Fhaiomys, a subgenus which difters
from the water rats or from Microtus (Microtns) nivalis and 31. (M.) rat-
ticeps (all of which were included b3' Blasius in Paludicola) in many
important characters. Because there is a general likeness in the pat-
tern of enamel folding they are united under one superspecific name.
Again, Blanford places in the section Neodon the species Microtus siJxl-i-
>«e»i/.s', which is a slightly abnormal member of tbe subgenus J/ /cro/»s,
and Microtus mchinogastcr, a species with the bony j^alate formed exactly
as in the red-backed mice {Evotomys). These members of widely dif-
ferent groups are brought together on account of a very superficial
likeness in enamel pattern. Blanford's section Alticola is probably
equal to the subgenera Alticola and Ryperacrius of the present paper,
thongh it is still a matter of doubt whether it actually included any
members of the latter.

The most recent classification of the subgenera of Microtus is that
proposed by Lataste. Tbis author has published two important papers
on the subject, the first in Le Xaturaliste (Tome II, pp. 323, 324, 332-
334, 342, 343, 347-349, 1883), and the second in the Annali del Museo
Civico di Storia Xaturale di Genova fSerie 2a, Vol. IV, pp. 259-274,
1887). While recognizing the unsatisfactory nature of the artificial
classification adoi)ted by Blasius, Lataste subdivides the voles in
accordance with a system fully as arbitrary as that followed by any of
his predecessors. According to Lataste the characters derived from
tbe teeth of the voles are of no value except in distinguishing between
genera.2 Tbe subgenera he arranges according to tbe number of maiu-

' Ijlanford adopted Biasing's classification of the voles at large (pp. 91, 92). Except
in the case of Paludicola, however, he supposed that none of the European sec-
tions of the genus Microtus are represented in the region with which he deals.

""Chez les Eongeurs du uioins, sinon chez tons les Mammiferes, les characteres do
la denture me serablent d'ordre gcnerique quand ils sont sxiffisamnient nets ct
tranches, mais sans aucune importance taxouoini(£ue quand ils sont aussi mininies
que ceux que I'on invoque d'ordinaire, a la suite de Blasius, chez les Campagnols, et

24 NORTH AMERICAN FAUNA. [No. 12.

rnip and plantar tiibeicles. Altbougli tliis system leads to a tolerably
satisfactory arrangement of tlie European voles, it can not be applied
to the genns at large, since it would unite sucli distinct groups as
Arvieohi and Chilotus, or JS/'eoJiher and Piiymys. Lataste's classification
is as follows :

Genus Alicrotus: '

Siibgeuus Myodes {rtitilus, glareohis).

Subgenus Microtus (grefjalis, arvaJis, afffesiis, ratiiceps, pennsiilvanicus, nhalia)-

Subgenus Arvieohi (ierrestris, mnsigiiani).

Subgenus Pityniys (p'nietorum suliterrmieiis, socialis, middcndorffi).

Subgenus Lasiopodomys (hrnndti).

The subgenera Myodes and Lasiopodomys are equal, respectively, to
tlie genus Evotomys and the subgenus Fhaiomys of the present paper.
The subgenera Microtus and Arvieohi coincide with groups here recog-
nized under the same names, while the subgenus Pifymys is essentially
the same as that defined on page 58. Lataste, however, includes in
ritymys the species middeiidorffii, which is probably not a member of
that group as now understood.

CHARACTERS ON WHICH THE PRESENT CLASSIFICATION OF THE
SUBGENERA OF MICROTUS IS BASED.

In the discussion of the systems of classification hitherto adopted,
the impracticability of subdividing the genus Microtus according to
the variations in any one set of characters has been shown. The
highly artificial systems of Blasius and Lataste give the best examples
of the unnatural results to which any such course must inevitably lead.
In the present paper the classification used is based on an assemblage
of characters. The more important of these, or the ones least adapted
to the special needs of the different animals, and hence least likely to
vary, are: Form of skull, structure of bony palate, pattern of enamel
folding, number of mamma*, number of plantar tubercles, and presence
or absence of musk glands on the sides. Characters of less importance,
because more readily modified to fit a species to the special requirements
of its environment, and hence more unstable, are: Quality of fur, hair-
iness of soles, length of tail, form of front feet, size of eyes, and form
of external ear. It is only through careful consideration of all these
that a satisfactory arrangement of the species can be obtained.

Nearly all of the characters now used have been recognized in classi-
fications already proposed. In every case, however, they have been
assigned degrees of importance different from those which they now
receive. To take the three most consi^icuous examples: De Selys
Loiigchamps arranged the voles with regard to their external form;

qui portent sur les extr^mit^s mal definies et 6mineniment variables, soit j^osterieur
d(i la derniore molaire snperieure, eoit anterieur de la prcmii-re molaire inforieure."
(Ann. del Mus. Civ. di Genova, Ser. 2a, Vol. IV, p. 260 footnote.)
Compare with this the oiiinion expressed by Biichner. (See footnote, p. 25.)
' To Lataste is due the credit of recognizing tlie true status of the name MkroUia.

,u-LY,l896.] CHARACTERS OF ENAMEL PATTERN. 25

Blasins based liis classification on the pattern of enamel folding ^itli-
ont regard to external characters, and Lataste snbdivided the group
according to the numbers of mammte and plantar tubercles, disregard-
ing everj^thing else. The impossibility of reaching satisfactory results
by any of these methods has been pointed out by Biichuer, who, how-
ever, takes an equally extreme position in his reluctance in any way to
subdivide the genus Microtus.

Biichuer was first to recognize the important fact that tlie enamel
pattern, while variable within certain limits and hence of little value
taken by itself, is nevertheless of considerable systematic importance
when considered in connection with other characters.'

In about 75 per cent of the specimens of a given species the enamel
pattern conforms to a type which maybe considered normal.^ Among
the abnormal specimens constituting the remainder, the variation, how-
ever, is very considerable. In the accompanying illustrations (figs. 1,
2, 3, 4, o, and 6) are shown some of the conspicuous aberrations in the
form of the teeth of Microtus pennsylvanicus.^ In the descriptions which
follow the normal enamel pattern is alone considered.

1 After meutiouing Lataste's view (see footnote, p. 23), Biichner says: "Meiner
Ausiclit uach liefert im Gegentheil der Ban der Backenziihue, obwolil derselbe
zn weileu aucli im Bereiche einer Art leicbt variirt, eiu vorziiglielies Merkmal,. welches
allein geuommen fiir die Charalvteristik eiuer Art nicht geniigt, iu Yerljiudung aber
niit (leu iibrigen Merkiualen sebr grosse Dieuste leistet und von bedentendem sys-
tciuatiscben Wertbe ist." (Wissenschaftlicbe Resultate der von N. M. Przewalski
nach Ceutral-Asien nuternommenen Reisen. Zool. Theil, Bd. I, Siiugethiere, Lief. 3,
1889, p. 97.)

-Among 285 specimens of Microtus pennsylvanicus 71, or 21.9 per cent have tbe
enamel pattern in some way abnormal. Of these, 26, or 9.1 per cent, have the first
outer triangle in ia^3 communicating more or less freely with the inner triangle
(fig. 3); one has the second outer triangle opening into the posterior loop (fig. 3);
two have the posterior loop of very unusual shape (iig. 3) ; one has a second inner
closed triangle iu in3 (fig. 3), and 14, or 4.9 per cent, show a distinct fourth salient
angle on the outer side of the same tooth. In the first lower molar 24, or 8.3 per
cent, liave 6 closed triangles (fig. 4), one has only 3, still another has 7 (fig.4),
while in 5, or 1.7per cent, there are 4 (fig.4). Of these 28.5 specimens m 3 is abnormal
in 44 cases, or 15.4 per cent, iiTl in 31 cases, or 10.8 per cent. C4rouping the abnor-
mnlities according to their frequency, they may be arranged as follows:

m_3 has first outer triangle open in 26 cases, or 9.1 per cent.

m 1 has one additional triangle in 24 cases, or 8.3 per cent.

in 3 has an additional salient angle on the outer side in 14 cases, or 4 per cent.

Ill 1 has one less triangle than usual in 5 cases, or 1.7 per cent.

Ill 3 has the posterior loop of very unusual shape in 2 cases, or 0.7 per cent.

m 3 has the second outer triangle abnormal in 1 case, or 0.35 per cent.

Ill 3 has an additional inner triangle in 1 case, or 0.35 2)er cent.

iiU has two additional closed triangles in 1 case, or 0.35 per cent.

Ill 1 lias two less closed triangles than usual in 1 case, or 0.35 per cent.

■* The drawings here reproduced are all from specimens taken iu the eastern and
central parts of the United States and adjoining British Provinces. They are
selected from the series of about 170 l)elonging to the United States Department of
Agriculture.

26

NORTH AMERICAN FAUNA.

[No. 12.

Tlie value of the structure of the bony palate as a taxonomic charac-
ter was first pointed out by Cones,' who, however, considered it of rather
more importance than it really is. It was at first supposed that the
bony palate of all the members of the genus Microtus differed in a con-
stant way from those of Evotomys. Mr. Oldfield Thomas has, however,
recently described a Microtus {M. chinoisis)
in which the palate structure of JEvotomys is
almost exactly reproduced ; and on further

.J.J

Fig. 1. — First upper molar
iu six specimens of J/t-
crotiis pennsiilvaiiiens.

Fig. 2.— Second up-
per molar in .six
specimens otMicro-

tuS2H'nnsi/lra»!ciis.

Fig. 3. — Third ujiper molar iw
eighteen specimens of Microtus

pennnjhmnictis.

study it appears that several well-marked types may be recognized
among the species of the genus. These forms of jialate furnish char-
acters of considerable worth iu defining many subgenera. In all,
several structures remain sufficiently constant to serve as convenient
landmarks. The anterior portion of the bony palate, or that formed

exclusively by thepremaxillaries and
maxillaries, has no special interest, as
it shows very trifling variations. All
the characters of importance are de-
rived from the part lying behind the
maxillo-palatine suture. This suture
in the typi' al palate, or that occurring

Fig. 4. — First lower molar iu
eighteen specimens of Microtus
peniLsijlranicxis.

Fig. 5. — Second lower
molar in foiir speci-
mens of Microtus penn-
sylvanicus.

Pig. 6.— Third lower
molar in four specimens
oi Microtus pcimmjlran-
icug.

in true Microius and iu the great majority of species and subgenera
(lig. 7 A) forms a broad, U -shaped loop, the convexity of which is directed
forward and whose apex lies about opposite the middle of the second
molar. From this point the suture on each side sweeps rapidly back-
ward and outward until, at the level of the anterior edge of the posterior
molar, practically the whole width of the palate is occupied by the pala-
tine, and the maxillaries are reduced to a narroAV rim around the edges
of the alveoli.

Monogr. N. Am. Rodeiitia, p. 133, 1877.

July, 180G.]

CHARACTERS OF BONY PALATE.

27

I'ntil just before acquiring- its greatest width, the surface of the pal-
atine is on the same level with the rest of the bony palate, but imme-
diately on reaching this point it changes abruptly at the sides, more
gradually iu the median line, to the level of the anterior border of the
nteri^terygoid fossa, which lies about 0.5 mm. dorsad of the maiu part
of the bony palate. In the median line the palatiue slopes gently dor-
socauda<l to the edge of the interpterygoid fossa, a distance usually of
about 1 mm., but at the sides it l)realis away suddenly, and tlie spaces
between the median sloping ridge and maxillaries are occuijied by con-
spicuous pits (fig. 7 A, Lp). The floor of each pit is continuous with
the backward projection of the palatine, which runs out to join the

Fig. 7. — Palatal view of skull of MicroHis {Microtis) aivalis (A) and Evotomyg gappeii (B). (x 3).
i./a., interpterygoid fossa (reference line crosses pterygoid fossa); i.fn., incisive foramen; l.hr.,
lateral bridge; Z. r/r., lateral groove ; L^J-, lateral pit; in. r., median ridge; wz., maxillary ; ^jL, ^/Z'.,
palatine; pmx., premaxillary ; lit., pterygoid (reference line crosses jiterygoid fossa); s. in. »•., slop-
ing portion of median ridge.

pterygoid of its side (fig. 7 X^pt.). The ventral outline of the inter-
pterygoid fossa (fig. 7 A, i.fa.) forms three sides of a figure, which is
nearly a parallelogram, open at one end, the longer axis parallel with
the main axis of the skull, and the length more than double the width.
In front and for a short distance at the sides the fossa is limited by the
palatines (fig. 7 A, j>/',), l)ut the greater i)art of its boundary is formed
by the pterygoids (fig. 7 X^pt.). The open end lies between the haniu
lar processes of the pterygoids. Extending back from the incisive for-
au)iua are two distinct lateral grooves (fig. 7 A, /. </r.), which traverse
the bony palate longitudinally, leaving between them a ridge which pos-
teriorly is continuous with the sloping median ridge already described.
In these grooves open numerous foramina, larger and more crowded
just in front of the region from which the bony palate slopes away to

28 NORTH AMERICAN FAUNA. [No. 12.

the level of the pterygoids. Tlie median ridge just here widens
abruptly and sends out on each side a short process, which is met l)y a
similar one arising from the jjalatine on the op])osite side of the groove
(fig. 7 A, /. hr.). These processes usually meet and fuse, thus com-
pletely obliterating the groove, though they are frequently separated
by a narrow space. In Uvotomys (fig. 7 B) tlie sloi>ing part of the
median ridge has disappeared, together with the lateral pits, but
traces of the median ridge (fig. 7 B, m. r.), tlie lateral grooves (fig.
7 B, /. gr.), and the bridges (fig. 7 B, /. br.) may still be recognized.

At different times subgeneric weight has been given to the form of
the external ear, and to the proportional length of the tail to the head
and body. Neither one, however, is of any value, except in special,
isolated cases. The form of the ear is essentially the same in all the
subgenera, though there are slight modifications in length and in the
development of the valvular fold by which the meatus is closed.

The relative length of the tail is far too variable to serve as a useful
diagnostic character.

KEYS.

The following keys to the genera and subgenera of Mkrothuv are
wholly artificial and do not bring the groups together according to
natural affinities. Since analytical keys are of no value except as aids
in identifying specimens, it is necessary that they should be based on
characters that may be studied without difficulty in ordinary museum
material. Such material, however, is usually so imperfect that a single
key made with reference to one set of characters (as, for instance, the
form of the bony palate or the number of mammse) might be of little
use. Hence several keys are here introduced, each based primarily ou
a special set of structures. Of the tliree keys to the genera, No. 1 is
made, so far as possible, with reference to the skull alone; No. 2, with
reference to the teeth, and No. 3, with reference to external characters.
Of the keys to the subgenera of Microtus^ No. 5 is based primarily on
characters derived from the structure of the bony i)alate, and is thus
useless for the identification of specimens the skulls of which are not
available for stud}'. Key No. G is based on the pattern of enamel fold-
ing and may be used with specimens having broken skulls. The lines
in italics inserted in parentheses in this key are for the identification of
individuals with abnormal enamel patterns. These usually occur in the
proportion of about one to four (see p. 25). Hence, one fourth of any
given lot of specimens will agree with the characters given in paren-
theses-, the great majority, however, with those in heavy type. Key
No. 7, based primarily on the mammie and footpads, is made almost
exclusively with reference to external characters. It is necessarily
incomplete, since the number of mamnuTB and footpads is in several
instances unknown. It is, of course, impossible to use this key except
with alcoholic si)ecimens or freshly killed animals. Key No. 8 — if it

JULY.1S9C.] KEYS TO GENERA. 29

may be called a key — is a rough grouping' of the subgenera of Micrgtus
according to the essential characters used iu the chissilicatiou here
adopted. The keys are in all cases based on the characters of adults
only.

1. KEY TO THE GENEEA OF MICKOTlNJi.

[Based primarily on tlie sliull.l

Skull of adult more thau 50 mm. loug Fiher

Skull of adult less than 45 mm. loug.

Molars rooted; skull always less than 30 mm. long.

Posterioi' border of j)alate a thin-edged shelf, contiuuous between

alveoli of posterior molars Evotomys

Posterior border of palate not forming a shelf Phenacomys

Molars rootless; skiiU often more than 30 uuii. long.

Middle part of zygoma expanded so as to form an oblique

plate about 4 mm. broad Leminus

Middle part of zygoma only slightly expanded.

Rostrum about \ total length of skull Synajitomys

Eostrum more thau ^ total length of skull.

Postorbital process of squamosal peg-like Dicrostonyx

Postorbital process of squamosal shelf-like Microlua

2. KEY TO THE GE2fEEA OF MICEOTIX^.

[Based priniarily on tlu- teeth.]

Length of maxillary tooth row in adult more than 14 mm Fiher

Length of maxillary tooth row iu adult less than 13 mm.

Roots of lower incisors or inner (liugual) side of molar roots.

Upper iueisors grooved Synaptomys

Upper incisors not grooped.

m 1 with 3 closed triangles Lemmus

m 1 with 7 closed triangles Dicrostonyx

Roots of lower incisors on outer side of molar roots,
^lolars rooted.

Teeth weak; triangles tending to remaiu open; salient angles

rounded Evotomys

Teeth strong; triangles (dosed; salient angles sharp Phenacomys

Molars rootless Microtus

3. KEY TO THE GEXEBA OF MICEOTIXJ^;.

[Based primarily ou external characters.]
Tail tlattened laterally Fiber

Tail terete.

Tail shorter than hind foot.

Thumb with strap-shaped nail Lemmus

Thumb without strap-shaped uail.

External ear rudimentary Dicrostonyx

External ear well developed Microtus (Asiatic species

of subgenus Laynrits)
Tail longer thau hind foot.

Upper incisors grooved Synaptomys

Ujiper incisors not grooved.

Color usually reddish; molars weak, with triangles tending

to remain opeu and with salient angles rounded Erotomys

Color brownish, grayish, or yellowish; very seldom reddish;
molars strong, with closed triangles and sharp salient angles.

Molars rooted Phenacomys

Molars rootless Microtiia

30 NORTH AMERICAN FAUNA. []^'o.i2.

4. KEY TO SUBGENEKA OF STXAPTOMYS.

Miuulibnlar molars with closed triangles on outer side Synaptomys

Mandibular molars without closed triangles on outer side Mictomys

5. KEY TO THE SUBGENERA OF MICEOTUS.

[Based primarily on the bony palate.]

Palate normal or nearly so (see p. 27).

Third lower molar with all triangles closed.. La(jnrns

Third lower molar normally without closed triangles.
Claws small, those on front feet always .'ihortest.

Plantar tubercles 6 - Micrntus

Plantar tubercles 5.

Tail more than 30 per cent of total length it-rirola

Tail less than 30 per cent of total length.

nTl with 5 closed triangles Cliilotus

inl with 3 closed triangles Pedomys

Claws large, those on front feet usually longest.

Fur long and soft Fhaiomys

Fur dense and nude-like ntymys

Palate highly abnormal.

Palate ending in a broad median plate cut off from maxillaries at the sides.

Third lower molar with all triangles clo.sed Neofiher

Third lower molar without closed triangles.

Skull flat ; andital bulhe small Hyperacriits

Skull high ; audital bulla} large Jlticola

Posterior border of palate continuous between maxillaries.

Posterior border of palate straight Fothenomys

Posterior border of palate with median projection JvicUomys

6. KEY TO THE SUBGENERA OF MICROTUS.'

[Based iirimarily on tlie t(*th.]
{m 1 with 6 or 7 closed irianglcn.)
{Plantar tiihcreles 5.)

(Small ; not aquatic ; fur short Cirilotns)

{Large ; aquatic ; fur long -^ rricola)

{Plantar tnbercles 6 Jlicrotus)

m i with 5 closed triangles.

m 3 with 3 closed triangles.

m 3 with triangles always closeu Neofiber

m 3 with triangles normally open.
Plantar tubercles 6.

Fur not specially modified, claws moderate.

Posterior loop of m 3 short cr strongly curved; palate normal. Microtus
{Posterior loop of m 3 long and straight : palate ahnormal.)

{Skull hroad and flat ; plantar tubercles -T Eyperacrins)

{Skull not broad and flat ; plantar tubercles 6 Jlticola)

{Fur rcry long and soft, aspect lemming-like, claws rery long. Phaiomys)
{Plantar tnbercles 5.)

{Small ; not aquatic : fur short -. Chilotns)

{Large ; aquatic ; fur long Jrricola)

' Characters in heavy-faced type are those of specimens with normal enamel pat
tern; characters in italics (inserted in parentheses) are those of specimens with
abnormal enamel pattern.

July, 189b-.] KEYS TO SUBGENERA. 31

m 3 with 2 closed triangles.

Triangles in m 3 alternate and closed.

Aquatic ; soles naked ; tail long Neofiber

Not aquatic ; soles hairy ; tail short Lagurus

Triangles in m 3 normally opposite and open.

Claws small, those on hind feet always longest.
Mammffi 8 ; foot pads 5.

Small ; not aquatic ; fur short Chilotus .

{Large; aquatic; fur Ion;/ Jrvivola)

{Mamma 4; foot pads 5; sk>iU liir/h Pedomys)

{Claws large, those on front feet of lev longest.)

{ Fur short and dense J'itjjmys)

( Fur long and soft I'haiomys)

m 1 with 4 closed triangles.

m 3 with posterior loop elongated in axis of jaw.

Skull broad and fiat ; plantar tubercles 5 Hyperacrius

Skull not broad and flat ; plantar tubercles 6 Alticola

(m 3 with posterior loop rounded or crescentiv. }

{m 3 with 3 closed triangles ^ficrotus)

(m 3 irith 2 closed triangles Jrricola)

m 1 with 3 closed triangles.

(m 3 with 3 closed triangles.)
{Plantar tubercles 6.)

{Posterior loop of m 3 .^hort or stronghj citrred: palate normal . . . Microtus)

{Posterior loop of m 3 long and straight: palate ahnormal Alticola)

{Plantar tubercles 5.)

{Mamnue S; palate normal Arvicola)

{Mam mce 4; p)alate abnormal Hyperacrius)

m 3 with 2 closed triangles.

Sole almost naked Arvicola

Sole hairy.

(Palate abnormal Hyperacrius)

Palate normal.

Claws long, all about equal in length Phaiomys

Claws short, those on front feet shortest Pedomys

(m S with 1 closed triangle Hyperacrius)

m 1 with 0 closed triangles.

m 2 and m 3 of approximately the same form Eothenomys

m 2 and m 3 very different in form Anteliomys

7. KEY TO THE SUBGENERA OF MICROTUS.
[Based prim.irily on inamiiiie ami foot pads.]

^lanmia' 10 Phaiomys

Mamma' 8.

Plantar tubercles 6.

Palate normal Microtus

Palate abnormal A Iticola

Plantar tubercles 5.

Conspicuous musk glands on sides Arricola

No musk glands on sides.

Color dark brown Chilotus

Color light grayish or yellowish Lagurus

32 NORTH AMERICAN FAUNA. [Xo.12.

Maiuiu;e 4.

Size very large - - - Xiojiber

Size medium or smal'.

Plantar tubercles 6 AnteUomija

Plantar tubercles 5.

Skull not flattened I'edonuis

Skull flattened.

Palai e normal I'Hymija

Palate abnormal Hijjicracrins

8. SUBGENERA OF MICROTUS GROUPED BY ESSENTIAL CHARACTERS.

Palate normal. — Microtus, Pedomys, Pltymys, ChUotus, Phaiomys, Ai-vicola, Lagunts.

Palate abnormal. — Xeojiher, JUirola, Hyperacrius, Eoihenomys, Anteliomys.

Third lower molar always with closed triangles. — Neofiher, Lagurus.

Third lower molar normally without closed triangles. — Microtus, Pedomys, Pitymys,
ChUotus, Phaiomys, Arvicola, Eothenomys, Anteliomys, Alticola, Hyperacrius.

First lower molar normally with 5 closed triangles and 9salieut augles. — Microtus,
ChUotus, Xeojiher, Lagurus.

First lower molar normally with 3 or 4 closed triangles and 9 salient iingles —
Pedomys, Pitymys, Phaiomys, Alticola, Hyperacrius.

First lower molar normally with 3 closed triangles and 7 salient angles. — Arvicola.

First lower molar without closed triangles. — Anteliomys, Eothenomys.

Third upper molar normally with 3 closed triangles and 7 to 8 salient angles. —
Microtus.

Third upper molar normally with 2 closed triangles and 6 salient angles. — Xeojiher,
Arvicola, Pitymys, Pedomys, Phaiomys, ChUotus.

Third upper molar without closed triangles. — Anteliomys, Eothenomys.

Mammic 10. — Phaiomys.

Mammse 8. — Arvicola, Microtus, Alticola, ChUotus, Lagurus.

Mamm;e 4. — Xeojiher, Pitymys, Pedomys, Anteliomys, Hyperacrius.

Plantar tubercles 6. — Microtus, Phaiomys, Anteliomys, Alticola.

Plantar tubercles 5. — Xeojiher, Arvicola, Pitymys, Pedomys, ChUotus, Lagurus, Hyper-
acrius.

DESCRIPTIONS OF LIVING GENERA AND SUBGENERA.

Genus SYNAPTOMYS Baird.
Synaptomys Baird, Mamm. N. Am., p. 558, 1857. Type Synapiomys cooperi Baird.

Geographic (listrihiition ofty^ye species. — Boreal, Transition, and north-
ern edge of Upper Anstral Zone in eastern Kortli America from the
Atlantic coast to Minnesota.

Geographic (listribution of genus. — jSTortli America from northern edge
of Lower Austral Zone northward.

Essential characters :

Upper incisors with distinct longitudinal grooves.

Lower incisors with roots on inner (lingualj side of molars.

Molars rootless.

Enamel pattern characterized by great depth of reentrant angles en outer side
of maxillary teeth and on inner side of mandibular teeth.

m 1 with three closed triangles and two transverse loops, or with four trans-
verse loops and no closed triangles.

m 3 with four transverse loops and no closed triangles.

Feet not specially modified.

Soles and palms with well-developed tubercles.

Thumb with large flattened ligulate nail.

Tail very slightly longer than hind foot, terete.

External ear well developed.

July, 1896.] GENUS SYNAPTOMYS. 33

Skull. — The skull of Si/najjtomy.s (fig. 9 and PL I, figs. 12, 13) is moder-
ately broad, flat, aud massive, much less so than iu the other Lemmi.
Eostrum short (nasal bones about one-fourth occipito-nasal length) and
strougly deflexed; zygomatic arches not broadly flaring as iu Lemmus
and Dicrostonyx,^ though more so than in the voles; middle portion of
zygoma very slightly expanded, the outer surface nearly vertical; brain
case not greatly broadened or flattened, and seldom if ever conspicu-
ously ridged or furrowed; interparietal with rounded corners, the
antero-posterior diameter more than half the transverse diameter;
pterygoids short; interpterygoid fossa about one-sixth basilar length
of skull; posterior border of bony palate ending nearly as in typical
Microtus. (See p. 2G, PI. II, fig. 5, and fig. 7, p. 27.) Front edge of
squamosal forming a narrow, shelf-like postorbital process.

Teeth. — Anterior faces of upper incisors with distinct longitudinal
grooves. Lower incisor terminating posteriorly a little in front of the
hinder edge of the back molar. Throughout its length each mandibu-
lar incisor lies wholly on the inner (lingual) side of the molar series.
(PI. Ill, fig. 1.)

The molars are all rootless. The upper molar series is about one-third
the basilar length of skull, the lower series slightly less. The enamel
pattern (figs. S and 10) is characterized by the great depth of the outer
reentrant angles in the maxillary teeth and of the inner reentrant
angles in the mandibular teeth. Of the maxillary teeth m 1 and m 2
show no important peculiarities of form except that the outer reentrant
angles cut across to the enamel of the extreme inner side, a feature
shared by Lemmus alone. The posterior upi)er molar, however, like
that of Lemmus, differs widely from the corresponding tooth in all other
Microtina'. It is formed of four transverse loops. The first and second
of these loops are isolated by two deep reentrant angles on the outer
side of the tooth, while the third is formed by an equally deep dein-es-
sion on the inner side. The reentrant angles and closed triangles on
the inner side of the mandibular molars are greatly developed at the
expense of those on the outer side. In the subgenus Mictomijs the lat-
ter wholly disappear except in the last tooth. This has a reentrant
angle near the middle, but no closed triangle.

External form. — In general appearance Synaptomys resembles the
Microti much more closely than it does the Lemmi, a fact which has
given rise to the rather inappropriate names 'lemming- vole' and 'false
lemming.' The species of Synaptomys are thick-set microtines with
large heads, ears that just appear above the moderately long fur, short
tails, aud small feet. In color they are all dull brownish, darker on the
back, paler on the belly. The palms and soles are tuberculate, as in
the voles.

General reniarks.— Synaptomys differs from all the other genera of

'Tlip ratio of zygomatic breadth to basilar l<'njj;th is approximately 70 in SynaptomySy
75 iu Lemmus aud Dicrostonijx, and 65 iu Microtus.
16933— No. 12 3

34 NOKTH AMERICAN FAUNA. [No. 12.

Microthuc in its grooved incisors. From tlie other leniniiiig'S it may be
known by its numodified external form, and from the A'oles by the
characters of its molars.

Subgeuns SYNAPTOMYS Baird.
Sijna2)tomi/s Baird, Mamm. X. Am., p. 558, 1857. Type Si/napiomiis cooperi ]5aird.

Geo(/rapMc distribution of type species. — Boreal, Transition, and north-
ernmost edge of Austral zone in eastern United States and adjoining
British Provinces; west to Minnesota, south to Iowa, Indiana, Ohio,
and Maryland.

Geograpliic distribution of subgeuns. — Boreal zone to northern edge of
Lower Austral zone in eastern Canada and eastern United States;
west to Minnesota, south to Kansas and Virginia.

Essential cliaractcrs :

Eostrum very heavy.

Palate nearly as in trne Microtus.

Mandibular molars Avith closed triangles on outer side

Mamm* 6.

Sl-uU. — The skull of true ^Synaptomys (flg. 9 and PI. I, flg. 13) differs
from that of Mictomys in the remarkably heav^^ rostrum and in certain
slight details in the form of the bony palate. The latter is almost
exactly as in typical Microtus, the slight peculiarities in form being
well within the limits of variation in the latter.

Teeth. — The incisors in true Synaptomys are, like the rostrum, exces-
sively strongly built. The grooves are usu-
ally sharply defined and placed near the
outer edges of the teeth.

The maxillarj^ teeth differ in no way from
those of the species of Mictomys. In the
Fig. 8.-Enamei pattern of molar molars of the lowcr jaw, howevcr, the outer

teeth of Synaptomys cooperi. (s 5.) ^ „ ■,,,■,'• , i 1

edge of each tooth is cut by a deep reen-
trant angle which isolates a large outer triangle (fig. 8).

Mammw. — The nitmber of mamm;e in Synaptomys has been variously
recorded as four and six. Dr. Cones, in his monograi)h of the American
Microtinw, states that he finds six, four pectoral and two inguinal, in a
female from Brookville, Ind.' Quick and Butler,^ however, noted only
four, two pectoral and two inguinal, in specimens from the same local-
ity. Mr. Vernon Bailey records six mammae in a female collected for
the United States Department of Agriculture at Ann Arbor, Mich.,
and I find the same number in an alcoholic specimen taken at Eogerfe-
ville, Tenn. It is probable that six is the normal number, and that
Quick and Butler overlooked the ]»osterior pair on the breast, as these
are smaller than the others, at least in the alcoholic specimen from
Tennessee.

1 Monogr. N. Am. Rodentia. p. 236.

2 American Naturalist, XIX, p. 114.

July, 1896.]

SUBGENUS MICTOMYS.

35

General remarlxS. — The characters distiugiiishing- the subgenera Syn-
aiitomys and Mictomys are discussed under the latter.

Three species of true Synapiomys are now known: *S'. cooperi Baird,
S.fatuHs Bangs, and *S'. hdaletes Merriam.^

Subgenus MICTOMYS True.

1894. Alictomtjs True, Proc. U. S. Nat. Mus., XVII, No. 999, p. 242. Advance sheet,
April 26, 1894 (full genus). Type Mictomj/s inriuitus True.

1896. Mictomys Merriam, Proc. Biol. Soc. Washington, X, p. 57, March 19, 1896
(subgenus).

Geographic distribution of type species. — Synaptomys inmiitus is known
from the type locality only, Fort Chimo, Ungava, Labrador.

Geographic distribution of subgenus. — Hudsonian zone from Labrador
to Alaska, south to northern California.

Essential characters :

Eostrum slender.

Palate not as in true Miciotus.

Mandibular molars without closed triangles ou outer side.

Mammte 8.

Slcull. — The skull of Mictomys is in general much like that of Synap-
tomys proper, but the whole rostral part (including incisors) is dispro-
portionally slender and weak
(fig. 9, and PI. I, fig. 12). The
bony palate is formed on the same
plan as that of true Synaptomys
or of Microtus i^roper, but differs
from both of these in the prolon-
gation of the median ridge as a
spine projecting into the inter-
pterygoid fossa.

The i^terygoids are usually
longer and more slender than in
Synaptomys, and the hamular i^ro-
cesses less strongly bent outward.

Teeth. — The incisors in Mictomys are much smaller in proportion to
the size of the skull than in the subgenus Synaptomys. The grooves
in the upper incisors are usually nearer the middle of the tooth, and
less well defined than in true Syna2)tomys.

The maxillary teeth (fig. 10) are exactly as in the subgenus Synap-
tomys. The lower molars, however, difter from those of true Synaptomys
in the absence of reentrant angles on the outer borders of all but the
hindermost. Even in this the reentrant angle is never deep enough to
isolate an outer triangle.

Mamma\ — In the type of Synaptomys innuitus there are eight mammse,
two more than have been recorded in Synaptomys i^roper. ^Yhether

Fig. 9. — a. Hitnaptoiiiys helaleteg; b. Si/Haptomys
v:rancjeli.

See Merriam, Proc. Biol. Soc. Washington, X, p. 57, 1896.

36 NORTH AMERICAN FAUNA. [X0.12.

tliis difference is constant or otherwise, it is, liowever, impossible to

say.

General remarlxS. — Mictomys was lirst described as a full genus, but
tbe characters ou which it rests are of no more
than subgeueric importance. The group is
distinguished from true Syna;ptomys by the
slender rostrum and incisors, slightly different
form of bony palate, creuulate outer border of

Fig. 10— Enamel pattern of lower molars, and probably by the number of

molar ic.ih Synaptomys in- ^^.^^^^^^^ ^^^^^
nmtus. (x5.)

Four species of Mictomys have thus far been
described, Synaptomys innnitns (True), ^S. wravgeli Merriam, /S'. dalli
Merriam, and S. truei Merriam.^

Genus LEMMCS Link.

1795. Lemmus Link, Zool. Beytriige, I, Pt. II, ]). 75, 1795. Type by elimination Mas

lenunus Liuu.
1811. Ml/odes Pallas, Zoogr. Rosso- Asiat., I, p. 172, 1811 (part).
1877. Ml/odes Cones, Mouogr. N. Am. Rodentia, p. 237, 1877, anil most subseijuent

anthers.

Geographic distribution of type species. — Arctic region in Asia and
eastern Europe.

Geographic distribution of genus. — Arctic region in both hemis-
pheres.

Essential characters :

Upper incisors without grooves.

Lower incisors with roots on inner (lingual) side of molars.

Molars rootless.

Enamel pattern as in tSijnajytomys.

Feet highly modified.

Palms and soles without well-developed tubercles.

Thumb witli large flattened ' strap-shaped ' nail.

Tail shorter than hind foot, terete.

External ear small but well developed.

SI-hU. — The skull of Lemmus (PI, I, tig. G) is perhaps the most highly
modified in the family Microtina\ The rostrum, like that of Synapto-
mys^ is short in proportion to the length of the skull (nasal bones con-
tained about three and one-half times in occipito-nasal length), the
dorsal profile bent abruptly downward. Zygomatic arches very ab-
ruptly and broadly tlaring, each expanded near the middle into a wide,
strongly oblique plate.^ Brain case broad, flat, and subquadrate iu
outline, but dwarfed in appearance by contrast with the large zygomata.
Pterygoids short (about as iu Synaptomys). 13ony i)alate terminating
essentially as in Synaptomys, but lateral pits very deep and anterior

' See Merriam, Proc. Biol. Soc. Washington, X, p. 61, 1896.
^ These plates may be nearly 5 mm. across iu the widest part.

Jl-LY, 1896.]

GENUS LEMMUS.

37

Fig. 11.— Enamel pattern of molar
teeth, Lemmus leinmus. (x5.)

edgeof interpterygoicl fossa carried forward over (dorsad to) overliang-
iiig edge of palate (PI. 11, fig. 14). The anterior edge of the squamosal
forms a narrow but distinct shelf-like postorbital process, much as in
Synaptomys^ but more strongly developed.

Teeth. — The dentition of Lemmus is essentially the same as that of
Synaptomys. The upper incisors are, however, much more slender in
proportion to the size of the skull, and are without the peculiar grooves
always present in Synapiomys. In the pattern of enamel folding, the
only difference between the two genera is that the third transverse loop
in the hindermost maxillary tooth is iso-
lated by a single reentrant angle in l>iynap-
iomys, and by the contact of two reentrant
angles in Lemmus (fig. 11).

External form. — In external form the
species of Ljemmus diflter very widely from
all other microtiues except Dicrostonyx.
The head is disproportionately large for

the short thick body,^ while the tail is reduced to a mere rudiment only
about two-thirds as long as the hind foot. The feet are highly modified
to fit the animals to their fossorial habits. While the hind feet are
unusually large and strong, the front feet are even more specialized.
The thumb is provided with a large ligulate nail and the fingers are
armed with long, sharp claws (fig. 12). The claws are, however, sim-
ple in form and are not subject to the periodic changes that occur in
those of Dicrostonyx.
In the alcoholic specimens that I have examined the palms show no
trace of tubercles, but the soles bear indications of sev-
eral very small and exceedingly rudimentary pads close
to the base of the toes. The fur is remarkably long and
dense, the palms and soles densely furred, and the tail
provided with a pencil of stiff bristle-like hairs longer
than the tail vertebrie.

General remarls. — The species of Lemmus are true
lemmings with highly modified skull and external form.
With these characters they combine the dentition of
Synaptomys without, however, the peculiar incisors of
the latter. Ljemmus difiers from Synaptomys in its highly
modified skull and external form as well as in the dental character just
mentioned. From Dk-rostonyx it is distinguished by cranial and dental
characters and by the well-developed external ears (fig. 15), as well as
by the simple claws and large thumb nail.

The species of Lemmus at present recognized are L. lemmus (Lin-
iiieus), Tj. ohensis (Brants), L. sehistieolor (Lilljeborg), and L. niqripes
(True).

Fig. 12.— Left front
foot, Lemmus
li'mmux (hair re-
moved).

This peculiarity is carried even further iu Lemmus than in Synaptomys.

38 NORTH AMERICAN FAUNA. [No. 12.

Genus DICROSTONYX Gloger.

1830. Cuniculus Wagler, Nat. Syst. d. Amphibien, p. 31, 1830 (part).
1877. Cuniculus Cones, Moiiogr. X. Am. Rodeutia, p. 243, 1877.

1841. Dicroxioniix Gloger, Gemeinn. Hand- u. Hilfsbucli d. Naturgesch., pp. XXXI, 97,
1841. Type, an American species, probably Mus hudsonius Pall.

1854. " MyoJemmus Pomel, Ann. Sci. Soc. Auvergne, 1854 " (fide Trouessart).

1855. Misothermus Hensel, Zeitscbr. der Deutsch. geolog. Gesellscb., VII, p. 492, 1855.

Type Myodes forquatus Pall.
1881. Borioikon Polyakoff, Mem. Acad. Imp. Sci. St. Petersbourg, XXXIX, snppl. p. 34,
1881. Type Jlyodex torquatus Pall.

Geographic distribution of type species. — Arctic America.
Geographic distribution, of genus. — Arctic region in both liemisplieres.
Essential characters:

Upxier incisors without grooves.

Lower incisors with roots on inner (lingual) side of molars.

Molars rootless.

Enamel pattern characterized by approximate equality of reentrant angles.

m 1 with 7 closed triangles and 2 transverse loops.

m 3 with 3 or 4 closed triangles and 2 transverse loops.

Feet highly modified.

Palms smooth; soles with rudimentary tubercles.

Thumb with a rudimentary nail.

Tail shorter than hind foot, terete.

External ear rudimentary.

STciill. — The skull of Dicrostonyx (PI. I, fig. 14) in a general way
resembles that of Lemmus, but is smaller and more lightly built. The
zygomata are less broadly flaring and the expansion uear the middle is
comparatively slight. The outer face of the expanded portion, as in
Lemmus, is strongly oblique. The rostrum is also lighter and more
slender. While the pterygoids are proportionally longer than in Lem-
mus, the posterior edge of the bony palate is formed exactly as in the
latter (PI. II, figs. 12 and 14), The anterior edge of the squamosal
gives off a conspicuous peg-shaped postorbital
process very different from the postorbital ])r()C-
ess in Lemmus or any of the other Microtinc:.
These pegs are especially conspicuous when the
skull is viewed from the ventral aspect.
Fig. i3.-Enamei pattern of Tccth. — lucisors csseutially as in Lemmus.
molar teeth, Dicrostoni/x from ]\iolars Tootlcss. Pattern of cuamel folding ( lig.

Uugava, Labrador. (x5.) ^ ^, -, . m, , r. .-• . ^ • -, <• i i j-i

13) very different from that of either of the other
genera of Lemmi and in some respects resembling that of the Jlicroti.
The reentrant angles on the opposite sides of the teeth are approxi-
mately equal in depth,«thus producing closed triangles of nearly the
same size on the two sides. The first lower molar contains seven closed
triangles in addition to a transverse loop at each end. The second
lower molar contains a posterior loop followed by four alternating
closed triangles and an anterior transverse loop, which is much flattened

JCLY, 1896.]

GENUS DICROSTONYX.

39

\T"

'^^^^

Fig. 14. — Ear, (a) Dicrostonyx, (b) Lemmus
(double natural size'

aud so small that the tips of the salient angles do not reach to the level
of the tips of tlie other salient angles of the tooth. Occasionally the
anterior outer triangle opens into the transverse loop. The posterior
lower molar has a posterior transverse loop followed by three large
closed or nearly closed triangles (two on the inner side), and a fourth
smaller triangle on the outer side.' The maxillary teeth have each a
large anterior loop. This is followed in the first hy five alternating
closed triangles and a small postero-
external loop, in the second by four ^,^;Sxt>. a.
closed triangles and a small postero-
external loop, and in the third by fonr
closed triangles and a small ronnded
terminal loop.

External form. — In external form the
species of Dicrostonyx are even more
specialized than the members of the
genus Lemmus. As in the latter, the
head is very large, the tail is reduced to
a stub, shorter than the hind foot, and
the feet are highly modified for digging.
The external ears are, however, mere naked folds of integument lying
just behind the meatus (fig. 14 a). The fur is long and dense, much as in
Lemmus. The palms and soles are densely furred, and the tail is pro-
vided with a stiff pencil of bristle-like hairs, longer than the tail vertebrae.
The hind feet are very broad, the breadth at base of toes being about
one-half length of foot.- On the hind foot there are several minute,
faintly developed tubercles near the base of the toes. The palms are^

however, perfectly smooth. The claws on
the hind feet are large and well formed,
though in no way different from those of
Lem rims. Those on the front feet are very
highly modified, and present seasonal
changes in size and form unknown else-
where among the 2licrotiniv. The thumb
(fig. 15) is greatly reduced in size. The
thumb nail is so small as readily to escape
notice, but the ball of the thumb projects
as a distinct tubercle, the surface of which
is covered with a thick layer of corneous tissue. The claws on the
second and fifth fingers are large, though not peculiar in form. The
two middle claws, on the contrary, while in summer not different
from those of Lemmus, are in winter very greatly enlarged (fig. 15), and

'lu IJicrostoiiyj- torquatus there is a niiiuite supplemental anterior internal loop
wbich is absent in the species that occurs in Labrador.
-In Lemmus this breadth is only about one-third length of foot.

Fig. 15. — Left front foot of three speci-
mens of Z)icro*<o)ii/a; from Alaska, show-
ing successive stages in the develop-
ment of the claws (hair removed).

40 NORTH AMERICAN FAUNA. [X0.12.

wliollj' unlike those of any other microtiue. Dr. Coues's description
of tlie claws of Bicrostonyx is so interesting that it may be quoted
almost entire. He says (Monogr. IsT. Am. Eodentia, pp. 248, 240) :

The two middle fore claws attain their maximum of development iu winter. In
spring and early summer these claws do not appear very different from those of
Mijodi's [= Lemmus'], though averaging larger, more bulbous at base underneath, with
the terminal portion slenderer, straighter, and sharper. This bulbous portion under-
neath grows out simultaneously with increase in length and amount of curvature of
the main jiortion of the claw, until it equals or even exceeds the length of the latter,
and is quite as stout, or even stouter, being somewhat broad and pad-like. At this
period it runs the whole length of the claw, from which it is separated by a groove
along the sides, and by a notch at the end, both of varying depth. The claw then
looks nearly like two claws, one underneath the other. The pad woubl then seem to
gradually sever its connection with the main claw by progressive increase iu depth
of the constriction marked by the lateral groove and terminal notch, as well as by
loosening from the base, when it appears like an excrescence ; it is finally lost. Thus
the process appears to be a jieriodical one, like the shedding of the horns of rumi-
nants, and not continually progressive with age; and would seem to be connected
with the particularly fossorial habits of the quasi-hibernating animal that digs gal-
l,eries under ground in which to reside during the cold season, as compared with its
freer and more active mode of life in summer. At the period of the maximum
development of the claws these equal or surpass half an inch in length. » * #

General remarls. — Divrostonyx is so readily distinguished by its
peculiar dentition, highly modiiied feet, and rudimentary external ears,
that it requires no detailed comparison with any other genus.

While Dicrostonyx torquatus (Pallas) is the only species now recog-
nized, there are doubtless several others.

Genus PHENACOMYS Merriam.
1889. Phenacomi/s Merriam, North Ameijican Fauna No. 2, p. 28, October 30, 1889.

GeograijliiG distribution of type species. — Phenacomys intermedins is
known only from the type locality, Kamloops, British Columbia.

Geographic distribution of genus. — Boreal North America; also
recorded from the bone breccia of Ber.emend, southern Hungary, and
the Forest Beds of Norfolk and Suffolk, England (Nehring, Naturwis-
senschaftliche Wochenschrift, Xr. 28, ]). 346, July 15, 1894.)^

Essentia I cha ra cters :

Upper incisors without grooves.

Lower incisors with roots on outer side of molars.

Molars rooted.

Enamel pattern characterized by approximate equality of reentrant angles in
maxillary teeth and great depth of reentrant angles on inner side of mandibular
teeth.

m 1 "with five closed triangles.

m3 with two or three closed triangles,

' I have not seen the original description of the remains from Beremend (described
by Nehring in Naturwissenschaftliche Wochenschrift, 1883). The teeth from the For-
est Beds represent an animal which is certainly not Phenacomys. (Seenoteou JrricoZa
iniermcdiKS Newton ou page T.o.)

Jtn-r, 1896.]

GENUS PHENACOMYS.

41

Fig. 16. — Side view of molars,
Phenacomijs. (a) young, (6)
adult. (x3.)

Bony i)alate not terminating in a tliin-edged sbelf continuous between alveoli
of posterior incisors.
Feet not specially modified.
Thumb with a small pointed nail.
Tail longer than hind foot, terete.
Fur not specially modified.

Sl-nJl. — The skull of rhenaeomys (PI. I, fig-. 5) dififer.s very slightly in
general form from tlmt of typical Microtus. The brain case is, how-
ever, flatter and more quadrate (but no more
so than in the subgenera X^(;r/?(r».s' iindl'iti/iuys),
and the zygomata bend down somewhat more
abruptly in front. The expansion of the zygo-
ma at the region of contact between the malar
and the zygomatic process of the maxillary is
rather more abrupt than is usual in MicroUis,
but the difference is very trifling. The postor-
bital processes of the squamosals are slightly
more prominent and angular than in il//c>-o/*^s
arvalis or M.])ennsylvaniciis,hut scarcely more
developed than in 71/. agrestis, and consider-
ably less so than in M. alien i. The audital
bullne are proportionally about the same size
as or slightly smaller than in Microtus arvalis.
They are more globular and less ' subfusiform '
than in the typical species of true Microtus, but closely resemble those
of M. agrestis. The palate (PI. II, fig. 1) is formed essentially as in the
members of the subgenus Lagurus (PI. II, figs. 3 and 4).

Teeth. — The teeth of Phenaconiys differ in many ways from those of
the other voles. In young individuals the molars (fig. 10) are rootless,
but by the time the animals are full grown each molar has developed
two distinct roots, which, however, remain open until an advanced age,
though not so long as in the genus Evotomys.
The pattern of enamel folding (fig. 17) is essen-
tially the same as that of the voles of the sub-
genera Fedomys and Fhaiomys. (See pp. 50 and
57.) The differences are to be found in the lower
molars where the reentrant angles on the inner
side are proportionally deeper and those on the
outer side proportionally shallower than in Ted-
omys. There is a corresponding difference in the size of the closed
triangles on the opjiosite sides of the teeth. The anterior outer loop
in the second lower molar is especially reduced.

In PItenacomys the root of the lower incisor runs back between the
roots of the second and third molars, and terminates on the outer side
of the tooth row in the ascending ramus of the jaw, at about the level of
the middle of the posterior molar, and distinctly below the dental fora-
men, (PI. Ill, fig. 2.) While exactly this condition is not found else-
where except in Evotomys, it is somewhat closely approached in Fiber.

Tig. 17.— Enamel pattern of
molar teeth, Phenacomys
cclatvs. (x5.)

42

NORTH AMERICAN FAUNA.

[Xo. 12.

External form. — In external form the species of Fhenacomys show no
peculiarities to distinguish them from the other voles. The body, tail,
feet, ears, and eyes are usually jiroportioned about as in Microtns arra-
lis or ]\[. ansierus. In F. longicauda, however, the tail is proportionally
longer than in any of the other known species.

General remarTcs. — Phenacomys is readily distinguished from Microtns
by the rooted molars. From Uvotomys, rhenacomys is separated by
certain characters in the form of the skull, and more especially of the
bony palate, as well as by peculiarities in the teeth. The differences
between the three genera may be comjiared in detail as follows:

Mierottis.

£votomys.

Phenacomys.

Koot of lower incisor above den-

Eoot of lower incisor below den-

Root of lower incisor bolow den-

tal foramen.

tal foramen.

tal foramen.

Molars rootless throughout life..

Molars rooted in the adult, the

Molars rooted in the adult, the

roots closed in extreme old

roots closed in extreme old age.

Molars large and strong, the

age.
Molars small and weak, the sal-

Molars large and strong, the sal-

salient angles sharp.

ient angle rounded.

ient angles sharp.

Reentrant angles on oviter and

Reentrant angles on outer and

Reentrant angles on inner side of

inner sides of lower molars ap-

inner sides of lower molars

lower molars verj' much deeper

proximately equal in depth.

approximately equal in depth.

than those on outer side.

Skull strong and angular

Skull weak and rounded

Skull strong and angular.

Posterior border of bony palate

Posterior border of bony palate

Posterior border of palate never

extremely variable.

a thin-edged shelf continuous
between alveoli of posterior
molars.

a thin-edged shelf.

Middle portion of zygoma dis-

Middle portion of zygoma

Middle portion of zygoma dis-

tinctly expanded.

scarcely expanded.

tinctly expanded.

Since the discovery of the genus Phenacomys the following species
have been described: P. intermedins Merriam, P. celatus Merriam, P.
ungara Merriam, P. latimanus Merriam, P. orophilus 3Ierriam, P. longi-
caiida True, P. trnei Allen, and P. oramontis Ehoads. The status of
these forms is wholly a matter of conjecture.

C4enus EVOTOMYS C'oues.

1839. Myodes DeS61y8 Lougcbamp.s, fitudes de Micromammalogie, p. 87, 1839 '(sec-

tion).
1883. Ifyodcs Lataste, Le Natnraliste, Tome II, p. 349, 1883 (subgenus).

1840. H'jpuda'us Keyserling and Blasius, Die Wirbelthiere. Europa.s, p. 34, 1810 (sub-

genus). Type Miis glareolns Scbreber. (Not Hypnduus Illiger, 1811.)
1857. Hypudaus Baird, Mamm. N. Am., p. 513, 18.57 (subgenus).
1874. Evotomys Coues, Proc. Acad. Nat. Sci. Pbila., p. 186, 1874 (genusj. Type J/h«

rutilus Pall

Geographic distribution of type species. — xVrctic region in Europe and
Asia, possibly in America also.

Geographic distrihntion of genus. — Boreal Xorth America, Asia, and
Europe.

JCLY, 1896.]

GENUS EVOTOMYS.

43

Essential characters:

Upper incisors ■without grooves.

Lower incisors with roots on ourer side of molars.

Molars rooted.

Enamel pattern characterized by approximate eiiuality of reentrant angles.

m 1 with five closed or nearly closed triangles.

m 3 with three closed triangles.

Feet not specially" modified.

Thumb Avith a small, pointed claw.

Fur not specially modified.

Tail longer than hind foot, terete.

Sicull. — The skttll of Urotomi/s (PI. I, tig. -t), as compared with that

of the other voles, is characterized by a general weakness and lack of

angularity. All the outlines are full and

rounded, and the ridges and furrows are slightly

developed, even in extreme old age. The in-

terorbital region is broader and the audital

bulliie are larger and more inflated than usual in

Microtus and Phenacomys. On the other hand,

the zygomata are very slender and scarcely

widened in the region of contact between the

jugal and the zygomatic process of the maxillary.

The mandible also is slender and weak. The

bony palate terminates in a thin-edged shelf,

continuous between the alveoli of the posterior

incisors (fig. 7 and PI. II, fig. 10). The structure

is very different from that found in Flienacomys

and in typical Microtus}

Teeth. — The incisors are exactly as in Phenacomys. The lower incisor

runs back along the lingual side of the first and second molars, but

crosvses the line of the molar tooth row between the second and third

molars, terminating in the ascending ramus of the mandible at about

the level of the middle of the posterior molar and distinctly below the

dental foramen. The molars are rootless in the young (fig. 18), but in
the adult each is provided with two distinct roots
which eventually becomefull}- closed.^ In one very
old individual the crowns of the lower molars are
completely worn away, so that each root, with the
exception of the anterior root of m 3 (which lias
been shed) stands alone like a simple, round-topped
tooth (PI. Ill, fig. 1). The molars are all very

narrow and weak, in this character strongly contrasted with the strong,

broad teeth of Microtus and Phenacomys.

' For detailed comparison of the palates of Evoioniys and Microtus see pages 26-28.

^lu the original description of the genus Phenacomys (North Am. Fauna No. 2,
p. 30) it is stated that " Phenacomijs has genuine rooted molars, not half-rooted
molars like those of Ecofomys, which grow from persistent pulps." Erotomys, how-
ever, has as perfectly rooted molars as Phenacomys, though the roots do not close sa
early in life.

Fig. 18.— Side view of molars,
Evotomys. (a) youny:, ^&)
adult. (x3.)

FiQ. 19.— Enamel pattern
of molar toetli, Evotumyg
gapperi. (x5.)

44 NORTH AMERICAN FAUNA. [No. 12.

In the number and arranoement of triangles the enamel pattern
{fig. 19) is the same as that of the tetramerodont species of Microtus
(see p. Co). The salient angles are, however, for the most part rounded,
and so placed that the triangles are seldom fully closed.

External form. — In external form Evotomys does not differ essentially
from Microtus, although the ears are usually larger. The red or rufous
color of most of the species gives them a very different appearance
from the other voles.

General remarks. — The characters which separate Evotomys from Mi-
crotus and Phenacomys have been presented in such detail under the lat-
ter that it is unnecessary to consider them further. The peculiar bony
palate of Evotomys has been considered one of the best generic charac.
ters. Since the discovery that it is perfectly reproduced in two subgenera
of Microtus [Anteliomys and Eothenomys) it loses much of its importance.

The genus Evotomys is represented in Europe, Asia, and North
America by numerous species and subspecies whose interrelationships
are at present little understood. Among the American species may be
mentioned E. (japperi (Vigors), E. fuscodorsaUs Allen, E. galei Merriam,
E. idahoensis Merriam, E. californicus Merriam, and E. occidentalis Mer-
riam ; among those found in the Old World are E. rutilus (Pallas), E.
glareolns (Schreber), and E. rufocanns (Sundevall).

Geiinw MICROTUS Schrank.

1798. Microtus Schrank, Fauna Boica, I, Iste Abtb., p. 72, 1798. Typt- by elimination

Microtus terrestris Schrank = J/h8 arvalis Pall.
1883. Microtus Lataste, Le Naturaliste, Tome II, p. 348, 1883.
1801. Arvicola Lacdpede, Mem. de ITustitut, III, p. 489, 1801. Type ' Arvicola amplii-

hiu8'^=Miis terrestris Linn.

Geographic distribution of type species. — Central Europe and parts of
Asia.

Geographic distribution of genus. — In both hemispheres the genus
Microtus ranges from near the northern limit of mammalian life to the
edge of the tropics.

Essential characters :

Upper incisors without grooves.

Lower incisors with roots on outer side of molar series.

Molars rootless.

Enamel pattern characterized by approximate equality of reentrant angles.

m 1 usually with five closed or nearly closed triangles.

m 3 with one, two, or three closed triangles.

Tail nearly always longer than hind foot, terete.

Feet, fur, eyes, and ears very variable.

Thumb never with a well-developed ligulate nail.

Skull. — The skull of Microtus varies greatly in shape among the dif-
ferent subgenera. Eull descriptions will be given under each of these.
Considering the genus at large it is difficult to frame any diagnosis by
which the skull may be in every case distinguished from that of the
other voles. Most of the characters which at various times have been

Jl-LV, ISSIO.J

GENUS MICROTUS.

45

adult Microtus alleni. (x 2.)

brouyht forward for this purpose prove to be either wholly iueoustant
or coustant only when particular subgenera are held in view.

Tecih. — Although the skull of Microtus presents no tangible diag-
nostic characters, the teeth are readily distinguishable from those of
all other members of the subfamily. The upper incisors are never
grooved except in oc<;asional abnormal specimens. The root of the
lower incisor crosses the line of the molar
series between the second and third molars,
causing a greater displacement of the roots
of the latter (PI. Ill, fig. 3) than occurs in
anj'" other genus. It terminates in the ascend-
ing ramus of the mandible at a point slightly
above and behind the dental foramen (PI. Ill,
fig. 3). The molars, even in extreme old age,
are never rooted (fig. 20). This character
alone distinguishes them from the molars of
the other voles. The pattern of enamel folding-
varies considerably in the different subgenera, and forms one of the
numerous charactets by which the latter may be separated. Detailed
descriptions of the enamel i)atterus are given in the accounts of the
subgenera.

External form. — In external form the members of the genus jl/icro^Hs
vary excessively. Some resemble lemmings so closely that they have
been associated with these by certain writers. Others are modified for
an aquatic life and in consequence have more the appearance of musk-
rats {Fiber). Still others pass most of their time under
ground. In these the ears, eyes, and tail are reduced,
the front feet enlarged, and the fur so modified as to
suggest that of the moles. The great majority of spe-
cies, however, show none of these special adaptations,
but resemble in a general way the members of the genera
Phenacomys and Evotomys. Whatever may be the modi-
fications in form, the tail is almost invariably longer
than the hind foot and the thumb is armed with a small
or rudimentary i)ointed nail (fig, 21).
General remarks. — The characters of Microtus, as contrasted with
Evotomys and Phenacomys^ have already been given (p. 42) and need not
be repeated here.

-Subgenus EOTHENOMYS ^ Miller.
New subgeuus. Type Arvicola mc1a»o<jaster Milue-Edwards.

Geographic (UstriMition of type species. — Moupin, western Sechuen,
and western Fokien, China. (Blanford.)

Geographic distribution of subgenus. — Microtus melanogaster is the only
known species of Eothenomys, hence the geographic distribution of the
subgeuus is the same as that of the type species.

FlCJ.21.— Left front
foot, Microtxta
terrcstris.

^'H(b?, the morniug (eastern); Qev, from; yui??, mouse.

46 NORTH AMERICAN FAUNA. [No.i-.

Essential characters :

Palate abuormal.

m 3 without closed triangles.

m 1 witli triangles frequently open and 8 or 9 salient angles.

m 3 -with triangles usually open and G salient angles.

Mamma', (number not known).

Plantar tubercles, 5.

Sole hairy.

Claws on hind feet longest.

Fur apparently somewhat modified.

SJiidl. — In tlie spechneus of Eothenomys that I have examined the
skull is not in sufficiently good, condition to permit of auj^ detailed
description. The peculiar structure of the bony palate taken in con-
nection with. the teeth is, however, of itself enough to characterize the
group.

Bony palate. — Unfortunately in the two specimens of Microtus melano-
(jaster that I have seen (82.6.16.11 and 92.10.12.5, British Museum
Register) the basal part of the skull is so injured that the form of the
interpterygoid fossa can not be determined.
The bony palate, however, is sufficiently pre-
served to show the essential details of its
structure (PI. II, fig. 11). That part of the pal-
ate which lies in the level of the roof of the
riQ. 22— Enamel pattern of moutli cuds abruptly oppositc the front end of

molar teeth Microtu. (Eo. ^ ^ ^ j .^^ Straight-cdged Shclf

the7iomys)inclanogaster. (yi 5.) ii o e>

which extends without notch or projection from
alveolus to alveolus. Although the form is thus strikingly different
from that of the typical microtine palate, the vestiges of the structure
there i^resent may still be recognized. The lateral grooves and median
ridge are present, though slightly developed. The former terminate in
two depressions lying just in front of the Avide, flat, lateral bridges
which comjdetely obliterate the iDOsterior ends of the grooves, and
together with the terminal part of the median ridge form the edge of
the palatal shelf. The palate in all its essential characters is thus
exactly like that of Evotomys.

Enamel jyattern in general. — The enamel pattern in Eothenomys (ftg.
22) is in many ways remarkable. The triangles in all the teeth tend to
remain open, the points of the salient angles are blunt and rounded as
in Evotomys, the triangles on the outer and inner sides of the teeth are
subequal in size, and the maxillary teeth are especially noticeable for
their likeness to each other. The figures published by Blanford^ fail
to do justice to the teeth of this species. These are better represented
in Milne-Edwards's original iilate,^ in which there is also a hint at the
palate structure.

ijourn. Asiatic See. Bengal, L, pt. II, PI. II, fig. A.

2Recherches p. servir a I'histoire nat. d. Mammiferes, Vol. I, PL XLVI, figs. Ic
and Id.

July, 1896.] SUBGENUS ANTELIOMYS. 47

Front loicer molar. — The lirst lower molar Las the usual trausverse
posterior loop autl a moderately loug rounded anterior loop, with a
strong salient angle at each side of the base. It has five lateral tii-
angles, three on the inner side, two on the outer side. These may be
perfectly isolated, or more often Avidely open. Except for the greater
tendency to equality in the triangles, the teeth in the lower jaw do not
differ very greatly from the mandibular teeth of true Microtus.

Baclx upper molar. — The posterior maxillary tooth most nearly resem-
bles that of Fedomys. The anterior loop is followed by two lateral
triangles, subequal in size and more or less completely isolated from
each other and from the anterior loop. The third lateral triangle is
reduced to a strongly developed salient angle on the inner side of the
posterior trausverse looj). A second salient angle is formed on the
outer side of this loop, which thus appears as a crescent joined near
the middle of its coucavitj' to the rest of the tooth.

Other teeth. — The middle upper molar has a posterointernal loop
nearly as large as the postero- external loop, the two placed opposite
each other. The result is a tooth of practically the same shape as the
one behind it. The anterior upper molar is likewise provided with a
very large postero- internal loop opposite the loop on the outer side,
normally terminating the tooth. Tiius it very closely resembles the
two other maxillary teeth, differing only in its one more closed triangle
at the front end.

Mamma\ — The number of mammae in Eothenomys is unknown.

Feet. — The feet are moderately hairy, in this respect not differ-
ing from true Microtus. Blanford states that there are five well-
developed pads on the sole and a rudimentary sixth. The claws are
not greatly developed on any of the feet; those on the hind feet are the
longest.

Fur. — A skin in the British Museum has the fur of a iieculiar, dense,
mole-like quality suggestive of Pitymys. The specimen appears to be
in worn coat, however, and this character may not be normal.

General remarls. — Eotheaomya is such a Avell-jnarked subgenus that
it is surprising to find that it has hitherto received no name. In tooth
pattern it agrees iu a general way with Microtus siMimensiSj a circum-
stance which induced Blanford to place it in the subgenus '■Xeodon ;^
but the palate structure is widely different from that of the subgenus
Microtus, to which ^[. sil'lcijnensis I'eally belongs, while the similarity
in the enamel pattern of the two species is very superficial.

Snbgeuns ANTELIOMYS i Miller.
New subgenus. Type Microtus dtinensis Tlioiuas.

Geographic distribution of type species. — Microtus chinensis is known
from one specimen collected at Kiating-fu, west Sze-chuen, China.

' 'yJrr?)/\/o?, eastern; /iv?, mouse.

48 NORTH AMEEICAN FAUNA. [No. 12.

Geographic distribufion of subgenus. — Microtus chinensis is the ouly
known si^ecies of the subgenus.
Essential characters :

Palate abnormal.

m 3 without closed, triangles.

HI 1 with triangles mostly open, and with 9 salient angles.

Ill 3 with triangles mostly open, and with 9 salient angles.

Mamma', 4.

Plantar tubercles, 6.

Sole moderately hairy.

Claws on hind feet longest.

Fur not specially modiiied.

Skull. — As remarked by Mr. Tliomas in the original description of
Microtus chinensiSy the skull of Anteliomys resembles in a general way
that of Evotomys. Unfortunately, I am nnable to add any more deiiuite
information concerning its characters.

Bony palate. — The palate of Anteliomys (PI. II, tig. 8) is similar to that
of Eothenomys, except that the median ridge is produced backward as a
distinct spike lying perfectly in the plain of the roof of the mouth.

Just in front of the strongly developed lateral
bridges, the posterior edges of which form the
back rim of the bony palate, lie two pits, iu
which terminate the lateral grooves. These
pits communicate freely over (dorsad to) the
riG.23.— Enamel pattern of mo- lateral bridgcs with the anterior end of the

lar teeth, Microtus (Aiiteli- i i , • i x-

' . ' . , , , broad mesopterygoid lossa.

owyx) chmensis. (x 5.) ^ ••' ^

Enamel pattern in (/encral. — The enamel pat-
tern in Anteliomys (fig. 23) is characterized by rounded angles, imper-
fectly closed triangles, and great complexity in the prisms of the back
upper molar.

Front lower molar. — The anterior lower molar is made up of four
transverse, perfectly isolated loops. The anterior loop is much the
largest and contains three salient angles (two on the inner side). Each
of the succeeding looi)S has two salient angles. The tooth thus con-
tains exactly the same elements as the corresi)onding one in Microtus,
the difference in form being due to the fact that in Anteliomys the
prisms are placed ojiposito each other instead of alternately. The
Ijrisras on the opposite sides of the tooth are nearly equal in size,
thus producing the bilaterally symmetrical appearance found to a less
degree developed in Alticola and Eothenomys. The figures iu the
original description of Microtus chinensis^ give a very poor idea of the
teeth.

Back npper molar. — The posterior maxillary tooth is like that of true
Microtus except that the posterior loop is greatly lengthened and on
the lingual side cut by two reentrant angles, of which the anterior is

' Ann. & Mag. Nat. Hist., Ser. 6, Vol. VIII, p.- 118, August, 1891.

July, 180G.] SUBGENUS LAGURUS. 49

the deeper. TJiere is a salient augie at the outer base of the posterior
loop and the outer border is faintly crenulate. A tooth with nine well-
developed salient angles is the result.

Other teeth. — The front maxillary teeth are exactly as in tetranierodont
Microtus. The back molars of the lower jaw are likewise in no way
peculiar. Tliey both, however, have the prisms on the two sides oppo-
site, tlius lacking all ch)sed triangles.

There is nothing worthy of note in the form of the incisors.

Mamma'. — In the unique type specimen of Microtus chinensi-s, which is
a female, there are four teats, all inguinal.

Feet. — The sole is well haired from heel to tubercles. There are six
pads on tlie sole, all well develoi)ed.

Fur. — The fur is not specially modified.

General remarls. — In its i>alate structure AnteUomys is related to
Fothenomys, and more remotely to Alticola, together with Avhich it
bridges the gap (so far as the palate alone is concerned) between Micro-
tits and Erotomijs. Tliese facts were in part noticed by Mr. Thomas,
who says in the original account of il/. chinensis :

Iji some respects it seems to be .■uiuecteiit between Evotomijs and the rest of the
voles, the structure of its palate and some of its dental characters [opposite prisms
and rounded anj^les] showing striking affinities to the former, far as itsrootless teeth,
fewer mamma-, and difterent external form separate it from any of the known mem-
bers of that group.

The enamel r)attern is, however, very difterent from that of Frotomys,
while the resemblance to that of its nearest relative, FotJtenomys, is
almost equally remote.

Mirrotn.s chinensis is the only species of Anteliomys thus far known,
i\u](iSii 2Iicroti(s m?V/r7e«fZor/f/^" (Polyakofty from Siberia- proves to be
a member of the same group. The figure of the teeth in the original
description of M. mid (lend or ff it is suggestive of Microtus chinensis,
though the triangles are very strongly isolated. Neither the palate
structure nor the number of mammte is given by Polyakoff, so it is
impossible to come to any conclusion on the subject of the animal's
true status.

Subgenus LAGURUS Gloger.

1841. Lagurus Gloger, Gemeinu. Hand-u. Hilfsbuch d. Naturgesch., p. 97, 1841

(genus). Type, Lagurus viigratorius Gloger^ If hs lagurus Pallas f^
1895. Lagurus Merriam, Am, Naturalist, XXIX, p. 758, Aug., 1895 (subgenus).

' Mem. Acad. Imp. Sci., St. Petersbourg, XXXIX suppl., p. 70, 1881.

-Polyakoff gives the following localities: Taimur, Vilui River, Ayan, and Kara
River.

^In restoring the generic name Lagurus (Aim. & Mag. Nat. Hist., 6th ser., XV,
Feb. 1, 1895) Mr. Thomas gives the species lagurus as the type. It appears highly
probal)le, however, that Gloger's Lagurus migraiorius is the Llypudaus migratorius
of Lichtenstein (Eversmann's Reise nach Buchara, p. 123, 1^2^)^=^ Microtus {Lagurus)
luteus (Eversmaun).

16933— Xo, 12 4

50

NORTH AMERICAN FAUNA.

[No. 12.

1881. Eremiomiis Polvakoft', Mem. Acad. Imp. Sci., St. Petersbourg, XXXIX ^uppl.,p.
34, 1881 (genus). Type J/(ts latjnrns Pall.

Geographic distnhution of type species. — Plateaus of western and
central Asia.

GcoyrapMc distribution of suhgenua. — The range of the subgenus
Lagurus is very imperfectly known, but probably extends over a large
liart of the Boreal region in Asia and in western Xortli America.
Essential characters:

Palate slightly abnormal.

m 3 normallj' witli 2 or 3 tightly closed triaugles.

m 1 normally with 5 closed triangles and 8 or 9 salient angles.

m 3 normally with '2 or 3 closed triaugles and 5 or 6 salient angles.

Mamma', 8.

Plantar tubercles, 5.

Sole very hairy.

Claws on hind ieet longest.

Fur not specially modihed.

Slaill. — The skull of Lagurus (PI. I, fig. 7') may be at once recognized
by the form of the audital bulla? (fig. 24). These are larger than in any
other subgenus of Microtus, and are especially remarkable on account
of the way in which they project backward behind the x)lane of the

occiput. Aside from the audital bullae,
the skull does not differ very notice-
ably from that of I'iti/niys or Chilotus.
As compared with that of Pitymys,
however, tlie rostrnm is considerably
more slender. The dorsal outline is
flat, as in Chilotus.

Bony palate. — The bony palate (PI.
II, fig. 1') is normal in structure but
there is less diiference than usual between the levels of the portions
lying in front of and behind the lateral bridges. A peculiar flat palate
with shallow lateral pits and broad, ill-defined median sloping ridge
is the result. This form of j)alate is much like that of Fhenacomys
(PI. II, fig. 1).

Efiamel pattern in
general. — The enamel
pattern of Lagurus
(fig. 25) is character-
ized by the tight clos-
ure of all triangles,
notably in the back
lower molar, and the
great width of the reentrant angles

Fig. 24.— Audital buUsB, (a) Microtus {Mi
crotui) arvalis ; (6) M. (Lagurus) pallidus
(X 2.)

Fig. 25.— Enamel pattern of molar teeth : (a) Microtug (Lagurus)
pallidus; (b) 21. (L.) lagurus; (c) M. (L.) luteus. (x5.)

The latter peculiarity gives the

'See also Naturwissenschaftliche Resiiltate der von N. M. Przewalski unternom-
meneu Reiseu, PI. XIII, figs. 1, 2, 3, 12, 13, and 14.

July, 1896.] SUBGENUS LAGURUS. 51

teetb a drawn-out appearance, wliicli is liighly cliaracteristic. Wide
reeutraut angles occur in the teeth of the young- of all Microti; in
Laf/urus this embryonic character is retained by the adults.

Front lower molar. — The number of loops and triangles in the first
mandibular tooth is the same as in true Microtus. In Microtus prze-
icalslcii and Microtus Inteiis the anterior loop is simple and much
reduced, while in Microtus lagiirus and M. pallid us the loop is exactly
as in Microtus arvalis.

Bad- tipper molar.— The posterior maxillary tooth differs considerably
in form among the various species. In certain American species the
loops and angles are arranged exactly as in M. (Arvicola) terrestris,
while in M. przeicalslvH and M. lutens the tooth, although retaining the
same number of elements, is remarkably like that of some of the spe-
cies of Alticola. (See PI. XIII, Wissensch. Eesnltate der von K.
M, Przewalski nach Cent.-Asien nntern. Keisen. Zool. Theil, Ed. I,
Lief. 3.) This resemblance to Alticola results from the unusual elonga
tion of the i)osterior loop. In Microtus lagurus there are three tightly
closed triangles, and the terminal loop has a well developed salient
angle on each side at the base.

Other teeth. — In the Old World species (fig. 25) the back lower molar
contains four tightly closed triangles. The American species, however
(fig. 25), so far as known, have only three closed triangles in this tooth.
The other molars are always formed as in tetramerodont Microtus.
There is nothing peculiar about the incisors.

Mam)tia\ — In 2[icrotus pallidus, or a closely related form, there are
eight mamnipe, four pectoral and four inguinal. I have been able to
find no statement of the number of mamm;^ in the Asiatic species.

Feet. — Soles densely hairy as in Phaioinys and the lemmings; plantar
tubercles, five; claws moderately developed, those on hind feet longest.

Fur.-^ThQ fur is full and soft, but not highly modified. In color
most of the species are dull yellowish or grayish. The marking of
Microtus lagurus is unique in the genus Microtus on account of the
strongly developed and sharply defined dark dorsal streak.

(General remarhs. — The subgenus Lagurus is a strongly characterized
group, but, as Dr. Merriam has remarked,' the species show no pecul-
iarities to separate them generically from Microtus arvalis. In Microtus
lagurus, M. luteus, and M. przeicalslii, the tail is usually shorter than
the hind foot, thus adding to the superficial resemblance to the lem-
mings. No other voles have the tail so short.

The subgenus Lagurus is represented in the Old World by Microtus
lagurus (Pallas), 21. luteus (Eversmann), and M. przeicalsldi (Biichner).
In America there are probably numerous species and subspecies.
Among- these may be mentioned Microtus pauperrimus (Cooper), il7.
curtatus (Cope), and M. pallidus (Merriam).

'American Naturalist, XXIX, p. 758, August, 1895.

52 NORTH AMERICAN FAUNA. [Jfo.i2.

Subgenus ALTICOLA Blanford.

1884. AUicoIa Blanford, Jouru. Asiat. Soc. Bengal, L, Pt. II, p. 89, 1884. Type
Arvicola stolicskaiius Blanford.

Geographic distribuiion of type species. — " High plateaus of Northern
Ladali (Western Tibet)" (Blanford).

Oeographic disirihuiion of subgenus. — Boreal Zone in the Himalayas.
Essential characters :

Palate abnormal.

m 3 without dosed triangles.

m 1 with 4 or .5 closed triangles and 7 salient angles.

m 3 normally with 2 closed triangles and 5 or 6 salient angles; posterior loop
produced backward iu line of jaw.

Mamnut, §.

Plantar tubercles, 6.

Sole, hairy.

Claws on hind feet longest.

Fur long and soft but not highly modified.

Skull. — The skull in this subgenus (PI. I, fig. 10) shows no striking
peculiarities to distinguish it from that of trne Microtus. The general
shape is usually much as in Microtus arvalis,
but the zygomatic arches are more flaring and
the brain case is somewhat broader and flatter.
The rostrum is proportionally longer than iu
rio.26.-Enan,eipatternofmo- Microtus propcr, and the audital bnllaj (fig. 27)
lar teeth, Microtus {.AUicoia) are more inflated aud papery.
aibicauda {type), (x 5.) i5oH7/ ^;o /a ^6.— The median palatal ridge (PI.

II, fig. 4) widens at a point opposite the space between the second and
third molars and is approached, as in the typical microtine palate, by
outgrowths from the opposite sides of the lateral grooves. These out-
growths, however, do not meet the median ridge, but leave the lateral
grooves open. Just at its widest point the median ridge is squarely
truncated. The sloping terminal ridge is entirely lacking and the si)ace
that it usually occupies forms the anterior end of the very long rectang-
ular interpterygoid fossa. A structure of nuich the same appearance
could be produced by widening tlie anterior end of such a hastate
interpterygoid fossa as that often present in 'Aulacomys'' (PI. II, fig. 7)
until the whole space acquired an equal breadth. The floors and median
w^alls of the lateral pits would then be so encroached upon as to oblit-
erate the pits, while a few slight further modifications would give a
l^alate indistinguishable from that of AUicoia. The palate of Alticola
resembles that of JS^eofiber more closely than it does that of any other
subgenera excei)t Hyperacrius.

Enamel pattern in general. — The enamel pattern in Alticola (fig. 26)
differs in many ways from that of any subgenus of Microtus. In gen-
eral it is characterized bj^ {a) a tendency to reduction in the number of
prisms in the variable teeth; (&) by a peculiar irregularity and iiulefl-
niteness in outline j (c) by a strong tendency toward bilatei'al symmetry

July, 1896.] SUBGENUS ALTICOLA. 53

caused by tlie approximately equal size of tlie triangles on the opposite
sides of the teetli, and (d) by the form of the posterior upper molar.

While the figures published by Blanford ' in his paper on the voles of
the Himalayas, Tibet, and Afghanistan are in many ways inaccurate,
they give an excellent idea of the general appearance of the teeth in
the voles of this group.

Front lower molar. — The tirst mandibular molar has normally four
closed triangles and seven or eight salient angles. Earely a fifth closed
triangle is isolated at the inner basal angle of the anterior loop. The
form, relative position, and degree of isolation of the triangles and
transverse loops vary greatly witli the different species. Any one of
the reentrant enamel folds may fail to reach the enamel of the opposite
side, and consequently any of tlie triangles may be open at one or both
ends.

BacTi upper molar. — The posterior maxillary tooth varies in form in the
dift'erent species. It is, however, always recognizable by the backward
prolongation of the posterior loop in the line of the jaw, a character
which is found elsewhere in HyperacrhiSj OhilotuSy and Laguru.s only,
and in all but the first of these developed to a much less degree. This
attenuate posterior loop is followed by three or four more or less incom-
pletely isolated lateral triangles, these by an anterior loop of the usual
form. The tooth is most complex in M. roylei and i¥. hhmfordi, in each
of which it has six salient angles and two or three closed triangles.

Other teeth. — Except for the stronger tendency to bilateral symmetry
combined with slight irregularity of outline the other molars do not
differ from those of ordinary tetramerodont Microtiis.

Mamma\ — The number of mamnnt^ in the species of Alfivola has
apparently not been recorded. Blanford does not mention it in his
descriptions of any of the species, and Mr. G. E. H. Barrett-Hamilton,
who has made at my re<xuest a special examination of the material in
the British ^Museum, is able to add nothing on the subject. In an
adult nursing female of a sjiecies of Alt icola closely allied to Microtus
albicauda (Xo. 62162, U. S. Nat. ]Mus. Ladak side of Kara Korum
Pass, Kashmir) there are eight well-developed mamma?. Hence there
is little doubt that eight is the normal number in the subgenus.

Feet. — The feet are very hairy, the long hairs on the dorsal surface
often nearly concealing the claws. Plantar tubercles six. The claws
on all the feet are long and slender, those on the hind feet longer than
those in front.

Fur. — As in most high boreal micro tines the fur is long and full.
Otherwise it is not peculiar.

General remarls. — The subgenus Alt icola is one of the best character-
ized groups in the genus Microtus. The pattern of enamel folding is
uulike that of any of the other subgenera, except Hyperacrius, while
the palate structure is approached by that of Hyperacrius and the

' Jouru. Asiatic Soc, Bengal, L, Pt. II, PI. I, tigs. B, C, D, and E.

54 NORTH AMERICAN FAUNA. [No. 12.

otherwise widely difterent Xeojiher ouly. The tendency to bilateral

symmetry in the molars is sliared by three other Asiatic subgenera,

Hi/peracriufi, Uothcnomi/s, and AnteJloiny.s.

Alticola, like Ei/jieracriu.s, is apparently a strictly boreal subgenus.

The following species are known : Mkrotus stoliczlcanus Blanford, M.

roylii (Gray), .If. stracheyi (Thomas), Tlf. hlanfordi (Scully), and M.

albieauda (True).

Subgenus HYPERACRIUSi Miller.

New subgenus. Type Jrvicola fertHis True.

Geographic distrihuiUm of type species. — "Central Kashmir, the Pir
Panjal Range and the Kaj Nag Mountains." (True.)

Geoyraplilc distrlhiition of suhgenns. — Mountains of central and south-
western Kashmir at elevations ranging mostly from 7,000 to 12,000 feet.

JEssential characters :

Palate abnormal.

m 3 without closed triaugies.

Ml 1 noruially witli 4 or 5 closed triaugies aud 7 salient angles.

m 3 normally with 1 or 2 closed triangles and 4 salieut angles.

Mammji' 4.

Plantar tubercles 5.

Sole hairy.

Claws on hind feet longest.

Fur short and dense.

Skull. — The skull in the subgenus Byperacrlus (PL I, fig. 11) differs
from that of Alticola in its longer rostrum, strongly cuneate nasals,
narrower interorbital constriction, more abrui^tly flaring zygomata, and

flatter brain case. The whole dorsal
outline of the skull is depressed so
that the zygomata are more nearly on
the level with the top of the skull
than in any other subgenus oi Micro-
tus. The audital bullne (fig. 27) are

Fig. 27.— Audita] bullae, (a) Microhis {Alti- proportionally Smaller than in Alti-
cola) albieauda; (h) M. {Hinjeracnus} fer- 7 , ^r- , -jf, rri ,

tiiis (x2) cola, true lUicrotus, or rdymys. ihe

brain case is much more depressed
than in Microtus i)roper (flatter even than in Fitymys), and viewed from
above it has a peculiar subcircular outline not known elsewhere in the
genus. Parietals proi^ortionally smaller than in Microtus proper;
squamosals and interparietal proportionally larger. The latter in old
individuals has much the same shape as in fully adidt Arvicola,
Keojiher, aud Fiber.

Bony palate. — The bony palate is exactly as in Alticola.

Enamel pattern in general. — The enamel pattern (fig. 28) has the gen-
eral appearance of that of Alticola.

Front loicer molar. — The first mandibular tooth is indistinguishable
from the corresponding tooth iu Alticola.

* Oi vnepdnpioi, inhabitants of the heights.

July, 1800] SUBGENUS PEDOMYS. 55

Bad' upper molar. — The last maxillary tooth has the same general
form as that of Alt i col a, but is simpler iu structure, thus recalling
the corresponding tooth in Lcuiurus (flg. 25). There are usually only
two lateral triangles and four salient angles. The posterior loop is
lengthened in the axis of the jaw as in AliicoJa.

Ma mm (('.—There are four maramte, all inguinal.

Feet. — The feet are well haired, but rather less densely than in ^1/^/-
cola. Plantar tubercles five — the faintest possible trace of a sixth
sometimes present. Claws on all four feet well developed, those on
hind feet longest.

Fur. — The fur is much shorter and more dense than in Alticola.

MisceUaiicous characters. — The ears, and a])parently the eyes, also,
are smaller than in Alticola. The whiskers are very short, reaching
scarcely to the ears, while in Alticola they are probably longer than iu
any other subgenus of Microtus.

General remarls. — Hyperacrius is most closely related to Alticola, from
which it differs chiefly in its highly modified skull and reduced number
of footpads and mammjc. Minor differences are to be found in the rela-
tive size of the ears and in the character of the feet. Hyperacrius
appears to be modified for a more strictly underground life than Alticola.
It requires no close comparison with any other
subgenus, though it bears a superficial likeness
both in external form and in cranial characters to
Pitymys. The structure of the bony palate and
the pattern of enamel folding readily distinguish fig. 28.-Enamei pattern of
it from the latter, however. '""i-'^"' ^'^^'^i'' ^ierotus (Hy-

v\ hether Microtus irynnei may be associated
with Microtus fertil is in the subgenus Hyperacrius is a matter of doubt.
Atmy request Mr. G. E. 11. Barrett-Hamilton has examined the speci-
mens of AW/coZa in the British Museum with special reference to the
relationships of M. a-ynnei. He finds that this species, as already
noticed by Blanford, has only five plantar tubercles, but that in other
characters it does not agree with the brief diagnosis of Hyperacrius, that
I sent him. The fur is long, as iu the species of Alticola, and the skull
apparently lacks the peculiar form seen in Hyperacrius. The number of
mamm^cannot be determined in M. wynnei nor in any of the species
of Alticola in the British Museum. For the present it is not safe to
attempt to refer ilitcrofws ^cynnei definitely to one subgenus or the other.

Subgeuns PEDOMYS Baiid.
1857. Pedomys Baird, Mamm. N. A., p. 517, 1857. Type Jrvicola aiis1e7-us LeCoute.

Geographic distribution of type species. — Transition and Upper Austral
zones in the central United States and adjoining British Provinces.

Geographic (listribution of subgenus. — The range of this subgenus is
the same as that of Microtus austerus, the only known species.
Essential characters:
. Palate normal,
m 3 without closed triautfles.

56 NORTH AMERICAN FAUNA. [No. 12.

ui 1 normally witli 3 closed truiuglcs and 8 or 9 salient angles.

ni 3 normally with 2 closed triangles and 6 salient angles.

Mamma- 4.

Plantar tubercles 5.

Sole thickly haired between heel and tubercles.

Claws moderate in length, those on hind foot longest.

Fur not specially modified.

Sl'uIL — The skull of Microins austerus, the ouly kuown si^ecies of
Pedomys, is remarkable for the subcyliiidric brain case, and great depth
of all that part back of the rostrnm. While the skull of Pedomy.s is not
strikingly different from that of true Microtus,^ it is very unlike the
flattened skulls of Phaiomys, Pitymys, and CJiilohis, the other groups of
small voles resembliug* Pedomys in tooth cliaracters and in number of
mamma? and' footpads.

Bony palate. — The bony palate is typical, though the interpterygoid
fossa is seldom squarely truncate anteriorly.

Enamel pattern in general. — The enamel i)attern (fig. 29) is charac-
terized by simplification in the structure of the variable teeth.

Front loircr molar. — The first mandibular molar has a posterior trans-
verse loop followed by three closed triangles and an anterior loop.

The anterior loop is deeply indented by two
reentrant angles, one on each side. These
sometimes cut deep enough to isolate a fourth
or even a fifth closed triangle, but this rarely
takes i)lace. There is often a very faintly de-
FiG. 29.— Enamel pattern of mo- vclopcd reentrant angle close to each side of
lar teeth, Microtus (redomys) ^^^ ^{^-^ ^^^ ^^^^ autcrior loop. lu cascs wlicre

these are strongly marked a front tooth pre-
cisely resembling that of Microtns is the result.

Bacli upper molar. — The posterior maxillary tooth is exactly like that
of Neojiher^ Pitymys, Phalomys, Chilotus, and typical Arrlcola, havhig
an anterior transverse loop, two closed triangles and a short posterior
loop, from the outer base of which a third closed triangle may some-
times be cut.

Other teeth. — With the exception of the two teeth just described, the
dentition of Pedomys is like that of the tetranierodont species of the
subgenus Microtus.

Mam))ia'. — There are four mamm.e, all inguinal.

Feet. — Soles densely hairy between heel and tubercles; pads five,
with no indication of a rudimentary sixth.

General remarls. — Pedomys agrees in tooth pattern with Pitymys,
Chilotns, and Phalomys, but dilfers from all three in the shape of the
skull, and from the last in the short claws and unmodified fur also.

Subgenus PHAIOMYS P.lytli.

1863. Phaiomi/s Blyth, Jouru. Asiat. Soc. Bengal, XXXII, p. 89, 1863. Type I'liaiomys
leucttriis Plyth^l/^/cro^Ms blythl Blaudibrd.

'A skull of Microtns ratticeps from Norway exactly resembles skulls of M. austerus
except that the rostrum is more slender.

July, 1896.] SUBGENUS PHAIOxMYS. 57

1887. Lasiopodomys Lataste, Annali dtl Mvis. Civ. di Storia Naturale di Geuova, ser.
2a, Vol. IV, i>. 268, 1887. Type Arricola In-anti Radde.

Ge(i(ircq)hic distribution of type sjjecies. — -'Banks of Tslio Morari and
Pankoug lakes, Western Tibet, also between Seh and the PankongLake
at elevations above 13,0C0 feet." (Blanford.)

GeiKirapltic distyihutioii of sulxjcnus. — High i)lateau region of central
and southern Asia. Probably does not occur below the Boreal zone.

Esse ntial cha racters :

Palate normal.

m 3 Avitliout closed triangles.

m 1 normally with 3 to 5 closed triangles and 8 or 9 salient angles.

m 3 normally with 2 to 3 closed triangles and 6 salient angles.

Mamma' j^robably 10.

Plantar tubercles, 6.

Sole very hairy.

Claws very long and of abont 0(jual length on all four feet.

Fur remarkal)ly long and soft.

*S'A-»7/.— Tlie skull of Phaiomys as compared with tliat of Pedomys is
readily distinguished by its very different form. The brain case in
Pedowys is higb, long, and almost cylindrical, while that of Phaiomys
is short, broad, and flat. The zygomatic arches are more broadly flar-
ing in Phaiomys than in Pedomys, while the
upper incisors are usually more promiuent.
The latter character is, however, inconstant.

Bony palate. — The bony palate is perfectly
normal and requires no detailed description.

Enamel pattern in yeneral. — The enamel ]iat- i^k;. 30.— Enamel pattern of
tern (fig. 30) is exactly like that of Pedomys, moi^r teeth, Microtus (Phai-

J. XI i. . 1 . " . . 1 • — .> omi/s) strauchi. (x 5.)

except that the outer reentrant angles m m 3

are somewhatlessdeveloped, while the anterior outer reentrant angle in
m 2 usually divides tlie anterior loop into two closed triangles. These
dijferences, however, are trivial and inconstant.

Other teetli. — In some of tbe members of the subgenus the incisors
are directed more forward than usual. The character is, as already
stated, wholly inconstant.

Mamma'. — There is still doubt as to the normal number of mamma?
in the subgenus Phaiomys. Milne-Edwards found only four in a skin
of .1/. mandrianus ; Biichner found six in a skin of M. straxchi, and ten
in a skin of M./uscus. I am inclined to think that ten will prove to be
the correct number.^ In the specimen of M. fuscns just referred to
there were six pectoral mamma*, the rest inguinal.

Feti. — The feet are large and densely haired. The number of tuber-
cles on the sole is still a matter of doubt. Biichner records six in both

* That Phaiomys probably has a large number of mammae — at least more than four —
■was suspected by Lataste, who in 1887 (Annali del Museo Civico di Storia Xatnrale
di Genova, Serie 2a, Vol. IV, p. 270) called attention to the fact that Blyth found
ten embryos in a female Microtiis blyihi.

58 NORTH AMERICAN FAUNA. [No. 12.

31. hramlti and M. strauchi, but I am able to find only five iu a skin of
the latter, even after tliorouglily relaxing the foot. It is probable that
six is tlie real number, as Bilchner's determinations were made from
alcoholic vspecimens. The claws on all four feet are large and about
e(]ual in length. That on the thumb is well develoj)ed — in this resi)ect
perhaps surpassing all other subgenera of Microtiis.

Fur. — The fur is long and soft, suggesting that of a lemming rather
than that of a vole.

General remarls. — In many respects Fhaiomys resembles Fedomys so
closely that I should hesitate to sei)arate the two groups were they not
already named. There are, however, such differences between them
that it is impossible to call them the same, while in all probability more
satisfactory material than that now available would show additional
characters. In external ai)pearance the two subgenera differ consider-
ably. While Fedomys is a typical vole, Fhaiomys bears a general resem-
blance to the lemmings. The peculiar aspect of the species of Fhaiomys
is caused by their short tails, large feet, and long, soft fur. The like-
ness between the species of Fhaiomys and the yellowish species of the
subgenus Lagurus is even more striking. From the latter, however,
they are readily separable by dental characters.

Mlcrotus hlythi (Blanford), M. mandarinus (Milne-Edwards), M. strauchi
Biichner, Jlf. /?/,s'c?/s (Biichner), and M. hrandti (Iladde), are perhaps the
best-known species of the subgenus Fhaiomys.

Subgeuns PITYMY8 Mclvlurtrie.

1830. J'sammomys LeCoute, Ann. Lye. Nat. Hist., New York, HI, p. 132, 1830 (genus).

Type rsammomijs pinetorum Le Conte (uoc rsammomys Cretzsclimar 1828).

1831. Pltymijs McMurtrio, American edition, Cuvier Ri-gne Animal, I, p. 434, 1831

(genus). Type rsammomys pinetorum LeCoute.
1857. rUymys Bainl, jMamm. N. Km., p. 517, 1857 (section;
1887. Tilymys Latasto, Aunali del Mus. Civ. di Storia Naturale di Cenova, serie2rt, IV,

p. 266, 1887 (subgenus).

1831. Ammomys Bonaparte, Saggio Distrib. Metod. degli Auim. "\'ert., p. 20, footnote,

1831 (genus). Tj'pe Psammomys 2>i»etornm Le Conte.
1836. Finemys Lesson, Hist. Nat. d. Mamm. et Ois d(^coiiv. depuis 1788, Compl.

Oouvres de Bufl'ou, Y, p. 436, 1836 (genus). Type rsammomys piuetorum

LeConte.
1867. TerricoJa Fatio, Les Canipaguols du liassin du Leman, ji. 36, 1867 (subgenus)

{sul)terranens and savii).
1876. Micrunis Forsyth Major, Atti dclla Societa Toscana di Sci. Nat., Ill, fasc. I,

p. 126, 1876 (subgenus). Type Arricola nehrodensrs Mina Palumbo.

Geoyraphic distribution of type s2)eeies. — Austral Zone in the eastern
United States.

Geographic distribution of subgenus. — Central and southern Europe,
eastern United States, i)arts of Mexico.
Essential characters :
Palate, normal,
m 3 without closed triangles,
m 1 uormallv with 5 closed triangles and 9 salient angles.

July, 189(3.] SUBGENUS PITYMYS. 59

111 3 normally with 2 or B closed triangles and 6 salient angles.

Maninne, 4.

Plantar tubercles, 5.

Sole moderately liairy.

Claws on front feet longest.

Fur short, dense, and mole like.

Sk-ulL — The skulls of the species of Pitj/mys differ considerably among
themselves. In Microtus i)metorum (PI. I, fig. 2), the most highly modi-
fied, the brain case is very broad and flat and the interorbital region is
remarkably wide. The brain case is like that of Lagurus, but the broad
anterior x^art of the skull is very different from the latter. The dorsal
outline is strongly arched, esj)ecially anteriorly from the region between
the orbits to the tips of the nasals. The arching is, however, no more
strongly marked than in Microtus arvaUs. In Microtua suhUrraneus
the skull is like that of 2[. pinetorum^ but the i^eculiarities are less
accentuated. In the Mexican species of Fiiymys the brain case is
narrower and higher than in J/, pinctorum, and the anterior part of
the skull is less heavily built. The zygomatic
processes of the inaxilhe stand out more
nearly at right angles with the side of the
skull, thus ])ringiug the broadest part of the
zygomatic arch farther forward than in 21.
pinetorum.

Bony pahde, — The palate is normal, though
the region between the posterior molars is in
M. pinetorum rather flatter than. usual in true -pm. si.— Enamel pattern of
Microtus, and the anterior outline of the inter- ^"^^^ teeth, (a) Jiicrotus
pterygoid tossa is otten somewhat hastate. j^ {P)savii (x5 )

Enamel pattern in general. — With the excep-
tion of the front lower molar and back upi:)er molar, the enamel ])attern
(fig. 31) is that of tetramerodont Microtus.

Front lower molar. — The anterior mandibular tooth contains the same
number of looj)s and angles as the corresponding tooth in Microtus
arralls. As a rule, however, the first and second triangles are not
comx)letely isolated from each other or from the anterior loov). The
tooth is therefore exactly as in Fedomys.

Back upper molar. — The i)osterior maxillary tooth is simplest in the
American species of the subgenus. In these it is like the back upper
tooth in Fedomys and Arvlcola, which contain two closed triangles
and an anterior and posterior loop. In M. subterranetis, however, the
tooth is formed exactly as in M. arralis, while in M. saril it is some-
what intermediate. In the last-named species the terminal loop is
slightly larger than in M. pinetorum^ and a third closed triangle is
usually cut oft' from the outer base.

Other teeth. — There is nothing peculiar about the incisors or remain-
ing molars.

Mamma'. — In Fitymys there are only four nuimmte — all inguinal.

60 NORTH AMERICAN FAUNA. [No. 12.

Feet. — The soles are moderately hairy. They have five well-devel-
oped tubercles, l)ut no trace of a sixth. The claws are well developed
on all the feet, those on the front feet either equaling or exceeding
those on the hind feet.

In M. pinetorum the front feet are much larger and the front legs
shorter than in true Microfus. These peculiarities are less developed
in M. suhtemmeus and M. savii. Of the other species 1 have not seen
alcoholic specimens, and so am unable to say which of those mentioned
they most closely resemble.

Fur. — The fur in all the Icnown species is remarkably short and dense.
This character is most noticeable in M. pinetorion, which has an almost
mole-like coat.

Miscellaneous eharacters. — The tail, eyes, and external ears are much
reduced in all the species of nti/mys. These characters, as well as the
peculiarities of the fur and front feet, are distinctly adaptive and fit
the animals for their underground life.

General remarks. — While Pitijinys agrees with P<;^Z(:>^>?_(/s in the number
of mammie and footpads, it is readily distinguished by its highly mod-
ified fur, small eyes and ears, and flattened skull. The type and most
extremely developed species is further characterized by its greatly
shortened front legs.

Pitymys is represented in Anierica by Microtus ijinetoruin (Le Conte)
and several forms related to 21. qiiaxlater (Ooues). In Europe a num-
ber of species and subspecies occur. Among these the best known
are M. suhterr<iHeus (De Selys Longchamps) and J/, savii (De Selys
Longchamps).

Subgenus CHILOTIIS Bairtl.

1857. Chilotu-s Baird, Mamni. X. Am., j). 510, 1857. Type, Arvicola oregoni Baclimau.

Geographic distribution of type species. — Oregon, Washington, aud
British Columbia.

Geographic distrihution of suhf/enus. — The range of the subgenus
Chilotus is coincident with that of the tyjie aud only known species.

Fssential characters :

Palate noimal.

m 3 normally without closed triangles.

m 1 with 5 closed triangles and 9 or 10 salient angles.

m 3 with 2 or 3 closed triangles and 6 salient angles.

Mamm;e 8.

Plantar tubercles 5.

Sole moderately hairy.

Claws on hind feet longest.

Fur short and dense.

Slnlh— The skull of Chilotus (PI. I, fig. 8) is low and flat, the dorsal
outline nearly straight, and the brain case not widened, as in Fedomys.
As compared with Fedomys, the rostrum is remarkably long and slender
in proportion to the rest of the skull.

jrLv,i89C.l SUBGENUS CHILOTUS. 61

Bony pen ate. — The palate is normal and calls for no further remark.

Enamel pattern in general. — The enamel folding (flg. 32) is like that
of the tetramerodont species of Microtns, except that the back upper
tooth is a little simplified.

Front lower molar. — The first mandibular molar is exactly like that
of typical Microtus.

BacJc upper molar. — The back maxillary tooth contains a transverse
anterior loop, two lateral closed triangles, and a somewhat lengthened
terminal loop. The latter has at each side of its base a conspicuous
augle, the outer one of which is often isolated as a third closed triangle.
The tooth has six salient angles, two to each of the transverse looi)S
and one to each of the closed triangles.

Other teeth. — As already stated, the remaining teeth are formed
exactly as in tetramerodont Microtus. One specimen from British
Columbia has the lateral triangles closed in the back lower molar.

2ramma\ — There are eight mamune, four pectoral and four inguinal.

Feet. — Soles moderateh^ hairy from heel to tubercles; plantar tuber-
cles five, all "well developed; claws on hind feet
longest ; front feet not modified like those of typical
Pitnmy.s.

Fur. — The fur is shorter and more dense than
iu true Mi c rot us, hwt the modification is not car- na. 32.— Enamel pattern of
ried so far as in Microtns {Pitymys) pinetornm. mov.ivte^th. Microtis icu-

General remarls. — Chilotus combines the mam- " us) 01 egom. (x .)
mic and foot pads of Arricola with the nearly typical enamel iiattern of
Mierotus and has a form of skull peculiarly its own. In general it is
modified in the same direction as Fitymys, but to a much less degree.

Great stress has been laid on the form of the ear as a character of
this subgenus. In the original description^ Baird says:

A speciiiK'U in alcohol, from Steilacoom, received since the preceding descriiitiou
■was prepared, is, in size, niucli as described, The ears are low, orbicular, the mem-
brane thickened, the margins or conchal portion much inflected or incurved, like a
half-open apple blossom, the concha being inflected all round. The antitragus is
well developed, but rather low. The surfaces of the ear appear perfectly naked,
with, however, a ciliatiou of long hairs toward the roots of the concha, on the dorsal
surface. A close examination of the auricle in the dried specimen shows a i'ew scat-
tered, very short, white hairs.

• The structure of the ear, though in many respects similar to that of A. innetorum,
is yet essentially different. Thus the upper and lower roots of tht; margin of the ear

' meet auteriorlj' so as to form even a low rim to the meatus anteriorly, completely
Inclosing the aperture ; the edge of the concha is inflected ; the region inside the
auricle, around the meatus, naked, and the antitragus so much develoijed as to be
capable of completely closing the meatus. In A. pinetorum the roots of the upper
and lower margins of the ear are widely separated, by a space of a quarter of an
inch, the space between these roots and anterior to the meatus perfectly plane ; the
edges of the concha, or of the auricle, not inflected at all ; the inner space around
the meatus partly hairy ; the antitragus A-ery slightly developed, not valvular, nor
capable of closing the meatus at all.

' Mamm. X. Am., p. 538, 1857.

62 NORTH AMERICAN FAUNA. [X0.12.

Through Mr. True's kinduess I have been able to examine one of the
alcoholic specimens on which Baird based this description. This speci-
men (No. 2533, from Tomales Bay, Cal.') is in good condition and shows
most of the peculiarities to which attention was called. The thicken-
ing of the edge of the auricle is, however, due to disease or to the
action of the parasites which often attack the rims of the ears in the
voles and other small rodents. The anterior base of the ear is not essen-
tially different from the same region iu Plfymys, though the valvular
fold is slightly more developed. It is ])robable that by means of this
fold the meatus in Fitynn/s, as well as in most if not all of the voles,
can be tightly clnsed.

Subgeuus MICROTUS Schraiik.

1798. ^icroms'Sclirauk, Fanua Boica, I, Iste Abtli., p. 72, 1798. Type l>y i-liminatiou

Microtus terresfris Schrauk ^ Mhs arralis Pall.
1817. Mynomes Rafinesque, Am. Monthly Magazine, II, p. 4.5, 1817. Type Mynomes

praiensis Raf. ^^ Arvicola pcnns)jlvaiiicus Ord.
1836. Hemiofomifs DeSolys Loiigchamps, Essai Monographique sur les Caiupagnols

(les environs de Liege, p. 7, 1836, part (included arralis and ierrestris).
1857. Remiotomi/s Baird, Mamra. N. Am., p. 515, 1857.
1819. Keodon Hodgson. Ann. and Mag. Nat. Hist., 2d ser., Ill, p. 203, 1849. Type

Neodon sikkimensis Hodgson.
1857. Paludicola Blasius, Fauna der Wirbelthiere Deutschlauds, I, p. 3.33, 1857, part

(included ierrestris, niralis, and rafficej)s).
1857. AgrieoJa Blasius, Fauna der AVirbeltliiere Deutsclilands, I. ]). 334, 18.57. Type

Arricola agrestis.
1867. Praticola Fatio, Les Campaguols du Bassin du L^mau, p. 36, 1867, part (included

ierrestris, nivalis, arvalis, raiiiceps, and eampestris).
1867. Sylvicola Fatio, Les Campaguols du Bassin du L^man, p. 63, 1867. Tyjje Jrfi-

cola agrestis.
1890. Camjncola Schulze, Schriften Naturwiss. Vereins d. Harzes in Wernigerode, V,

p. 24, 1890, part (included arvalis, snhterraneiis, and eampestris).
1894. Tetramerodon Rboads, Proc. Acad. Nat. Sci. Pliila., p. 282, 1894. Type Arricola

tetramerus Rlioads.

Geographic distrihution of type species. — Central Europe.
Geoiirapliic distribution of suhyenus. — Boreal region of both hemi-
spheres, south to Mexico, northern India, and southern Europe.
Essential characters :

Palate normal.

m 3 without closed triangles.

m 1 normally with 5 closed triangles and 9 salient angles.

m 3 normally with 3 closed triangles and 7 or 8 salient angles.

Mamnuc, 8.

Plantar tubercles, 6.

Sole moderately hairy.

Claws of hind feet longest.

Fur not specially modified.

Sladl. — In true Microtus (PI. I, fig. 3) the skull lacks the peculiar
modifications found in such subgenera as Layiirus, Fitymys, Chilotus,

' No. 2529 from Steilacoom, Wash., also mentioned by Baird, is lost.

July, 1806.]

SUBGENUS MICROTUS.

63

and others. Withiu certain limits, liowever, tlie skull varies consider-
ably in size and form, so that it is difficult to frame any accurate diag-
nosis. The skull of Microtus arralis figured on Plate I represents the
form characteristic of the great majoritj of species. One of the most
notable departures from this type is seen in the skull of Microtus nitvdis,
which has an unusually low, broad brain case, and flat dorsal outline.

Bony palate. — The bony palate in the subgenus Microtus (fig. 7 A, and
PI. II, fig. '■>) shows in its most perfect development the form which may
be considered the normal one in the genus, since it is characteristic of
most of the subgenera and of the vast majority of species. As this
palate has already been described (pp. 2G-27) it is necessary here to notice
a few departures from the type form only. In young individuals the
sloping ridge is broader than in the adults, while in very old individuals
it often becomes very abrupt and at the same time greatly narrowed.
Tliese two extremes, Avhich are usually characteristic of immaturity and
old age, occur as the normal condition in the adults of certain species.
In Microtus nivalis the ridge is broad and fiat, while in M. agrestis, M.
ratticeps, and most of
the American species
it is narrow and ab-
rupt. Occasionally (es-
pecially in M. ayrestis
and M. ratticeps) the
anterior edge of the
interpterygoid fossa is
encroached upon bj"
the projecting median
ridge. The latter, on
the other hand, may be slightly cleft in the median line, thus fore-
shadowing the first step in the series of changes which lead to the very
different palate of Evotomys.

Enamel pattern in f/eneral. — The enamel pattern in the subgenus
Microtus (fig. 3.j) is characterized by the large number of loops and
angles in the first lower molar and last upper molar.

Front hncer molar. — The first lower molar normally contains a pos-
terior transv^erse loop, five closed triangles, two of which are on the outer
side and three on the inner side, and finally an anterior loop which is
usually more or less deeply cut by two reentrant angles, one on each
side of the loop, the outer of which is always the more posterior of the
two. With these loops and triangles are usually associated nine well-
developed salient angles, two formed by the posterior transverse loop,
one by each of the five closed triangles, and one by each side of the
base of the anterior loop. That part of the anterior loop which lies in
front of the reentrant angles may develop a salient angle on its inner
side, less frequently one on the outer side. Very rarely the Ioojd may
be cut by a third reentrant angle. This condition occurs in adult spec-

FlG. 33. — Enamel pattern of molar teeth, (a) Microtus {Microtus)
arvalis ; {b) 3r. {M.) nivalis : (c) M. {M.) pennsijlvanictis ; (d) M.
(2[.) ratticeps. (xS.)

64 NORTH AMERICAN FAUNA. [Xo.:2.

imens of Microtus agrestis, M. pennsylvanicus, also in the type of M.
(Ariucola) arvieoloidcs {f\g. 35), and probably in any other species with
the tootli formed after tLe pattern of Microtus arvalis. The other varia-
tions in the form of the front lower molar are the result of the greater
or less development of the reentrant angles normally present at the
anterior end. Sometimes the fourth reentrant angle (counting from
behind) on the lingual side of the tooth fails to meet the third on the
opposite side. Very rarely the anterior outer triangle opens in a like
manner into the anterior inner triangle, and the latter at the same time
communicates with the anterior loop, thus producing a tooth like that
normally present in Pedomys and I'itymys. liather frequently a sixth
closed triangle is cut off from the outer basal corner of the anterior loop,
and occasionally a seventh triangle is isolated at the inner side of the
greatly reduced loop.

The variations just described are purely individual and occur in the
species having the tooth of the typical form. Two notable variations
from this form are normally found in Microtus ratticcps and M. nivalis.
In the former (fig. 33f?) the fifth triangle opens into the short, unindeuted
anterior loop. There is here an actual reduction in the elements of the
ooth, which has only eight salient angles, thus resembling the corre-
sponding tooth in Fedomys. In M. nivalis (fig. 33t), while there are five
closed triangles and nine salient angles, the anterior loop is small and
crescentic, much resembling the posterior loop in the maxillary teeth
of Eothenomys.

Bad' upper molar. — The last upper molar is noiinally made up as
follows : An anterior transverse loop, succeeded by three closed trian-
gles, two smaller ones on the outer side and a larger one on the inner
side, these in turn by a posterior loop of variable shape. The tooth
usually contains seven salient angles, two to each of the transverse
loops and one to each of the three closed triangles.

Variations in the form of this tooth are numerous. Beginning at the
anterior end where the structure is most definite, it is found that the
first outer triangle very frequently opens into the large inner trian-
gle, less often into the anterior loop. The second outer triangle very
rarely opens into the inner triangle, but is rather frequently in commu-
nication with the posterior loop. The i>osterior loop varies in form and
size, the variations being partly individual and partly characteristic of
species. For the present it is unnecessary to discriminate in all cases
between the two categories. The most usual form and that found in
the type species, Microtus arralis (fig. 33a) is an irregular crescent with
the concavity directed inward and backward and the j)osterior tip thick-
ened, the whole joined to the rest of the tooth nt a point on the con-
vexity midway between the middle and the anterior extremity. This
nearly crescentic form is usually distorted by the elongation and straight-
ening of the anterior limb, so that the resulting shape is more like that
of the letter J. The thickened posterior extremity of the loop is often

JvLv, 1^'jf,] SUBGENUS MICROTUS 65

extended and cut by a reentrant angle on the lingual side, so that the
crescent is modified into the form of a rude E. Occasionally the ante-
rior extremity of the crescent is isolated as a second inner triangle.
The convex side of the crescent may develop a more or less prominent
salient angle. This condition is normal in Microtus ratiiceiys and Micro-
txs dtrotorrhiniis, but occurs also in other species. In the aberrant
Microtus nivalis the structure of this tooth is simplified so that it is
essentially as in Arvicola, Pedomys^ and Fitymi/s.

Other teeth. — The first and second upper molars contain each an ante-
rior transverse loop and, respectively, three and t^YO closed triangles.
In Microtus agrestis, M. sihlimcnsls, M. pennsylrcmicus, M. tcrra'novce,
and .1/. aztecus the inner edge of m2 is i)roduced into a conspicuous
loop, which frequently becomes isolated, so as to form a closed triangle
about half the size of the others. The European species with m2
formed in this way have been placed in a subgenus called Agricola or
SylricoJa, while the American species have been referred to Mynomcs
in a restricted sense. The American species with m 2 exactly as in
2Iicrotus arvalis have received the name Tetrameroclon. While the
name Tctramerodon can not be used in a subgeneric sense, it is fre-
quently convenient to speak of the voles with the enamel pattern of
M. (irvaJis as the tetramerodont species to distinguish them from their
pentamerodont allies. In Microtus sillimensis a supplemental triangle
is developed in m 1 as well as in m 2. On account of this peculiarity the
animal has been made the type of the genus or subgenus '■Xeodon.^
Neither Xeodon nor Agricola are worthy of recognition as subgenera
distinct from Microtus. Their characters are of trifling importance,
while in other species of Microtus (as, for instance, M. nivalis, M. guen-
theri, and occasionally .1/. pennsylvanicus) intermediate conditions can
be found.

Mamma'. — In the subgenus Microtus the mammaj are always eight,
four pectoral and four inguinal. Xo exceptions to this number are
known.

Feet. — There are six turbercles on the sole. Five of these are always
well developed, but the sixth is variable in size, being especially large
in M. ratticeps. The sole is always moderately hairy from heel to
tubercles. It is never densely furred as in Vhaiomys or naked as in
Xeojioer. The claws on all four feet are moderately developed, those
on the hind feet always slightly larger than those on the front feet,
the latter never specially developed for digging (cf. Pitymys).

Fur. — The fur is moderately full and soft, neither long and silky as
in Fhaiomys nor dense and mole-like as in Pitymys.

General remarks. — The subgenus Microtus needs comparison with the
groups having normal or very slightly abnormal palates: Arvicola,
Pedomys, Pitymys, Chilotus, Fhaiomys, and Lagurus. From all the
others it differs too widely to give rise to confusion. Lagurus is dis-
tinguished from Microtus by the tightly closed triangles in the posterior
16933— Xo. 12 5

66 NORTH AMERICAN FAUNA. [No. 12.

mandibular tooth, Arvicola by tlie presence of large musk glands on
tLe sides, Fedomys and IHtymys by reduction in the numbers of both
mamma' and jdantar tubercles, Chilotiis by reduction in the latter only,
and I'haloniys by an increase in the number of mammte and by the
very large claws. More extended comparisons will be found under
each of these subgenera.

This subgenus is the most widely and generally distributed, as well
as the one containing the largest number of si^ecies. Although the
si)ecies of MicrotifKe are still very imperfectly known, there is little
doubt that the members of the subgenus Mierotus greatly outnumber
the species of all the other genera and subgenera together. Conspicu-
ous representatives of the subgenus Mierotus are (in the Old World) :
Microtvs arvalis (Pall.), M. a[/restis (Pall.), 71/. raUlceps (Keys. & Bias.),
M. nivalu (Martins), 71/. {/uentheri (Dansford & Alston), 71i. silJcimensis
(Hodgson) 5 (in America) : Ilicrotus pennsylvanicus (Ord), 71/. terrwnovce
(Bangs), 71/. .rantliOf/natJnt.s (Leach), 7li. ehrotorrhinus (Miller), 71/ Jongi-
cauda (Merriam), 71/. mor/oUo^iensis (Mearns), 71/ townsendi (Bachmau).

Subgenus ARVICOLA Laceiii-de.

1801. ArritoJa Lact_^pecle, Mem. de I'Institut, Paris, III, p. 489, 1801 (genus). Type,

'Arvicola anqjhibius^ = Mus terrestris Linn.
1883. Arvicola Lataste, Le Natnraliste, Tome, II, p. 349, 1883 (subgenus).
1836. Hemiotomys De S^lys Longchamps, Essai Mouograpbique sur les Campagnols

des environs de Liege, p. 7, 1836, part (included arvalis and ierresiris).
1857. Paludicola Blasius, Fauna der Wirbeltbiere Deutscblands, I, p. 333, 1857, part

(included terrestris, niralis, and ra if icejis).
1867. Oclietomys Fitzinger, Sitzungsber. K. Akad. Wiss. Wien, LVI, p. 47, 1867. (No

type mentioned, but genus intended to include all the water rats of Europe.)
1867. PraticoJa Fatio, Les Campagnols du Bassin du Leman, p. 36, 1867, part (included

terrestris, nivalis, arrnlis, ratiiceps, and caynpestris).
1894. Aulaconuis Rhoads, American Naturalist, XXVIII, p. 182, 1894. Type, Aitlacomys

arvicoloides Rhoads.

GcograpMc distribution of type species. — Northern Euroi^e.
Geographic distribution of subgenus. — Noi'thern x^ai't of Xorthern
Hemisphere, exclusive of America east of the Eocky Mountains.
Essential cliaracters :

Palate slightly abnormal.

m 3 occasionally with closed triangles.

m 1 normally with 3 to 5 closed triangles and 7 to 9 salient angles.

m 3 normally with 2 or 3 closed triangles and 6 to 8 salient angles.

Mamma^ 8.

Plantar tubercles 5.

Sole almost naked.

Claws on hind feet longest.

Fur slightly modified.

Musk glands iiresent on sides of body.

STculI. — The skull of the larger Old World species of Arvicola (PI. I,
fig. 9) is nearly as large as that of Neofiber. In the American species

July, 1896.

SUBGENUS ARVICOLA.

67

(PI. I, tig'. 1) it is smaller, though considerably larger than in most
species of jl7/cro/ Mi' proper. Aside from its large size and prominent
ridges, the skull of Arvicola differs from that of Microtus in its broader,
shorter brain case, more widely flaring zygomatic arches, and propor-
tionally slender rostrum. The peculiar appearance of the rostrum is
heightened by the fact that the incisors project more than usual. Some
of these characters are more noticeable in the American species, though
the latter show no cranial peculiarities of sufficient importance to sep-
arate them subgenerically from those of the Old World. In the Amer-
ican species the skull is usually more lightly built and less strongly
angular than in the typical members of the genus (compare figs. 1 and
Oofri. I).

Bonii palate. — The bony palate is usually normal, but ocasionally
the median sloping ridge is divided in the median line, so that the
iuterpterygoid fossa is hastate anteriorly (PI. Ill, fig. 7). This condi-
tion occurs most frequently in the Ameri-
can species, but even among these it is
inconstant.

Enamel pattern in (jeneral. — The enamel
pattern in iy\)ic,2i\ Arvicola (tig. 34Z>) is char-
acterized by the great reduction in the
number of closed triangles and salient
angles in the front lower molar and
back upper molar. In these peculiarities,
though closely approached by Pitymys,
Pedomys, and Phaiomys, it presents the
extreme conditions found in the genus.
The third lower molar shows the tendency
to closure of the lateral triangles charac-
teristic of all the larger members of the genus. The pattern of enamel
folding in the molar teeth of the American species of Arvicola (fig.olrt)
is, on the other hand, exactly like that of the tetramerodont species of
the subgenus Microtus (e. g., Microtus arvalis and most of the western
American species).

Front lower molar. — In the typical species the simplification in the
structure of the teeth is carried furthest in the first lower molar. This
tooth normally contains a i)osterior transverse loop followed by three
closed triangles (one on the outer side, two on the inner side) and a
terminal transverse loop which is deeply constricted in the middle.
Each transverse loop forms two salient angles and each lateral triangle
one, making seven in all. Deviations from this form are very rare. In
one or two specimens I have seen a fourth triangle isolated on the outer
side, thus producing a tooth much like the corresponding one in Micro-
tus {2Ilcrotus) ratticeps, a species which has the last upper molar very
complicated in structure. The front lower molar in typical Arvicola
differs from that of the other groups in which it has only three closed
triangles in the reduced number of salient angles — seven instead of

Fig. 34.— Enamel patteru of molar teeth,
(a) Microtun (Arvicola) macropus ;
(h) M. (A.) terrestris. (x5.)

68 NORTH AMERICAN FAUNA. [No. 12.

uiue. Since this tootli in the Americau species lias the same structure
as in Microtus arralis, no special description is necessary.

Bade upper molar. — In the typical species the last upper molar has
an anterior transverse loop, a closed triangle on each side, and a very
short, simple terminal loop. With these loops are associated six salient
angles, two on each of the terminal loops and one on each closed triangle.
Rarely the posterior terminal loop is reduced by the isolation of the
outer basal angle as a third closed triangle, but this seldom happens,
while the resulting form of tooth is quite difierent from that found in
any member of the subgenus Microtus except the aberrant Jf. nivalis. In
the American species this tooth is formed exactly as in Microtus arralis.

Mammw. — There are eight niammie in Arvicola, as in Microtus.

Feet. — In Arvicola the soles are very sparsely haired or almost naked
between the tubercles and the heel.

The tubercles are only five in number, as the small one which in
Microtus lies midway between the large proximal tubercle and the base
of the fifth toe is absent. Claws moderately developed, those on hind
feet slightly the larger.

Fur. — The fur is close, dense, and long, the under fur especially thick
and woolly. It thus resembles the fur of Neofiher^ though the modifica-
tion is not carried so far as in the latter.

Miscellaneous characters. — The species of Arvicola are provided with
a large musk gland on each side of the abdomen. These glands lie
immediately in front of the hind legs and are very conspicuous in alco-
holic specimens. In a half-grown male Microtus terrestris from St.
Petersburg, Russia, each gland is 13 mm. long by 6 mm. wide. They
are regularly oval in outline, the long axis parallel with the long axis
of the body. The surface, which is slightly raised above that of the
surrounding skin, is closely and irregularly wrinkled, and has much
the appearance of very finely honeycombed tripe. Each gland bears a
sprinkling of fine hairs much shorter than the fur, but at first sight
appears to be naked. In dried skins the positions of the glands are
indicated by tufts of grease-soaked fur.

General remarks. — The subgenus Arvicola is distinguished from all
other groui)S with similar enamel pattern or with like numbers of
mammte and foot pads by the presence of the large glandular masses
on the sides of the body. The species are all water rats, and, with the
exception of Microtus [Neojiber) allem., they considerably exceed the
other members of the genus in size.

Although this subgenus is now for the first time recorded from
America, at least three species of Arvicola inhabiting the western
United States have been described within the past five years. These
are 3[ierotns macropus (Merriam), M, arvicoloides (Rhoads), and M.
principalis Rhoads. Microtus macropus was supposed to be "one of
the western members of the subgenus or section Mynomes,^' that is, a
tetramerodont Microtus.^ Microtus arvicoloides was made by its descri-

' North American Fauna No. 5. p. 60, July, 1891.

July. 1896] SUBGENUS NEOFIBER. 69

ber the type of a new genns, Aulacomys,^ while M. principalis, closely-
allied to botli M. maeropns and M. arvicoloides, was referred by the same
author to true J/icro/M.v.'- This coufusion arose from the fact that the
subgeiieric aud generic determinations were based chiefly on dental
characters. Hence Mierotus maeropns and M. principalis were naturally
considered members of the subgenus Mierotus, since both have the
enamel pattern characteristic of the tetramerodont species of that group.
The teeth of the type and only kuown specimen of Mierotus arvico-
loides, on the other hand, show certain characters which, although clearly
abnormal, led to an entire misunderstanding of the animal's true rela-
tionships. The first of these abnormal characters, and the one which
suggested the name Aulaeoniys, is seen in the upper incisors. Each of
these has a narrow longitudinal median groove. They can not, however,
be considered as entitling the species to generic rank, since similar
though fainter grooves are occasionally found in almost any species of
Mierotus, while they are absent in the vast majority of specimens of
'■Aulaeomys.'' The second abnormality in the type of Mierotus arvico-
loides is in the form of the front lower molar. This tooth (fig. 35) has
two reentrant angles on the outer side of the anterior
loop instead of one as usual in Mierotus. The supi)le-
mental reentrant angle, like the grooves in the incisors,
is purely an individual character, which may crop out ^^^"' ^^--^^J^iormai

'■ "^ ' ^ L front lower molar

in any species of Mierotus, with the front lower molar of type specimen of
formed as in M. arvalis, and which is absent in all the 'Mdacomys- arvico-

• > I 7 ITT loides. (x4.)

other thirty or more specimens ot ^ Aulaeomys^ that 1
have seen. The subgenus Aulaeomys if retained as distinct from Arvi-
eoJa must rest on characters of enamel pattern alone, since in all other
peculiarities it agrees perfectly with the latter. The difterences in
enamel folds are rather considerable, since ^Aulacomys^ has the highly
complicated pattern of true Mierotus, while the species of typical Arvi-
cola have the simplest pattern of any known. While it seems highly
inadvisable to base subgeneric divisions on such characters, the deci-
sion rests on purely individual judgment.

In the Old World numerous species and subspecies are probably
confused under the name ^Arvieola amphibius.'' Mierotus musignani (De
Sclys Longchamps) and M. monticola (De Selys Longchamps) appear to
be especially distinct from ^1/. terrestris (Linn.).

Snbgeuus NEOFIBER True.

1884. Xeqfiber True, Science, IV, p. 34, July 11, 1884 (full genus). Type Neofiher alleni

True.
1891. Neofiher Merriam, North American Fauna, No. a, p. 59, July, 1891 (snl)geuus).

Geoyraphical distribution of type species. — Florida. "Doubtless a com-
mon animal in favorable localities throughout the State." (Chapman.)

'American Naturalist, XXVIII, p. 182, February. 1894.
^American Naturalist, XXIX, p. 940, October, 1895.

70 NORTH AMERICAN FAUNA, [No. 12.

Geographical distrihutioyi of suhgenvs. — The range of the subgenus
Neofiher is the same as that of the tyi)e and only known species.
EssentUd characters :

Palate abnorm:il.

m 3 with all triangles closed.

m 1 with 5 closed triangles and 9 salient angles.

m 3 with 2 closed triangles and (3 salient angles.

MamniiTB 4.

Plantar tubercles ;">.

Sole naked.

Claws on hind feet longest.

Fur highly modified.

SJx-idl. — The skull of Xeo fiber is characterized by its large size, great
dei^th through the frontal region, and conspicuous development of
postorbital processes. The ratio of fronto-palatal depth to basilar
length is about 41 in Xeojiher, while in true Microtus it is onlj^ about
35. As the occiput in Xeojiher is not correspondingly high the dorsal
outline of the skull curves gently and regularly from front to back,
with the higliest point Just behind the orbits. When viewed from

above the skull of Neofiber differs from that
of Microins chiefly in the larger scpiamosals,
smaller parietals and interparietal, and in
the sharp-pointed postorbital processes.
The latter project over the orbital cavity
as square-cornered shelves, which are espe-

FiQ. S6.— Enamel pattern of molar cially UOticeablC whcil vicWcd from bclOW.

teeth, Microtus (Xeofiber) aiieni. Palate.— T\\& bouy pahitc iu Xcofiber (PI.

II, fig. 9) differs widely from that of MicrotuSy
and exactly resembles that of Fiber (p. 72).

Enamel liattem in general. — In general the enamel pattern of Xeo-
fiber (fig. 36) is characterized by a tendency to reduction in the number
of angles in the variable teeth and to the tight closure of all triangles.
The latter peculiarity gives the teeth the greatest possible strength.

Front lower molar. — The first molar in the lower jaw exactly resem-
bles the corresponding tooth in Microtus except that the anterior loop
is rather shorter than in the typical members of that subgenus. In
one specimen (jS"o. 23153, U. S. Nat. Mus.) the anterior loop has two
indentations on the outer side, thus suggesting Anaptogonia.

Back upper molar. — The third maxillary tooth is like that in the
subgenera Fitymijs, Fedomys, Fhaiomys, Chilotus, and typical Arvicolafi
as it has only two closed triangles and six salient angles.

Other teeth. — The back lower molar has all the triangles tightly closed,
in this respect differing from all other subgenera except Lagnriis.
Closed triangles are sometimes formed in the third lower molar of
almost any of the larger voles, but Xeofiber awd Lagurns are the only
groups in which they are always present. (Outside the subgenus
Lagurns, most of the known species of which are small, the tendency to

July, 1896.] GENUS FIBER. 71

closure of tlie triangles in this tooth increases with the size of the
animals until in such large species as Microtus alleni and the niembers^
of the genus Fiber they are always tightly closed. Microtus terrestrisj.
the only species approaching ^1/. alleni in size, has closed triangles in
m o very often, while in one specimen the tooth is formed exactly as in
Xeofiber. M. 'princiimlis Rhoads, another large species, also rather
frequently shows closed triangles in this tooth. The incisors, like those
of Fiber, are short, broad, and very strong, in this respect reaching the
opposite extreme from that attained by ^Anlacomy.s.^

2famm(c. — Ajiparently the number of mamm.Te in Neojiber has never
been stated in print. Mr. Outram Bangs writes me, however, that he
found four inguinal teats in an adult female Microtus alleni which he
took in Brevard County, Fla., during February, 1895.

Feet. — Soles wholly naked, foot pads five, as in Arvicola; claws on
hind feet longest.

Fur. — The fur is modified to meet the requirements of an aquatic life
in the same way and to almost the same extent as in the genus Fiber.
The under fur is exceedingly thick, woolly, and dense, while the longer
hairs are very glossy and lustrous. This condition is suggested in
Arvicola, where, however, the moditication is not carried so far.

Miscellaneous characters. — Whether Xeofiber is i^rovided with musk
glands like those of the other water rats is at present uncertain. Col-
lectors have failed to notice them, but they might easily escape detec-
tion in the thick fur unless specially searched for. The only alcoholic
specimen that I have examined is not iwW grown. This shows no trace
of the glands even when the skin of the sides is raised and examined
from beneath.

General remarks. — In Xeofiber are combined the mandibular enamel
pattern of Lagurns with the maxillary enamel pattern and external
characters of typical Arvicola, complicated by a reduction in the num-
ber of mammae as in Pedomys and Pitymys.

Genus FIBER Cuvicr.

Fiber Cnvier [Tabl. Ele'm. de I'Hist. Xat. des Auiin.. p. 141, 171»8], Lemons d'Auat.
Comp., I, tabl. I, 1800. Type Castor zihethicus Liuii.

Geographic distribution of type species. — Xorth America iu)rtli of the
southern border of the United States.

Geographic distribution of genus. — The range of the genus Fiber is
essentially the same as that given for the type species.

Essential characters :

Upper incisors witli antorior faces smooth.
Lower incisors with roots on outer side of molars.
Molars rooted.

Enani(d pattern characterized by approximatf e(iuality of reentrant angles on
outrr and inner sides of molars.
Feet modified for swimming-.
Tail flattened laterally.

72

NORTH AMERICAN FAUNA.

[No. 12.

Fig. 37.— Skull of Fiber zibethicus
(natural size).

Sl-ull. — The skull (fig. 37) differs very slightly from that of Microtus

except that it is cousiderably larger than in any known species of the

latter, and has a proportionally longer ros-
trum. The bony palate (PI. II, fig. 12) re-
sembles that of the species of Altk-oUi and
J^eofiher in the extension forward of the in-
terpterygoid fossa and suiipression of the
sloping part of the median ridge. The pos-
terior border is thus squarely cut off imme-
diately behind the lateral bridges. A vestige
of the sloping ridge usually persists in the
form of a median spine projecting into the
ill terpterygoid space. The skull of Fiber is
peculiar in the expansion of the squamosals
on the dorsal surface of the skull at the ex-
pense of the x)arietals. The postorbital proc-
esses of the squamosals form prominent
triangular projections closely resembling
those of Keofiher. The interparietal is
squarish in outline and usually somewhat

longer transversely than antero-posteriorly.

Teeth. — The molars are all rooted in the adults (fig. 38), though the

roots on the back lower tooth are usually

less w^ell developed than those on the

others. Otherwise the teeth are exactly

as in MicrottLS. The enamel pattern

(fig. 30), most closely resembles that of

Microtus [Xeofiber) aUeni, but differs in

the larger anterior loop of the first lower

molar. This loop is cut by two deep

reentrant angles, which often isolate

two additional closed triangles, making

seven in all.
Feet. — The feet are large and so formed

that they can be turned edgewise when carried forward, thus producing

the least possible resistance to the water Avhile the animal is swimming.

This character is, however, to a certain ex-
tent, reproduced in the more aquatic species
of Microtus and can not be considered diag-
nostic of Fiber.

Miscellaneous characters. — Thetail is strong-
ly compressed laterally, making an effective
rudder. The peculiar form of the tail is
scarcely noticeable in the young even when
large enough to leave the nest, but develops

rapidly as the animals increase in size.

The fur of the species of Fiber is highly modified to produce a

Fig. 38.— Sick' view of molars, Fiber zibeth-
icus. (X li.)

Fig. 39.— Enamel pattern of molar
teeth, Fiber zibethicus. (x 2^.)

JvLY, 1896.] GENUS BRAMUS. 73

tliorougiily waterproof covering. The long Lairs are remarkably close
and glossy, wliile the under fur is very dense. In the character of the
fur Fiber is approached by some of the aijuatic species of Microtus,
esi>ecially M. (Arricola) terrestris and M. (Xeojiher) alleni.

General remarls. — Fiber is very closely related to Microtu.s, from
which it is distinguished by its tlattened, rudder-like tail, and rooted
molars.

In addition to the well-known musk rat, Fiber sibethicus, three forms,
whose interrelationships are not yet understood, are uow recognized.
These are: Fiber zibethicns paUidns Mearns, F. obsciirus Bangs, aud
F. riralicins Bangs.

DESCRIPTIONS OF EXTINCT CtENERA AND SUBGENERA,

Three extinct rodents referred by authors to the family Microtinec
have been made the types of superspecific groups. Two of these, from
the Postpliocene of Pennsylvania, are subgenera of TIf /cro/».sf; the third,
from the Quaternary i)hosphorites of Trara de Nedroma, near Ain-
Mefta, Tunis, is a genus of doubtful affinities. As these groups are
necessarily based almost wholly on dental characters, it is impossible
to describe them in tlie same manner as the living genera and sub-
genera. It is furthermore impossible to form a clear judgment of the
validity of the groups in question without examination of the actual
specimens. Such examination I have not been able to make. Hence
the few conclusions here reached are necessarily incomi»lete and
unsatisfactory.

The genus Bramus Pomel (Comptes Rendus, Paris, CXIY, p. 1159,
1892), from the Quaternary Phospliorites of Tunis is represented by
one species, Bramus barbarus Pomel. Of this animal the mandible and
the teeth of both jaws are known. ^ These show characters which sug-
gest the Casioridce.

'Les molaires montrent sur leur couroiine la structure de celles flu r<ifc d'eau, dont
dies out ii peu pres les diuiensious. Ou y voit une double serie d'eucoches et d'angles
alteruatifs qui oorrespoudeut lateralemeut a des aretes saillautes, 5 eu dedans et 4 eu
dehors a la premiere dent iuferieure, 3 de ehaque cote aux deux suivantes iuferieures
et aux deux premieres sup^rieures et 2 seulement avec arete post^rieure a la troisieme
d'en-liaut. Chez Arvkola cette derniere est beaucoup plus conipliqu^e, ayant trois
paires d'aretes et un fort contrefort posterieur. Dans la fossile les sillons sout moins
profonds, a angles moins A-ifs, ainsi qne les aretes, et les lignes d'email ne se soudeut
pas d'un cote a I'autre de la couronne, ainsi qu'elles le font chez Arvicola ; il eu rosulte
nue lign. ni^diane continue de dcntiue sur la couronne, au lieu d'une s^rie alternative
de petits triangles hordes d'email; de sorte que la dent (V Arvicola est, en realite,
formee de deux rangces de prismes distiucts, tandis que celle du fossile est nn prisme
unique fortemcnt sillonne sur les cotes. II y a plus de ressemblance avec certains
Gerbilles, qui out cepeudant les molaires bien moins prismaticjues et antrenient
coustituees.

Les molaires des Arvicola ne sent jamais radiculees sauf peut-ctre chez les tres
vieux sujets. Dans notre fossile, je les ai trouvees toujours radiculees des qu'elles
percent ralvoole dentaire; leur fut, quoique franchement prismatique, est bieu moins
allonge. Ses deux racines, a la vdrite, sout tres longtemps ouvertes a leur extremite,

74 NORTH AMERICAN FAUNA. [No. 12.

The iiiolar.s, which are rooted, do not differ essentially in enamel pat-
tern from those of living species of 2ficrotus, except that the back
npper tooth is remarkably simple in structnre, and the reentrant angles
in all the teeth are so shallow that the triangles are open. While the
front lower molar has nine salient angles, as in tyi^ical Mkrotus, the pos-
terior maxillary tooth has only four and a very small terminal looj).
The author remarks that the open triangles give the teeth of Bramns a
resemblance to those of some of the Gerhiilida', but this likeness must
be very superficial. The most remarkable character of Bramus is the
form of the mandible, Avhich is like that of Castor and very unlike that
of any of the ]\[uri(la'. It is probable that Bramus is the tyi3e of a group
differing too widely from any of the recent Mlcrotimv to be united with
them in one subfamily.

The subgenera. Imdelta and Anaptogonia were described by Prof.
E. D. Cope in 1873 (Proc. Amer. Philos. Soc, XII, p. 87). Both are
based on teeth from the Postpliocene deposit in Port Kennedy Cave,
Pennsylvania. Anaptogonia is very different from any of the living
subgenera of Microtus — so different that, as Professor Cope suggests,
it may be eventually recoguized as a distinct genus. Isodelia, on the
other hand, is hardly separable from Fitymys, since the characters
pointed out as diagnostic of the two groups are not beyond the range
of variation among the species of one subgenus.

The original description of Microtus hiatidens, the type of the sub-
genus A«fl^^O(/oj(«V/, is as follows:

Represented by several molar teeth. These are several times as large as the teeth
occupying the same position in any of the species alreadj^ mentioned in this essay,
and suggest the genus Fiber. The distinctive features of the latter are the com-
pressed, oar-lilve tail, with rooted molars, and it is evident that the relationship of
this species is not to it. Perhaps it is neither an J r/coZa (sic.) l^Microtits'] nor a
Fiber, since it diifers in the structure of the teeth from the known species of both.
None of the triangles are isolated, but are connected by a narrow striii of dentine,
which is narrow posteriorly, but widens anteriorly until it opens out into the ter-
minal loop. Thus the sectional nsmie Anaplof/onia maybe found ultimately appli-
cable to a separate genus. The separation of the enamel folds merely carries to the
highest degree that which is seen in the anterior part of the tooth of ^. sigmodus.

In the inferior m 1 the triangles, which do not open on one side to the anterior
loop, are IJ, then one on each side, and the short, wide, terminal loop, which is
bilobed or emarginate in the middle of the end. The ridges, Avhich are very promi-
nent and acute, are, therefore, i ; at the extremity there are two short ones, between

mais elles sont de bonne heure parfaitement distinctes I'uue de I'autre. La troisifeme
molaire iuferieure, un peii plus arqu^e que dans ArvicoJa, ne descend pas a la face
interne de Tincisive, nuiis reste tout h fait au-dessus, et ses racines seules s'insiuueut
un peu lat^rulement sur cette face.

L'os maudibulaire presente des diltereuoes beaucoup plus importantes. Son
apophyse angulaire, restaut presque dans le plan general de l'os, ne fait en ai'riere
qu'uue l(5gere saillie bordaut la branche montante, (ju'elle suit tres haut sous le
condyle pour se terminer en simple petit cran. II y a une grande analogie de forme
avec ce que I'on voit chez les Castors. Dans Arvicola, au contraire, I'apophyse
angulaire est basse et se rejette oblitiuement en arriere en forme de cuilleron forte-
ment crocliu et tordu, rappelant du reste, sauf cette torsion la disposition de cette
partie chez les autres Murides.

July, isnfi.] ARVICOLA INTERMEDIUS NEWTON. 75

■which a third and more promiuent oue rises a little below the grinding snrfaec. A
little more attrition would give the distal loop a trilobate outline, and a little more,
an acuminate one, from the loss of the lateral angles; finally the median ridge
disappears also.

Tbe subgenus Isodelta is cousidered by Professor Cope to show au
exaggeration of the characters of Pitymy.s. Tbe type and only known
species, Microtus .sjyeothen, is described as follows:

This species is represented by the entire dentition of the left ramus mandibuli,
with a few fragments of the adjacent bone. As already pointed out, its characters
entitle "it to rank as a listinct section of the genus. Thus, the triangles of the inner
side of the anterior inferior molar are one less than in any species of the section
Anicola l = Mic7-otus']. The anterior loop presents two well-mai-ked angular basal
areas, while its terminal portion is regularly rounded. « * * That this is not one
of the species of Plti/miis, in which the basal lobe of the anterior trefoil has been cut
off by unusual inflexion of the enamel angle, is demonstrated by the structure of the
second molar, Avbich is precisely that of typical Arvicola [== J/)c/o/hs], all the tri-
angles from the posterior being isolated and alternating, producing the formula
1 1 0. The third molar has the usual formula, 1-1-1, the posterior two lobes being
crescentic, the anterior trapezoid.

NOTE ON ARYICOLA INTERMEDIUS NEWTON.

In a paper entitled ' Tbe Vertebrata of tbe Forest Bed Series of
Norfolk and Suftblk' ^ Mr. E. T. Newton describes nanierous remains
of a niicrotiue rodent with well-developed fangs on tbe molar teeth and
intermediate in size between Arvicola amphibius [ = Microtus ferres-
tris] and tbe smaller voles. This animal, which Mr. Newton named
Arvicola intermcdius, has been recently referred to the genus Phena-
comi/s.- While the species is certainly not an Arvicola [=Microtus], it
appears to be equally far removed from Phenacomys and ])robably from
Evotomys and Fiber also. The teeth are described as follows:

I have now before me about 40 vole jaws from the " Forest Bed '' which, altliough
dithering somewhat in size, agree precisely in the patterns of their teeth. Only 14
of these allow the bases of their teeth to be seen, but nine of these have more or
less distinct fangs; the other five have no fangs, but are most probably immature,
as in other particulars they agree precisely. I have likewise some hundreds of iso-
lated molar teeth, and a very large proportion of these are fanged. * * * The
great variation in the size of these fanged teeth would lead one to suspect that they
represent more than one species, but there are no sufficient grounds for their separa-
tion. * * * The patterns of the grinding teeth are so nearlj^ like those of J.
(unphihins as scarcely to need description, and it is on the presence of fangs in the
adult that the chief distinction between the two species I'ests; nevertheless, there
are a few points deserving of notice. In one of the largest and most perfect man-
dibular rami (figs. 3, 3a) the entire molar series, measured along the alveolar margin,
180.33 inch (8.5 mm.). Mr. Reeves's specimen, from the Bramertuu Crag (tig. 12), is a
little larger. The first molar has the five inner and four outer angles alternating,
but the anterior two are not so prominent as is usually the case in A. ampliihins, and
the front of the tooth is somewhat more rounded (fig. 3/^). In the Bramerton jaw
this is especially the case (fig. 12a). All the anterior lower teeth from the "Forest
Bed" series which I have seen have the infoldiugs of the enamel behind the ante-
rior prism less deep than in those examples of A. anipMhius which I have been able

'Memoirs of the Geological Survey, England and Wales. London, 1882.
= Nehring, Naturwissenschaftliche Wochenschrift, Xr. 28, July 15, 1894.

76 NORTH AMERICAN FAUNA. [No. 12.

to examine; aud hence tlie dentinal portion of tlie anterior prism is more widely
continent with the second inner and outer prisms; it is, in fact, an exaggeration of
the form indicated hy Blasins, tig. 186 (Slingethiere Deutschlands, p. 345). The second
molar has three inner and three outer angles alternating. The third molar has like-
wise three inner and three outer angles, hut the alternation of the prisms is so slight
that the opposing inner and outer prisms are confluent. ^ * '^ I am not acquainted
with any specimen which shows the three upper molars in place, hut Mr. Savin has
two specimens which retain the first and second upjier grinders (fig. 1), and Mr.
Reid hag obtained several isolated specimens of last upper molars. The anterior
upper molar (tig. la) has three inner and three outer angles alternating; the second
tooth has three outer and two inner angles alternating. The third upper molars
vary somewhat; in some only three inner and three outer angles can he counted
(fig. 2a), while others have three inner and four outer angles. The widely confluent
character of the front prisms of the lower anterior molar is repeated in these hinder
upper ones. It will he noticed that in all Blasius's figures of the last upjier teeth
(I.e., p. 345) the anterior inner fold (cement space) and the two anterior outer
folds extend across the teeth and meet the enamel of the opposite side, while in
one case (fig. 190) the two inner folds pass across. Now, in most of the teeth under
consideration it is only the one anterior inner and one anterior outer fold which pass
across; in some instances the second outer fold passes farther inward, hut I do not
think that in any instance it touches the opposite side.

The teetli of '■Arvicola'' intermedius dift'er in nuuierous characters
from tbose of Fiber, Uvotomys, aud Phenacomys, the only known living

microtines witb rooted molars. The small
size of the remains and the simple struc-
ture of the first lower molar are sufficient
to indicate that the animal is not closely
related to Fiber, although the chara(;terof
the roots of the molars, as shown in tigs.

TlG.40.-Enamelpatternofmolarteeth, 5 g ^^,j(^| 7 ^^f p|^ XIII, is StrOIlgly SUggeSt-

Arvicola intennedius. From Newton. ^ ^ <=<

ive of this genus. The figure of the inner
side of the lower jaw (PI, XIII, fig. 3ff) suggests that the posterior molar
is strongly displaced by the shaft of the incisor, as in Microtus. This
character alone would show that the species is neither an Evotomys nor
a PhenacomyH ; but the peculiarities of the enamel pattern furnisli addi-
tional reasons for its exclusion from these genera. The enamel pattern
(fig. 40) is, as Mr. Newton remarks, almost exactly like that of Mierotns
terresiris (see fig. 34). It thus lacks the deep reentrant angles on the
inner side of the lower molars characteristic of Phenacomy.s, and the
rounded salient angles aud opposite triangles characteristic of Fvo-
tomys. The last lower molar in particular is noticeably different from
that of either Evoiomys or PJienacomy.s. '■Arricolu ' intcnuediu.s is appar-
ently still further removed from Erotomys by the large size of the teetli
as compared with the jaw. There can be little doubt that the animal
represents a genus distinct from any now living.^ In the absence of
si)ecimens. however, nothing would be gained by an attempt to name
aud define the group.

' Whether the rooted microtine teeth mentioned hy Nehring (Naturuissenschaftliche
Wochenschrift, Nr. 28, .Tuly 1894) and hy Forsyth Major (Atti 80c. Ital. Bci. Nat.,
XV, p. 389) belong to animals congeneric with Jri-icola intermedius is purely a matter
of conjecture.

INDEX

[Synonyms in italics.]

i Aijricola. 11,10,21,62.
i Alticola, 9, 17. 19, 23, 5l'-54.
I Alviccola, 15.
' Arnmomys, 15, 58.
( Anaptogonia, 17, 74-75.
1 Antelioniys, 9.
Arctomys, 13.
Arvicola, 9, 14.
Arvicola, 14.

atiiphibuis, 14.
intermedins, 75-7t)
Arvicoiince, 8.
Avlacomys, 18, (59.

Bainl, classification adopted, by, 21-22.
Blanlord, classification adopted by, 23.
Blasius, classification adopted by, 21.
Bony palate, 26-28.
BorioUcon, 17, 38.
Brachyurus, 15.
Bramus, 18, 73-74.

barbarus, 73-74.
Biiclmer, opinion on taxononiic value of enamel

pattern, 25.
Camjiicola, 18, 62.
Castor, 12.

Chilotus, 9, 16, 19, 22, 60-62.
Coues, classification adopted by, 22-23.
Cuniculns, 12.
Cuniculus, 16, 38.
De Selys-Longcliaiiips, classification adopted Tiy,

19-21.
Dicrostonyx, 8, 9. 10, 38-40.

ton^uatus, 9, 40
Enamel pattern, 25.
Eothenomys, 9. 45-47.
Eremiomyg, 17, 50.
Evotomys, 8, 9, 17, 22, 28, 42-44.
californicus, 44.
fuscodorsalis, 44.
galei, 44.
gapperi, 44.
glareoliis, 44.
idahoensis, 44.
occidentalis, 44.
rufocauu.s, 44.
rntilus, 9, 44.
Fatio, classification adopted by, 22.
Fiber, 8, 9, 14, 71-73.
obscurus, 73.
rivalicius, 73.
zibethicus, 9,71-73.
Glis, 12.
Glis, 12, 13.

Hemiotomys, 16, 19, 20, 22, 62.
History of classifications, 19-24.

Hyperacrius, 9.

Hypudceus, 14, 21, 22.

Interpterygoid fossa, 27.

Isodelta, 17, 74-75.

Keys, 28-32.

Lagomys, 13.

Lagurus, 9, 16. 49-51.

Laiiox>odumys, 18, 24, 57.

Lataste, classification adopted by, 23-24.

Lateral bridges, 27.

Lateral grooves, 27.

Lemuii, 8.

Lemmings, 8.

Lenimus, 8, 9, 13, 36-37.

lenimus, 9. 37.

nigripes, 37.

obensis, 37.

.scliisticolo7-, 37.
5VIarinota, 13.

Maxillo-palatine siituri'. 26.
Microti, 8.

Microtina?, geographic distribution, 9-10.
habits, 10-11.

lists of genera and subgenera, 9.
subfamily and divisions, 8-9.
Microtus, 8, 9, 14, 19, 20, 21, 24, 44-71.

agrestis, 66.

albicauda, 54.

alleni,9, 69-71.

arvalis, 9, 62, 66.

arvicoloides. 08-69.

austerus, 9, 55-.56.

blanfordi. 54.

blythii, 9,57.58.

braudti, 58.

characters on which present classifica-
tion of subgenera is based, 24-28.

chinensis, 9, 47-49.

chrotorrhinus, 66.

curtains, 51.

fertilis, 9, 54-55.

fuscus, 58.

(genus), 44-15.

guentheri, 06.

hiatidens, 74-75.

iagurus, 9, 49, 51.

longicauda, 66.

Inteus, 49, 51.

macropus, 69.

mandarinus, 58.

melanogaster, 9, 45-47.

middendortfi, 24, 49.

mogollonensis, 66.

monticola, 69.

77

78

INDEX.

Microtiis musiguaui, 09.

nivalis, GO.

oregoni, 9, 60-C'J.

pallidas, 51.

liaunerriruus, 51.

peunsylvfinicns, 66.

(peutamerodont t>i)ecie.s), 65.

piueturuni, 9,58,59.

principalis, C9.

przewal-sliii, 51.

quasiater, 60.

ratticeps, 66.

roylii, 54.

.savii, 60.

sikkiiuensis, 65, 66.

speotlien, 75.

stolifzkanii.s, 9,52,5-1.

stradiej-i, 54.

.straucbi, 58.

(subgenus), 62-66.

subterraneus, 60.

terrsenova?, 66.

terre.stris, 9, 66, 69.

(tetramerodont species), 65.

townsendi, 66.

wyunei, 55.

xantliognatlius, 66.
Micruriis, 17, 58.
Mictomys, 9, 18, 35-36.
Misothermus, 16, 38.
Mu8, 11.

amphibins, 11.
terrestris, 11.
Mijnomes, 15, 19, 20, 21, 23, 62.
Myocastor, 13.
Myodes, 15, 20, 24.
Myolemmus, 16, 38.
Myotalpa, 8.
Myotalpiuae, 8.

iVeodow, 16, 19,23,62,65.

Neofiber, 9, 17, 19, 69-71.

Nomenclature, 11-19.

Ochetomys, 17, 66.

Ondatra, 13.

Palatine bone, 27.

Paludicola, 17, 19, 21, 23, 62.

Pedouiys, 9, 16, 19, 22, 55-56.

Pentamerodout species of Mierotus, 65.

Pbaiomys, 9, 17, 56-58.

Pbenacomys, 8, 9, 18, 40-42.

celatus, 42.

intermedius, 9,42.

latimanus, 42.

longicauda, 10, 42.

oramontis, 42.

orojihilus, 42.

truei, 42.

ungava, 42.
I'incmys, 16, 58.
Pitymys, 9, 15, 19, 22, 24, 58-60.
Praticola, 17, 22, 62.
I'aaminomys, 15, 58.
Siphneince, 8.
Sijfhneus, 8.
Sylvicola, 17, 22, 62.
.Synaptomys, 8, 9, 16, 32-36.

cooperi, 9, 35.

dalli, 36.

fatuus, 35.

helaletes, 35.

innuitus, 9.

(subgenus), 34-36.

truei, 36.

wrangeli, 36.
Terricola, 17, 58, 62.
Tetramenidon, 18,62.
Tetramerodont species of JUiCrotus, 65.
Voles, 8.

PLATE I.

[Enlarged cmo aiul om-half times.!

Fig. 1. Microius {Arricola) macropny. AVood Rivt-r, Idaho.
(No. 31630, U. S. Nat. Mus.)

2. Microtus (Pilymys) jjinetoriiiu. Wasliiugton, D. C.

(No. 30332, U. S. Nat. Mns.)

3. Microtus (Microtus) arralis. Cepiu, m^a.. Esszek, Slavouia.

(No. 3035, collectiou of Gerrit 8. Miller, jr.)

4. Evotomys. Portland, N. L)ak.

(No. 35835, U. S. Nat. Mus.)

5. Phenacoviys oramontls Rlioads. Mount Baker Range, British Columbia.

(No. 3562, collection of Gerrit S. Miller, jr.)

6. Lemmus nigripes. St. George Island, Alaska.

(No. 42680, U. S. Nat. Mus.)

7. Microtus (Lagurus) curtatus. Reese River, Nevada.

(No. 32498, U. S. Nat. Mus.)

8. Microtus (Chilotus) oregoni. Sinnas, British Columbia.

(No. 4160, collection of Gerrit S. Miller, jr.)

9. Microtus (Arricola) terrestris. Braunschweig, Germany.

(No. 1934, collection of C. Hart Merriam.)

10. Microtus (AUicola) alhicauda. Type. Braldu Valley, Ballistan.

(No. 36916, U. S. Nat. Mus.)

11. Microtus (Hyperacrius) fertilis. Pir Panjal Range, Kashmir.

(No. 35511, U. S. Nat. Mus.)

12. Synaptomys (Miciomys) wrangeli. Wrangel, Alaska.

(No. 74720, U. S. Nat. Mus.)

13. Synaptomys (Synaptomys) hclaleies. Dismal Swamp, Virginia.

(No. 75172, U. S. Nat. Mus.)

14. Dicrostonyx torquatus. Petschora, Russia.

(No. 3621, collection of Gerrit S. Miller, jr.)
80

North American Fauna, No. 12.

Plate I.

^ 5
||

5 5

C " ij

t-occi

Q fc^is^-l

16933—^0. 12 6

PLATE II.

[Enlarged two aud oue-half times.]

Fig. 1. Bony palate of Phenacoinys. Salmon River Mouuiains, Idaho.
(No. 31249, U. S. Nat.Mus.)

2. Bony palate of Microius {Lmjurus) pallidiin. Reese River, Nevada.

(No. 32498, U. S. Nat. Mus.)

3. Bony palate of ^('cro/MS (/'(7//7H^8) j>j«e<or((r«. Washington, D. C.

(No. 30.332, U. »S. Nat. Mus.)
t. l>ony palate of Microius (Alticula) hlanfurdi. Nultar Valley, Kashmir.
(British Museum Register, 81. 3. 1. 23.)

5. Bony palate of MicnAiis (Microtus) arvalis. Geneva, Switzerland.

(British Museum Register, 79. 9. 25. 52.)

6. Bony palate of Microtus {Lag urns) lagurus. Gurjeff, Russia.

(No. 3619, collection of Gerrit S. Miller, jr.)

7. Bony palate of Microtus (Arvicola) arvicoloides. Type. Lake Kichelos,

Washington.
(No. 1358, collection of S. N. Rhoads. )

8. \ionj -pal&te of Microtus {Anteliomys) chlnensis. Type. Western Sze-chuen,

China.

(British Museum Register.)

9. Bony palate of Microtus {Xeojiher) alleiii. Florida.

(No. 23452, U. S. Nat. Mus.)

10. Bonj iihIaIg of Erotomi/s glarcoJ us. Ghristiania, Norway.

(British Museum Register, 84. 10. 31. 11.)

10. View perpendicular to plain of palate.

10ft. View from below and behind at strong angle with jilain of
palate.

11. Bony jialate of Microtus {Eothenomijs) met anog aster. Western Fokien, China.

(British Museum Register, 92. 10. 12. 52.)

12. Bony palate of Dicrostonyx torquatus. Petschora, Russia.

(No. 3621, collection of Gerrit S. Miller, jr.)

13. Bony palate of Fiber. Lake George, New York.

(No. 67689, U. S. Nat. Mus.)

14. Bony palate of Lemmus lemnius. Vola. (From St. Petersburg Museum. ^

(No. 3620, collection of Gerrit S. Miller, jr.)

14. View perpendicular to i)laiu of palate.

14a. View from below and behind at strong angle with plain of
palate.
82

North American Fauna, No 12.

Plate II,

2-2
^ p

P 2
•S ?

Oh -J

PLATE III.

[Enlarged two and two-thirds times.]

Fig. 1. Synaptomys cooperi. Roan Mountain, North Carolina.
(No. 50865, U. S. Nat. Mas.)

1. Left mandible from beneath ; bone cut away to expose roots of

teeth.
la. Left mandible from inner side; bone cut away to expose roots of
teeth.

2. Pheiiacomys oramontis Rhoads. Mount Baker, British Columbia.

(No. 3562, collection of Gerrit S. Miller, jr.)

2. Left mandible from beneath ; bone cut away to expose roots of

teeth.
2a. Left mandible from inner side; bone cut away to expose roots of
teeth.

3. Microiii8 pennsylvanicus. West Tisbury, Mass.

(No. 1885, collection of Gerrit S. Miller, jr.)

3. Left mandible from beneath; bone cut away to expose roots of

teeth.
3a. Left mandible from inner side; bone cut away to exi)ose roots of
teeth.

4. Evotomys yapperi. Seekonk, Mass.

(No. 193, collection of Gerrit S. Miller, jr.)

Left mandible showing effect of excessive wear on teeth.
84

North American Fauna, No. 12.

Plate III.

^

1. Synajjtomys.

2. Phenacamys.

3. Micrdtiis.

4. Evotomys.

U. S. DEPARTMENT OF AGRICULTURE
DIVISION OF BIOLOGICAL SURVEY

NORTH AMERICAN FAUNA

IN"©. 13

[Actual tlate of jjublicatioii, October 16, 1897.]

.■.^:

y^

REVISION OF THE NORTH AMERICAN BATS OF THE
FAMILY VESPERTILI0N1D.E

BY

GERRIT S. MILLER, Jr.

Prepared under the direction of

Dr. C. HART MERRIAM

CHIEF OF DIVISION OF BIOLOGICAL SUllVEY

washii!^gto:n^

G-OVERNMENT PRINTING OFFICE
189 7

LETTER OF TRANSMITTAL.

U. IS. Department of Agriculture,

Division of Biological Survey,

Washinf/ton, D. C, July 1, 1897.
Sir: I have the honor to transmit herewith, and recommend for pub-
lication, the manuscript of No. 13 of North American Fauna, comprising
a monographic revision of the bats of the family Vespertilionidfv inhab-
itiug North America north of Panama, by Gerrit S. Miller, jr. It is
based mainly on material belonging to the Biological Survey, Avhere
tlie work has been done.

The Department is constantly in receipt of bats sent for identifica-
iou and of letters of inquiry concerning these animals; but heretofore,
owiDg to the chaotic state of the literature relating to this group and
the uncertainty respecting the status of the various species, it has been
impossible to answer such inquiries with any degree of certainty. The
present paper is intended to remove these difficulties.
Eespect fully,

C. Hart Merriam,

Chief, Biological Survey.
Hon, James Wilson,

Secretary of Agriculture.

CONTENTS.

Page.

Material 7

Changes iu color of s])ecimens preserved in alcohol 8

8exnal variation 8

Agt? variation 8

fieograpliic variation 9

(Geographic distribution 10

Migration 10

Measurements 11

Illustrations 12

Nonieuclature of North American Vespertiliouidie 12

1. Generic and subgeneric names 12

2. Specific and subspecilic names 20

Lists of North American Vespertilionidic 38

Descriptions 41

ILLUSTRATIONS.

PLATES.

1. Ears of Myotis velifer, il. californicus, M. ynmanensis, M. Ilijisanodes, M. evotis,

Xycticeius humeralis, Ehogeessa gracilis, B, tumida, Corynorhinus viacrotis town-
seiidii, Euderma maculatum, Jntrozous pallidiis.
Uropatagia of Ehogeessa fjvacilis, I\. tumida, Xycticeius hnvieralis.

2. Uropatagia of Ahjotis californicus. M. yumanensis, 21. evotis, M. thysauodes, M. relifer.

3. Wiugs of Plecotus, Corynorhinus, Euderma, aud Lasiurus.

TEXT FIGURES.

1. "Wings of Vespertilio serotinus.

2. Anterior part of rostrum of species of Phyllostomatidw aud Vespertilionida;.

3. ^IvLzzie of Antrozous.

4. Abnormal front teeth of Antrozous.

5. ^\i\\\\B of Antrozous.

6. Teeth of Antrozous.

7. Muzzles of riecot us and Corynorhinus.

8. Skulls of two subspecies of Corynorhinus.

9. Teeth of two subspecies of Corynorhinus.

10. Upper incisors of three specimens of Corynorhinus.

11. Skulls of My Otis myotis, M. thysanodes, aud M. nigricans (top).

12. Skulls of Myotis myotis, M. thysanodes, and M. nigricans (side).,

13. Ear of Myotis snhulatus, M. Jccenii, M. lucifugus, and M. alascensis.

14. Teeth of Myotis yumanensis, M. lucifugus, M. lucifugus longicrus, M. relifer,

15. Teeth of Myotis californicus, M. subulatus, M. evotis, M. thysanodes.

16. Maxillary teetli of four specimens of Myotis thysanodes.

17. Abnormal premolar of Myotis thysanodes.

18. iiknW of Lasionycteris noctivagans.

19. Teeth of Lasionycteris noctiragans.

20. Ear of Pipistrellus suhfavus aud P. hesperus.

21. Skull of Pipistrellus hesperus and P. subflacus (top).

22. Skull of Pi2ns1r ell us hesperus and P. siihflarus (side).

23. Tee1h of Pipistrellus hes2)erus and P. suhfiarus.

24. Skull of Vespertilio hahamensis, V. fuscus, aud V. serotinus (top).

25. Skull of Vespertilio hahamensis, V. fuscus, and V. serotinus (side).

26. Teeth of Vespertilio serotinus, aud four subspecies of V. fuscus.

27. Ear of Lasiurus horealis aud L. teliotis.

28. Skull of Lasiurus horealis and L. teliotis (top).

29. Skull of Lasiurus teliotis and L. horealis (side).

30. Teeth of Lasiurus teliotis and L. borealls.

31. Skull of L^aslurus cinereus.

32. Teeth of Lasiurus cinereus.

33. Skull of Dasypterus intermedins.

34. Teeth of Dasypterus intermedlus.

35. Skull of Xycticeius humeralis.

36. Teeth of Xycticeius humeralis.

37. Incisors of Ehogeessa aud Xycticeius (front").

38. Incisors of Ehogeessa aud Xycticeius (crowns).

39. Skull of ii'/(o_^e('8s« tumida.

40. Teeth of Ehogeessa tumida and E. gracilis.

6

No. 13. NOETH AMERICAN FAUNA. October, 1897.

REVISION OF THE xNORTH AMERICAN BATS OF THE FAMILY

VESPERTILIONID.E.

By Gerrit S. Miller, Jr.

Writers on American bats have published a large mass of facts con-
ceruiug the distribution and comparative anatomy of members of the
family Vesperfilionida'. Unfortunately, however, no work has yet
appeared in which the numerous species by which this group is now
known to be represented in North America^ are treated from the stand-
point of the systematic zoologist. In other words it has hitherto been
impossible for anyone not thoroughly acquainted with the extensive
and scattered literature of North American bats to identify specimens
correctly. The present paper has been prepared with special reference
to the loug-felt want of a ready means to accomplish this object.

MATERIAL.

The greater part of the material on which this revision is based is
contained in the collection of the Biological Survey of the IT. S. Depart-
ment of Agriculture. This collection of bats, which consists of more
than 0,000 specimens, chiefly in alcohol, has been brought together dur-
ing the past few years by the held naturalists of the Survey. In addition,
the writer has examined the bats in the United States National Museum,
the American Museum of Natural History, and several private collec-
tions, making a total of about 2,700 specimens of North American
Vesp(rtilio)tlda'. It is to be regretted that so few South American bats
are contained in the museums of the United States that no definite con-
clusions can be reached concerning the relationships of several Mexican
Species to the forms occurring farther south. For this reason certain
questions of nomenclature must for the present remain in a condition
of uncertainty. It is also to be regretted that comparatively fi^w well-
prepared skins are available for comparison. Without good series of
dry specimens it is impossible to determine the limits of individual
variation in color, as conclusions of the most general kind only can be
based on specimens that have been subjected to the action of alcohol.
Series of bat skins as extensive as those by which most groups of small
North American mammals are now represented will doubtless i)rove

'In the present paper the term North America is used to indicate the whole of the
North American continent and the West Indies.

8 NORTH AMERICAN FAUNA.

the existence of several well-marked geograpliic races in addition to
those now recognizable.

In tlie lists of specimens examined it lias not been thought necessary
to distinguish between those contained in the IS^ational Museum proper
and those in the collection of the Biological Survey. Specimens from
other collections, however, are always specially designated.

CHANGES IN COLOR OF SPECIMENS PRESERVED IN ALCOHOL.

Bats which have been kept in alcohol for a i)eriod of more than a
few mouths become so altered in color that they furnish reliable char-
acters of size and form only. The rate and amount of change appear
to vary with different species as well as with the strength of the jire-
servative fluid and the amount of exposure to light. 1 have seen two
lots of specimens of one species collected at the same place and on
practically the same date and su[)posedly treated in the same way, yet
after six years' immersion in alcohol those in one bottle still retained
essentially their normal color, as proved by comparison with skins col-
lected at the sanie time, while those in another bottle were so bleached
as to show scarcely a semblance of their original appearance.

While the details of the changes produced by alcohol are not known,
it may be said that a gradual bleaching and ultimate entire loss of
color is the general rule, though as a preliminary step browns are often
very noticeably reddened. The subject is one that merits exj^erimental
study.

SEXUAL VARIATION.

The range of sexual variation in North American Vesjiertilionida' is
always slight and in many cases scarcely api)reciable. For tlie most
part it consists in the slightly greater a\'erage size of the females.
Even this is often trifling or absent, as in the case ot Myotis hicifiifjus
longicnts from Nicasio, Cal., six males of which average: Total length,
Ito.l; tail vertebrie, 45.8 ; forearm, 37.8; ear, 11.8; tragus, 7.3; while six
females from the same locality average: Total length, 9G.3; tail ver-
tebra:', 44.1 ; forearm, 37.3; ear, 12.1; tragus, 7.2. In general, however,
it is necessary to take this factor into consideration when comparing
specimens from widely se])arated localities, I know of no instances
of constant sexual differences in color among North American Vesper-
tiIionid(v, and only one of differences in cutaneous structures, that of
L'hofjeissa gracilis^ in which the only known male has in each ear a
distinct glandular swelling, absent in the two females that I have
examined (see PI. I, fig. 7).

AGE VARIATION.

Young bats when nearly full grown often present characters different
enough from those of the adults to cause confusion in identification.
The fur of such immature specimens is usually shorter and more woolly

GEOGRAPHIC VARIATION.

than that of the adults and the color darker and duller. The immature
skull ditters in size and form from that of the adult, but as the sutures
disappear at an early age, it is often somewhat ditticult to recognize.

I have found that the best guide to the age of those bats that I have
studied is the condition of the linger joints. In specimens young
enough to furnish unreliable characters these are always large and
looselj' formed, with epiphyses sei^arate from the ends of the j)halanges
and metacarpals, both of which are distinctly enlarged for some dis-
tance from the joint (tig. I a). In adults the tinger joints are small and
compact, the epiphy-
ses no longer visible,
and the ])halanges of
essentially the same
diameter throughout
(tig. 1 h). These dif-
ferences are equally
a))parent in alcoholic
specimens and in
dried skins.

GEOGRAPHIC VARIA-
TION.

As comi)ared with
other small mammals,
bats show remarka-
bly little geogra])hic
variation in size, pro-
portion s, or color.
Thus bleeding indi-
viduals of Xycticeius
humeral is from Car-
lisle, Pa., Dismal
Swamp, Virginia, and
the extreme southern
point of Texas are
alike in color,' while
in size they agree almost as closely as any three lots of specimens
from one locality.'- The onl}- difference that can be found is a slight
northward increase in size of the ears. Specimens of Myotis Uici-
fufiihs from Washington, D. C, are not distinguishable from a series
taken on Kadiak Island, Alaska, and skins of Lasiurus clnereus from
Minnesota are exactly like others from southern California. While
such constancy of characters in wide ranging species is unparalleled
among American mammals, the only ones of which it is yet possible to

1 So far as cau be ascertained from comparison of specimens in alcohol.
^See table of measurements on page 120.

Fig. 1. — W)u<rs of Vespertilio serotinus: a, adult ; h. immature (natural
size).

10 NORTH AMERICAN FAUNA.

speak with certaiuty, the explanation of the fact is probably very sim-
ple. Living throughout the warmer part of the day in cool, dark, ami
for the most part damp situations, bats, even in widely separated locali-
ties, are exposed to comparatively little variation in temperature.
Feeding at a distance above the surface of the ground and during the
hours between sunset and sunrise, when colors are scarcely distin-
guishable, they are jiractically freed from that necessity for protective
coloratioii which binds the color of most mammals so closely to that of
their surroundings. From this reduction in the force of two of the
most powerful factors in the production of geographic variation — dif-
ferences in temperature and need for protective coloration — the com-
parative constancy in the characters of bats naturally results.

GEOGRAPHIC DISTRIBUTION.

From the peculiar habits of bats it results that the ranges of these
animals are less closely limited by life areas than in the case of most
mammals. To be more accurate, the frequent dampness and usual low,
even temperature of the retreats occupied by bats during the hot
part of the day expose the animals to essentially similar conditions
wherever they may be, so that a given region of like environment is
much more extended geographically for a bat than for most other
mammals.^

Therefore, although many species seemingly disregard the laws of
geographic distribution, their independence is more apparent than real.

MIGRATION.

A factor which introduces much uncertainty into the study of the
distribution of bats is the little understood migrations which some
species are known to make. That many bats migrate is a well-estab-
lished fact, but the extent to which migration aff'ects the apparent dis-
tribution of species is not known.

Although there are probably earlier references to the subject, the
first mention of bat migration that I have seen is by Dobson, in bis
Catalogue of the Chiroptera in the British Museum, published in 1878.
In his remarks on the geographic distribution of Pipistrellus abramuSj
Dobson says: "Found during the summer months in the Pahearctic
region throughout middle Europe; * * * evidently migrates north-
ward, * * * asithasnever been taken in Europe in winter" (p. 227).
In 1888 Dr. C. Hart Merriam published evidence in the Transactions of
Idae Eoyal Society of Canada (V, Section V, p. 85), which showed con-
clusively that two American bats, Lasiom/ctcri.s noctivagans and Lasi-
urus cinereus^ perform regular periodical migrations. Xo details of the

'Analogous condiMons are found in sphagnum bogs and heavy, damp woodlands,
in which animals of northern affinities, such as shrews, lemmings, and red-backed
mice, extend far south of the normal limit of their kind.

MEASUREMENTS. 11

extent or exact dates of tbe northward aud southward movements
could then be giveu further than that the known southern records of the
hoary bat (Soutli Carolina, Georgia, Bermuda Islands) were all during
autumn and winter, and that the silver-haired bat occurred in spring
aud fall about the lighthouse on Mount Desert Eock, 30 miles oft' the
coast of Maine, a treeless islet where bats were at other times unknown.
In August and September, 1890 and 1891, I liad an opportunity to
watch the appearance and disappearance of three species of bats, Lasi-
onycteris noctivagans, Lasiurus hoi-ealis, and LaHiurus cinereus^at High-
land Light, Cape Cod, Massachusetts. The animals, which were not to
be found during tlie early summer, suddenly became numerous shortly
after the middle of August aud remained abundant for about a month,
when they as suddenly disappeared. The regularity with which this
phenomenon occurred on the two successive years over which my obser-
vations extended shows that the migration of bats is probably as defi-
nite as to dates aud paths as that of birds. ^

MEASUREMENTS.

For general purposes of identification, ten measurements are useful.
Theseare : Total length, tail vertebni^, tibia, foot, forearm, thumb, longest
finger, height of ear from meatus, width of ear, and height of tragus.
The lengths of the separate phalanges of the fingers are important in
special cases only.

The tables which accompany the descriptions of the different forms
contain average measurements of specimens selected from as wide a
range of localities as possible. Whenever the full complement of meas-
urements is given, it is to be understood that all have been taken from
alcoholic specimens by the writer. When the total length, lengtli of
tail, and the three measurements of the ear are omitted, the measure-
ments have been taken from the dried skin. In a few cases the skin
measurements are supplemented by the collector's measurement of total
length and tail veitebra\ The use of specimens preserved in alcohol
introduces a source of error in two measurements — total length and
length of tail. According to the strength of the preservative fluid, both
body and tail are to a varying degree shrunk or relaxed, so that consid-
erable discrepancies in the averages of specimens taken at different
localities by different collectors may result. In general, it is probable
that these two measurements as given in the tables are a trifle shorter
than they would have been if taken from fresh material.

It is unfortunate that detailed measurements of individuals can not
be published, since averages are of use for comparison with averages
only, and it often happens that a single specimen must be identified.
Averages, moreover, give no indication of the normal range of indi-
vidual variation at a particular locality.

'A detailed accouut of the migration of bats on Cape Cod was iiublislied in Science,
N, S., V, No. 118, pp. 541-543, April 2, 1897.

12 NORTH AMERICAN FAUNA.

ILLUSTRATIONS.

Tlie illustrations in this paper are reproductions of pen-and-ink
drawings made under iny constant supervision bj'' Mr. Frank Miiller.
Special difficulty lias been encountered in obtaining satisfactory repre-
sentations of the external ear and of the crowns of the teeth.

The ears of alcoholic specimens are generally sufficiently altered in
form, by pressure and by the action of the preservative tluid, to
retain only approximately the appearance which they had in the living
animal. This is especially the case with such large-eared species as
AnirozouH imlUdns, Corynorhinus viacrotis, Myotis erotis, and others.
In the impossibility of reproducing their original appearance, it has
been thought best to represent the ears in a uniform but somewhat
unnatural i)osition, with the conch flattened and the external basal
lobe tui-ued outward. This will account for the apparently undue
width of certain drawings.

The crown views of the teeth were first sketched with the aid of a
camera lucida and afterwards corrected and finished by the use of hand
lenses. The great difficulty in obtaining accurate and uniform results
arose from the impossibility of kee])ing specimens in exactly com-
parable positions and from the considerable changes in outline result-
ing from every slight variation in the angle of vision. Therefore the
drawings are not wholly satisfactory. They are published, however,
in the belief that, such as they are, they nmy help to an understanding
of the characters of the species.

NOMENCLATURE OF NORTH AMERICAN YESPERTILIONIDyE.

To arrive at final conclusions in regard to the nomenclature of the
VesjiertUionuJw of North America, it will be necessary to consider in
detail all names that have been based on those members of the groui)
that inhabit the region in question, and also a few based on allied Old
World species. The names may best be taken up alphabetically.

1. Generic aud Subgeneric Names.

Adelonycteris H. Allen, 1892 ( Proc. Acad. Js^at. Sci., Phila., 1891, p. 4G6,
Jan. 19, 1892), was proposed as a substitute for Vesjferus Keys. & Bias.,
preoccupied in Entomology by Vesperus Latreille, 1829. The name is,
however, a synonym of YespertiUo Linmeus, 1758, Eptesicus Eatinesque,
1829, and also of Cneph(vus Kaup, 1829.

Aeorestes Fitzinger, 1870 (Sitzungsber. Math.-Xat. CI. K. Akad.Wiss.,
Wien, LXII, Abth., I, i)p. 427-430), is a synonym of Myotis Kaup, 1829.
The group included three South American species, Myotis viUosissimus,
M. nif/ricans, and M. albescens.

Antrozous H.Allen, 1862 (Proc. Acad. Xat. Sci. Phila., p. 248), is the
only generic name based on Vespertilio paUidus Le Conte.

NOMENCLATURE. 13

Atalapha Eafinesque, 1814 (Precis des Decouv. et Travaux Somio-
lofticiues, p. 12), is clearly based ou a Sicilian bat.' The use of the
name for a geuus coufiiied to America is therefore impossible.

Brachyotus Kolenati, 1856 (Allgem. Deutsch. Naturhist. Zeitg., Dres-
den, Xene Folge, II, pp. l.'U, 174-177), is a subgeiieric name based on
three European species of ' Vespertilio'' {my.stacinus, dauhentonii, and
dasycneme) with ears shorter than head.

Cateorus Kolenati, 1856 (Allgem. Dentsch. jSTaturhist. Zeitg., Dresden,
Xeue Folge, II, pp. 131, 102-103), a subgeneric name based ou '■Yefiperus'
serotinus, is a synonym of Vespertilio Linufeus.

Cnephaeus Kaup, 1829 (Skizzirte Entw.-Gesch. u. Natiirl. Syst. d.
Europ. Thierw.. Ister Theil, p. 103), is a generic name based on Yesper-
tilio serotinus Schreber, a species congeneric with Ves2)ertilio fuscus of
America. The name is a synonym of Vespertilio.

Cnephaiophilus Fitzinger, 1870 (Sitzungsber. K. Akad. Wiss., Wien,
LXII, Abth. I, p. 81), is a genus composed of very heterogeneous ele-
ments among which no type is mentioned. The species referred to it
are macellus ('Borneo'), peUucidus ('S. E. Asia, Philippines'), ferriigi-
?^e^^s•('Mittel Amerika, Surinam'), and the North American noctivaf/ans.
Whether or not the name may be available for some of the other spe-
cies, it certainly is not for the one which comes within the limits of the
present paper, since this was already provided with the generic name
Lasioni/eteris.

Comastes Fitzinger, 1870 (Sitzungsber. Math.-Xat. CI. K. Akad.
Wiss., Wien, LXII, Abth. I, p. 5Go), is a synonym of Myotis Kaup,
unless it may eventually be shown that the sijecies on which it was
based, capaccinii, megapodius, dasycneme and limnopMlus, are subgeu-
erically distinct from Myotis myotis.

Corynorhinus II. Allen, 1865 (Proc. Acad. Xat. Sci. Phila., p. 173),
proposed as a generic name for Flecotus maerotis Le Coute and F.
toirnsendi Cooper, is the only available name for the group of which
CorynorJiinus maerotis is the only known species.

Dasypterus Peters, 1871 (Monatsber. K. Akad. Wiss., Berlin, 1870, p.
912, published 187J ), was established as a subgenus of Atalapha { = Lasi-
tirns) to contain the species intermedia, eyregia, ega, and caudata. It
has recently been raised to full generic rank by Dr. Harrison Allen.

Eptesicus Eafinesque, 1820 (Annals of Nature, p. 2), originally cou-

'II. G. ATALAPHA (Chanve-sonris). Incisivesnullesaux deux nii'ichoires, canines
et uiachelieres aigues: aucuue crete sur le nez. queue lU'esqu'entierement unie aux
membranes.

2. Atalaplia sicitla. — Oreilles de la longueur de la tete, et auriculoes, une verrne
sous la ir-vre inferieure; corps roux brnnutre en dessus, roux cendro en dessous, ailes
et museau noiratre, queue saillaute par une pointe obtuse. — Obs. J'ai observo cette
espece en Sicile, elle ditt'cre de V Atalapha ameruona (Vexprrtilio tiorehoracenxis Lin.),
autre espece du menie genre, par ees deux premiers et son dernier caractere.

14 NORTH AMERICAN FAUNA.

taiued two species, E. melanopn aud E. mydas.^ Eptesimis melanops is
without doubt the Yespertilio fuscus of Beauvois. E. mydas, however,
cau not be identified (seep. 32). The first species must therefore be
taken as the tyi)e. Since this species is congeneric with Vespertilio
murinus Linna'us {=Vesperiigo discolor batterer), the type of the genus
Vespertilio, the name Eptesicns is a synonym of Vespertilio.

Euderma IT. Allen, 1892 (Proc. Acad. Nat. Sci. Phila., 1891, p. 467,
published Jan. 10, 1892), is the tenable name for the genus of which
Eistiotns maculatus J. A. Allen is the type and only known species.

Histiotus Gervais, 1855 (Exped. Comte de Castelnau Am.du Sud, Zool.,
Mamniif., p. 77, PI. XII), was based on the South American Plecotus
velatus of Geofifroy. Euderma maculatnm was originally described as a
member of this genus, the name of which has not otherwise appeared
in the literature of North American Vespertilionida\

Hypexodon Eatiuesque, 1819 (Journal de Physique, de Chlmie, d'His-
toire naturelle et des Arts, LXXXVIII, p. 417), can not be identified
with any known group of bats. The characters which Rafinesque
assigns to the type species^ may be those of a mutilated aud distorted
specimen of someof the small species of Xycticeius, Pipistrellvs, ovMyotis.

Hypsugo Kolenati, 1856 (Allgem. Deutsch. Xaturhist. Zeitg., Dres-
den, Xeue Folge, II, pp. 131, 167-l(i9), is a synonym of Pipistrellus
Kaup. It was based on ' Vesperugo'' maurvs Blasius aud ' V. ' Tcrascheni-
nil'oivii Eversmann.

Isotus Kolenati, 1856 (Allgem. Deutsch. Naturhist. Zeitg., Dresden,
Xeue Folge, II, pp. 131, 177-179), is a subgeneric name based on two
European species of ' Vespertilio ' {nattercri and eiliatus) which have the
ear about equal in length to the head. It is of course a synonym of
Myotis Kauj), 1829, and of Selysius Bonaparte, 1841.

Lasionycteris Peters, 1865 (Monatsber. K. Preuss. Akad.Wiss., Berlin,
1865, p. 648), is the first name proposed for the genus of which Vesper-
tilio noctivagnns Le Conte is the only known species.

Lasiurus Gray, 1831 (Zoological Miscellany, Xo. 1, p. 38), is the first

'The original diagnosis of the genus Eptesicns is as follows:

" I. N. G. EPTESICUS. Four acute foie-teeth to the upper jaw, in two equal ])air8,
separated by a great interval and a large flat wart, each pair has two unequal
teeth, the outside tooth is much larger and unequally bifid, the outside one much
larger, inside tooth small and entire. Six fore-teeth to the lower jaw, equal very
small, close and truncate. Canine teeth very sharp, curved and long. Grinders
unequally trifid. Snout plain, nose without appendages. Ears sei>arated, auricu-
lated. Tail mncronate. — This genus apjiears to differ from all those of Geoffroy and
Cuvier, among the extensive tribe of Bats. The name means house-flyer."

-1. Nouveau genre. HYPEXODON. (Chauvc-sourits.) Museau nu ; narines rondes,
saillantes; incisives snporienres nulles, 6 inferieures omarginoes, uue verrue ala base
extdrieure des canines inferieures. Queue eugagee dans la membrane. Le reste
comme le genre VesperlU'io. — I espece H. iiu/stax, entiJ'rement fauve, dessus de la tcte
brun, ailes et membranes nolres, queue mucrouee, des moustaches, oreilles brunes
auriculecs, nervures inferieures et transversales; longueur totale, 3 pouces, dont la
queue 2 pouces. En Kentucky.

NOMENCLATURE. 15

name based on the bats of the American genus commonly but wrongly
called Atalapha. It was introduced as follows : " The bats, the Vesper-
tilionc.s of Geoffroy, might for convenience be divided into three genera,
the true bats, VesperUUo * * *, the Facliyotus * * *, and the
hairy tailed species of America (Za-s'/ ;n- its)." As the only hairy-tailed
American bats known in 1838 were members of the modern genus
Lasiurus, this brief statement may be taken as a definite indication of
the author's meaning. In 1838 Gray referred the species pruinosus
{=cinen'U.s), lashiriis (= borealis), and hlos-sevillei (= horealis,Jif1e'Dobsou)
to the group, which he then regarded as a subgenus or section of
ScotophiluH (Mag. Zool. & Bot., II, p. 4!)8, Edinburgh, 1838).

Marsipolaemus Peters, 1872 (Monatsber. k. Preuss. Akad. Wiss., Berlin,
p. 200), was proposed in a subgeneric sense for a Mexican bat, Vesperns
aJhigularis Peters, about the size of Ve,s2)ertilio fuscus, with the denti-
tion of that species, but with the outer border of the ear continuous
with a fold of skin which extends back from the corner of the mouth,
under and behind which a distinct pocket is formed. I have never
seen this bat, and am unable to say what value is to be placed on the
characters described. (See \^. 101.)

Meteorus Kolenati, 1856 (Allgeni. Deutsch. Katurhist. Zeitg., Dresden,
Neue Folge, II, pp. 131, 107-101>), is a synonym of Yespertilio Linn;eus.
It was proposed as a subgenus of '•Yesperus'' to include the species
nilssimi, (lificojor, .s((vii, Ieucip2)e, and arisiippe.

Myotis Kanp, 1829 (Skizzirte Entw. Gesch. u. Natiirl. Syst. der Euroj).
Thierw., Ister Theil, p. 106), is the first name based on the large, long-
eared, thirty- eight toothed bat wrongly called YespertUio miirinus by
Schreber.^ It is therefore the tenable name for the genus of which this
animal is the type. As the Yespertilio murinus of Schreber is not the
Yespertilio murinns of Linna'us, another specific name must be applied
to the former. The name w^/oif/s Bechstein- is available for this i>ur-
pose. Hence the Yespertilio murinus of Schreber and of European
writers in general nmst stand as Myotis myotis (Bechstein).

Nannugo Kolenati, 1856 ( Allgem, Deutsch. Naturhist. Zeitg., Dresden,
Neue Folge, II, pp. 131, 169-172), is a synonym of Pijnstrell us Kanp,
1829. It was proposed as a subgenus of ^Yesperugo'' to include the
European species jnpistrellus, IcuhlH, and nattereri.

Noctula Bonaparte, 1837 (Iconografia Fauna Italica, I, fasc. XXI,
under Yespertilio alcythoe), based on Yespertilio serotinus Schreber is a
synonym of Yespertilio Linnteus.

Nycticeius Rafinesque, 1819 (Journal de Physique, de Chimie, d^His-
toire Xaturelle et des Arts, LXXXVIII, p. 417), contained two species,

'Kaup says: " Fledermihise von riesenmiissiger Grosse, mit nacktem Gesiclit,
getreuiiteu, kopfslangeii Ohreii, langeu lanzettformigen Ohrendeckelu, iind 38
Ziihnen."

-resperiilio mi/otis Beclisteiu, Gemeiniiiitz. Naturgesch. Deutschlauds, Bd. I, p.
1145, 1791 (fide Blasius).

16 NORTH AMERICAN FAUNA.

N. humeralis Eaf. aud N. tesselatus Bat JSTotliiug in the description'
indicates which of these the author considered as the type. Nycticeiiis
tesselatus Raf. is Lasiurns horealis (Miiller), and ^. humeralis may with
some degree of probability be identified with the small brown bat more
generally known as Nycticejus crepuscularis Le Conte,- There is cer
tainly nothing in the diagnosis of the genus or in the description of
yespertlUo humeralis previously published in the American Monthly
Magazine that precludes this possibility, while the size, the number of
incisors, and the naked uropatagium point directly toward it. As
horealis was removed to tlie genus Lasiurus by Gray in 1838, humeralis
becomes the type of Nycticeius. The orthography of this name has had
several emendations, as Xycticcus, Kycticejus, Xyeticea, and Nycticeyx.

Nyctilestes Marsh, 1872 (Amer. Journ. Sci. & Arts, 3d ser., IV, p.
215), is a fossil genus based on part of a lower jaw and molars from
Eocene or Lower Miocene strata near Henrys Fork, Wyoming. The
remains present no characters to distinguish them generically from
Vespertilio. Only one species, Nyctilestes serotinus, has been described.

Nyctitherinm Marsh, 1872 (Amer. Journ. Sci. & Arts, 3d ser., IV. p.
127), is a genus based on the fragments of two lower jaws found with
teeth in place, from Tertiary strata at Grizzly Bnttes, Wyoming. The
original description indicates no characters by which these teeth may be
distinguished from those of small species of Fipistrellus or Vespertilio.

Nystactes Kaup, 1829 (Skizzirte Entw.-Gesch. u. Natiirl. Syst. der
Europ. Thierw., Ister Theil, p. 108), based on Vespertilio bechsteinii
Leisler is strictly synonymous with the same author's Myotis?

Pachyotus Gray, 1831 (Zool. Misc., ISo. 1, p. 38), was first used as the
name for a genus made by the combination of Xycticeius aud !Scotophi-
lus. Later (Mag. Zool. & Bot., II, p. 498, 1838) Gray transferred it to
Vespertilio villosissimus Geoffroy in a subgeueric sense. The name is
of course untenable.*

Fipistrellus Kaup, 1829 (Skizzirte Entw.-Gesch. n, Xatiirl. Syst. der
Europ. Thierw., Ister Theil, p. 98). This name was based on Vespertilio
pipistrellus Schreber, a species strictly congeneric with the ' Vesper ugo

12. NYCTICEIUS. (Chaiive-souris.) Diftere du genre pr^cMent iHypexodon'] par
2 iucisives superieures separees par un graucl iutervalle, accoldes aiix cauiues et h
creuelures aigui-s, 6 incisives iuf6rieures trouqu(?es. point de verrues aux cauiues.—
Ce genre coutient an moius 2 espi'ces, X. humeralis et N. tesselatus, que j'ai dojii
d^crits dans VAmeriva)) Monthly Mafjaziue, sous la denomination geuerique Vespertilio,
avec plusieurs autres uouvelles espt'ces de ces contrces.

•See Tliouias, Anu. & Mag. Nat. Hist., 1891, 528.

• Kaup says : " Fledermiiuse luit sehr laugen getrennteu Ohren, langem zugespitzem
Ohreudeckel, 38 Ziihncn uud spitzmausiilinlicliem Kiissel."

■■The original reference is as follows: ''The bats, the VespertiJiones of Geott'roy,
might for convenience be divided into three genera, the true bats, Vespertilio, with
thin ears and membranes aud a hairy face, the Pachyotus, with thick ears and mem-
branes and bald swollen cheeks, including the genera Nycticejus and Scotophilus, aud
the hairy-tailed species of America {Lasiurus)."

NOMENCLATURE. 1 7

georgianus'' of tlie United States, It autedates the name Vesx)erugo by
exactlj' ten years,

Plecotus Geoflfroy, 1818 ' (Description de I'lOgypte, Mamniiferes, p. 112),
included tbree species, TOreillaid de Daubeuton,' 'la barbastelle,'and
a new species from Timor.^

As no American bats are congeneric with the species originally
included in this genus, the name can not be used for any of the genera
now under consideration. It has been applied to the species of Cory-
norhinus.

Rhogeessa H. Allen, 1866 (Proc. Acad. Nat. Sci. Phila., p. 285), was
proposed as a genus to contain the species B. parvnla II. Allen and R.
tumida H. Allen. The group, whose validity has not been questioned,
has received varying treatment at the hands of difl'ereiit writers.
Dobson placed it as a subgenus under ' Vesper ugo,^ but Thomas has
recently pointed out its close relationship to Xyeticeius. The latter
disposition appears to be the more natural.

The name has been amended to Bhogoessa by Marschall (Nomenclator
Zoologicus, Mamm., p. 11, 1873).

Scotophilns Leach, 1821 (Trans. Linn. Soc. London, XIII, pt. l,p. 00),
type iS. Jxuhlii Leach, is a genus peculiar to the Old World, where it
apparently replaces the Lasiiirus of America. Jt is mentioned here
merely because the mime has been used for the North American species
of Laslurus, Vespertilio, Lasionyeteris, and npisfrellns at times when
these bats were supposed to be congeneric with Old World species.

Selysius P.onaparte, 1841 (Iconografia Fauna Italica, I, Introduzione
[p. .'>] ), is a synonym of Myotis Kaup, 1829. It was based on the
common European YesperUlio mystacmus of Leisler.

Synotus Keyserling and Blasius, 1839 (Wiegmann's Archiv f. Natur.
geschichte, r)ter Jahrgang, Bd. I, pp. .'>0.">, 30(>), was based on the bar-
bastelle, a European bat representing a genus not known to occur in
America. The mime, however, has been applied to the American genus
afterwards called Gorynofhiniis. It is antedated by Barbastel la Grtky,
1821 (London Medical Kepository, XV, \}. 309. Type Vesper tilio barbas-
tellus Schreber).

Taphozous (leoli'roy, 1818' (Description de l'l5gypte, Mammiferes, p.
113), based on ' Le lerot-volant' and 'le V. lepturus,^ which are without
representatives in America. The red bat {LasiKvui'i borenUs) was, how-
ever, included in this genus by ( Jodnuin under the name Taphozous ru/us.-^

' See Sherborn, Proc. Zool. Soc. Londou, 1897, p. 288.

4 2 5-5

-Dents incisives g; canines ., ; niolaires ,,_,.. 'Nez simple et saillauf ; clianfrein

large el mciplat. Oreilles plus (jrandcs que la tote, et rdunies; oieilloii iut<_^rieur.
Membrane inteifcmorale ctendue et a angle saillaut. Queue longue et toute entiero
enveloppt'c.

Obs. Les trois espices do ce genre sont, Foreillard do Danbenton, la barbastello
et uue noiivelle osptcc de Timor.
^Fauua Americana, p. 23, 1825.
2772 No. 13 2

18 NORTH AMERICAN FAUNA.

Fitzinger ' refers to a ' Taphozous hracJimanus Ciodmau ' amoug the syn-
oiiyins of Last urns ^vk/ks^ (= horealis). This name, however, I have been
unable to And in any of Godnian's writings.

Vesperides Coues, 1875 (in Coues and Yarrow, Zool. of Wheeler's
Exped., p. 83), was proposed as a subgenus of YesperUUo based on Yes-
IwriUio noetivagans Le Conte. The name is antedated by Lasionycteris
refers, 18G5.

Vespertilio LinucTUS, 1758 (Syst. Nat., 10th ed., I, p. 31), eontaiued
seven species: vampiirus, spectrum, persiyiciUatus, spasma, leporinus,
auritus, and murinus. These have all been removed to other genera, as
follows: vampyrus to Ptcropus in 1702 (Brissou, llegn. Anim., ed. II,
pp. 13, 153), leporinus to NoctiUo in 1700 (Linna'us, Syst. Nat., 12th ed.,
p. 88), spasma to Mcgaderma in 1810 (Geoffroy, Ann. Mus. d'Hist. Nat.,
XV, p. 197), anritus to Plecotus in 1818 (Geoffroy, Descript. de I'Egyjjte,
Mammiferes, p. 112), murinus'^ to Eptesicns in 1820 (Eafines(iue, Annals
of Nature, 1820, p. 2), perspicillatus to Artihens in 1821 (Leach, Trans.
Linn. Soc. London, XIII, p. 75), and spectrum to Vumpyrus in 1821
(Leach, Trans. Linn. Soc. London, XIII, p. 79).

The only European species are anritus and murinus, one of which
nuist therefore become the ty])e of the genus. The species anritus was
removed to the genus Plecotus by Geottroy in 18! 8, leaving murinus as
tyi)e of the genus YcspertiUo. The YespcrtHio mnriuns of Linnanis is,
however, a totally different animal from the bat afterwards described
under the same name by Schreber. To understand the case fully it is
necessary to go back to the first and second editions of Linuicus's
Fauna Suecica. In the first ho records only one bat, the 'Laderlapp,'
'Fliidermus' or 'Nattblacka,' Yespertilio caudatus, naso oreque simplici
(No. 18, p. 7, 174G). In the second edition he mentions two, Y. caudatus,
naso oreque simplici, anriculis duplicatis, capite majoribus, and Y. cau-
datus, naso oreque simplici, anriculis capite minoribus (No. 2, pp. 1, 2,
1701). In the tenth edition of the Systema Naturae these had been
given binomial names, Yespertilio auritns and T'.w?/ri/y?(,s, respectively.
The account of the teeth of the latter in the second edition of Fauna
Suecica is as follows : ^

Deutes priiiiores superiores 6, aciiti tlistantes.
iiiferiores 4, acuti coiitigiii.
Laniarii superiores 2, anteriore innjore.

inferiores 3, antico iiiaximo.
Molares utriuque 3, tricuspidati.

' Sitziingsber. K. Akad. Wiss. Wien, LXII, Iste Abtli., p. 402, 1870.

"Altliougli Rafiuesque did not actually place the species murinus iu the genus
JEptesicus he based the latter ou a strictly cougouerio forui.

»In tbcfirst edition the dental formula is the same, except that the lower iucisors
are said to be five in number, an error corrected iu the second editiou.

NOMENCLATURE. 19

It therefore appears that the Vespertilio murimis of Linnaeus is a bat
with ears shorter than the liead, and with the deutal formula:
. 2-21 i_i i_i ;3_3

*' 3^ '^'1-1 '^'"'2-2 ''"'313 = ^^-

The only common Scandinavian bats which combine these characters
are the two usually known as Vespernyo nilssoni and Vesperuyo discolor.
To these strictly congeneric European species and their exotic repre-
sentatives the generic name Vespei't'dio must be applied, regardless of
its long misuse for a different genus.

The current misidentification of Linuaeus's Vespertilio murinus has
been recognized by at least three writers on European bats, Nilsson,
Blasius, and Lilljeborg. Nilsson^ discusses the matter at considerable
length and arrives at the conclusion that the name murinus nuist be
substituted for discolor^ while the bat commonly known as murinus
must take the si^ecific name myotis Bechstein. As this author unites
the genera ' Vesperugo ' and ' Vespertilio^^ he has nothing to say in regard
to tlie validity of the generic names used by Keyserling and Blasius.

Blasius' regarded Nilssou's identitication of Vespertilio uinrinus a^s
doubtful, though he admitted that the animal described by Linnaeus
under that name could not have been the one generally called Vesper-
tilio muriuKs by European authors at large, lie therefore reasoned
that Linua?us's name might be disregarded as undeterminable and in
no way invalidating Schrcber's later application.

Lilljeborg alone questioned the tenability of the generic name Vesper-
tilio for the thirty-eight toothed bats of Europe.^ He says:

* * ■ As regards modifying the Linuii'iiu _i;cueric name Vespertilio, it may be
viri!;ed that Liuna-us did not include in it any of the species referred to it by Keyser-
liiij; and Blasius. Furtber, it would have been more correct to apply the name
respcrtilio to the preceding genus [' f'cspcn<(jo'], since one of the si)ecies included in
the genus )»y Liuun-us ( i'espertilio murinus) agrees, in all important characters at
least, with the genus mentioned, as shown above. As, however, the niodihcatiou of
the name introduced by Keyserling and Blasius has become time-sanctioned, it will
be retained, although we consider the objections against it reasonable.'

Vesperugo Keyserling and Blasius, 1839 ( Wiegmann's Archiv f. Natur-
gesch., 5ter Jahrgaug, Bd, I, p. 312), was proposed as a genus to contain
the following species up to that time commonly associated with Vesper-

' In Linna'us's statement the figures 4 and 6 are evidently transposed.

'Skandinavisk Fauna, I, Diiggdjuren, 2ded., 1847, pp. 17-20.

'Naturgesch. d. Siiugetbiere Deutschlands, pp, 74, 84, 1857.

SSveriges och Norges Ryggradsdjur, I, Diiggdjuren, p. 144, footnote, 1874.

■'■'* * * I afsecnde pa tilliimpningen hlir af det Linneanska genus-namnet
Vespertilio, kan deremot inviludas, att Liun6 icke uti detta genus upptagit en enda af
de arter, som Keyserling & Blasius derunder beskrifvit, och att det hade varit
riittare, att auviinda detta namn ffir furegaende sliigte [' fesper;;*;©'], emedau en
af de af Linne uti si. Vespertilio npptagna arterna — Vespertilio murinits Lin. —
atmin^,toue till hufvudsaklig del, euligt hvad ofvan blifvit antVirdt tillhur niimde
sliigte. Da cmellertid den af Keyserling &- Blasius infiirda tilliimpningen af
naoinet vuuuit hiifd, vilja vi bibehalla den, ehuruviause inviiudningen vara befogad.

20 NORTH AMERICAN FAUNA.

tilio: serotinus, dhc(>l(>t\ iiils.soni, sdnii, leHcippc, (iri.stippe, noctula, leishri^
lultliij alboUmhatus, nathmii, iiiid pipistrellus. The first six were [)lace(l
iu tlie new subgenus Ves2)erus, the others in the subgenus Vesperiujo.
Hence the ty[)e must be a member of the second group. This group,
however, ccmtaiiis two modern genera, the first represented by the spe
cies itoctuht and leisleri, the second by JctiJilii, '■ albolimbatus'' {^^Jcuhlii,
fide Dobsou), '• natluisW [^=abramus, fide Dobson), and pipistreJliis.
These had ah^eady been named Ptrrygi.stes and PipistrelluH, respectively,
by Ivaup iu ISl*!). Hence Vespenujo is untenable in any connectiou.

Vesperus Keyserling and Blasius, 1839 (Wiegraann's Archiv f. Natur-
gesch., oter Jahrgang, Bd. I, p. 31.")), pr()[)Osed as a subgenus of M'c.s-
pcn((jo^ to incUide the species svrotiiiK.s, discolor, nilssoni, savii, leueippc,
and aristippe, is antedated by Cnephwus Kaup, 1829, Eplesieus Eafines-
(pie, 1820, and Vcspertilio Linnu'us, 1 TaS. It is moreover preoccupied iu
Entomology by Vesperus Latreille, 1829.

2. Specific and Subspecific Names.

Affinis (Vespertilio). H. Allen, Mouogr. Bats N. Am,, p. 53, 1864.
The type of Dr. Harri.son Allen's Vespertilio affinis, now in the United
States National JNIuseuin, proves to be a tyi)ical example of Myotis
luci/iKjus. It is therefore in no way related to the VespertUio nitidm
or V, (ilbrsceibs of Dr. Allen's second monograph.

Albescens (Vespertilio). E. (rcoftVoy, Ann. Mus. d'llist. Nat., Paris,
VIII, p. 204, 1806. This is a South American species of Vli//o<u, i)roba-
bly closely related to M.velifer (J. A. Allen). The measurements given
by Azara and <|uoted in the original descrii)tion are: Total length, 80
mm.; tail, 33; extent of wings, 235; ear, 14. The name albescens has
been used by Dr. Harrison Allen for Myotis yumanensis, M. evotis, M.
cali/oruicus {^Vespertilio albescens vielanorh inns''), M. reli/er, M. titysa-
nodes (under M..velifer),imd M. hicifnyus (' Vespertilio albescens affinis''),
which he unites as subspecies.

Albigularis (Vesperus). Peters, Monatsber. K. Preuss. Akad. Wiss.,
Berlin, p. 2(30, 1872, Vesperlilio aibigularis (Peters) is the ty[)e of the
subgenus MarsipoUvmus. The characters given in the original descrii)-
tion indicate a well marked species, with which, however, I am wholly
unacqmxinted. The type was collected in ftlexico.

Alleni (Rhogeessa). Thomas, Ann. & Mag. Nat, Hist., (Uh ser,, X,
p. 477, 1892. This is the only name for this species.

Americana (Atalapha). Kafinesque, Precis des Decouv. Somiologiques,
p. 12, 1814, This is a synonym of Lasinrus borealis (Miiller), though
proi)erly speakiug the name is a nomen nudum (see p. 10(5).

Arquatus (Vespertilio). Say, Long's Expedition to the Kocky Moun-
tains, I, p, 107, footnote, 1823. Tlie description clearly indicates Ves-
pertilio fuscus Beauvois.

Auduboni (Vespertilio). Harlan, Featherstonehaugh's Monthly Ameri-
can Journal of Geology and Natural History, I, p. 220, PI. II, November,
1831, Both description and plate indicate the silver-haired bat.

NOMKNCLATURPL 21

Austroriparius (Vespertilio lucifugus). lihoads, Proc. Acad. Nat. Soi.
Pbila., 1). 227, May, 1897. Vc.sjX'rtUio Jucifngus austroripariuH Rlioads
is a synonym of Myolis lud/w/ns (Le Coiite). The type, a two-thirds
grown young from Tarpon Si)ri'ngs, Florida, shows nuinerons characters
by which it may be distinguished from northern adults, but the full
grown topotypes are, as originally determined by Dr. Harrison Allen
(see IMioads, I.e.), indistinguishable from northern si)ecimensof /*<c//«-
(jKs that have been immersed in alcohol for a similar period. Even if it
Avere assumed that the Tarpon Springs bat difilerod in some way not
now discoverable from the ' IncifiKjus of North Carolina and northward,'
there could be little doubt that the southern form was the one originally
described by Le Conte. (See page G3).

Belli! (ScotopMlus). (iray. List Si)ec. IVlamm. Brit. Mus., p. 30, 1843.
Scotophilus hellii (Iray is anoinen nudum probably based on one of the
West Indian forms of Vespertilio fiiscus. Gray's account is as follows:
"Bell's Bat. Scotophilus Bellii. ^/Inspirits. West Indies. — Pre
sented by Thomas Bell, Esq., F. R. S."

Borealis (Vespertilio). Midler, Natursyst. Suppl., p. 21, 1776. Midler's
Vespertilio borealis is the first name based on the red bat, Lasiurus
borealis.

Brevirostris (Vespertilio). IVIaximilian, Wiegmann's Archiv. f. Natur-
geschichte, 1801, Bd. I, p. 19.5. VespertUio brevirostris of Maximilian is
probably Myotis lucifu(/ns (Le (\)nte). The original measurements
are: Total length, 3"; extent, 9" 4'"; ear from crown, 5J'"; tragus, Lj'".

Calcaratus (Vespertilio). Rafinesque, American Monthly Magazine,
III, ]). 445, 1818. No known bat agrees with the description of Rati-
nesque's Vespertilio calearatus^ which is as follows: "Tail one-third,
body dark brown above, dark fallow beneath, wings black, shafts rose-
coloured, a spur at the inner side of the elbow, hind feet black. Length
4 inches, breadth 12."

Californicus (Vespertilio). And. & Bachm., Journ. Acad. Nat. Sci.
Phila., VIII, Pt. II, p. 285, 1842. This is the earliest name based on
the small western bat commonly known as Vespertilio nitidiis H. Allen.
The original description is as follows : '

r. ralifoi-niciis (C'aliforni.in bat). — V. fnsco Iiitosceiis, vellere loiijioet iiiolli; trago
lnii<j;itu(lino diniidiuni anris cxccdcntc.

('aliforiiian hat. — Witb long silky bair.s; tragus more tban balf tbo Icugtb of the
ear; cobjr light yellowish brown.

Descripfiou. — Anterior upper fore teeth bilol)ate. Head small; nose sharp; ears
of moderate .lize, erect, rather narrow, and pointed. Trat/iis Ihuar, atleuiiatcd. \\'ings
of moderate length, which together with the ears are naked, [nterfemoral mcmhranc
Willi a. few scattered liairs; feet small; nails slightly hooked. Tail projecting a little
beyond the intei'fomoral membrane.

Color. — The peliige, which is uniisualhj lonf/ for the size of thehody, and very soft and
glossy, is, on the upper surface, dark pluml>eous from the base, and hroadly tipt with

' I have italicized statements specially applicable to ' f. nitidus.'

22 NORTH AMERICAN FAUNA.

lif/hf }iellowisli hrown; on the under siirfiico the color is a little darker, owing to the

outer extremities of the hairs being more narrowly edged with the prevailing color

on the back, exhibiting the darker shades beneath. The enrs and tragus are Idack-

ish — the nose, chin, wings, and interfemoral membrane dark brown.

Hah. — We have obtained but a single specimen, which was captured at California.

2-2 1-1

DenCition. — Incisors^. Canines :. |.

DimenHons. — Length of head and body, 1 inch 7 lines [40 mm.] ; length of tail, 1
inch 5 lines [35.8] ; length of spread, 7 inches C> lines [190] ; height of ear ]>nHteriorly,
3 lines [6.35] ; height of tragus, 2 lines [3.8].

The only other small bats known to occur in California are Pipistrellus
hesperus, Myotis thysaiiodes, M. ynmanensis, M. evotiSj and M. lucifiu/ns
lonyicrus. That YespertUio californicus can not be Pipistretliis hesperus
is shown by the description of the tragus. From Myotis tliysanodes it is
separated by its small size and nnfriiiged interfemoral membrane; from
M. yumanensis by its small feet; from M. evofis by its short ears, and
from M. lucifugus longicrus by its light color and small size. Myotis
thysanodes and 71/, lt(cifiigns lonr/icrtis are moreover comparatively rare
bats in California, while ' YespertUio nitidns'' is one of the most common
and universally distributed species.

Carolii (Vespertilio). Temminck, Monographies de Mammal., II, p.
237 (13me Monogr.), 1835-41. The YespertUio enrol H of Temminck is
without doubt Myotis lucifugus (Le Conte). That it is a Myotis is shown
by the number of teeth, six molars in each jaw, while that it is not M.
svhulatus, the only other s[)ecies known to occur in the vicinity of Phil-
adelphia or New York, is shown by the short ear, 11.5 mm. in length.'

Carolinensis (Vespertilio). Geoffroy, Ann. du Mus. d'llist. Nat., Paris,
VIII, p. 193, 1806.-

This species is YespertUio fuscus Beauvois. Dr. Harrison Allen in

'The essential part of the original description is as follows:

"Taille et formes de notro pipistrellc, mais les oreiiles plus longues. * * »
oreilles uK^diocres, ovoides, un pen docoupoes a Icnr herd exterleur, sans lobe on
prolongement en avant; tragus en feiiille de saule * * *. Dents iucisives 4 par
paire en haut et 6 en bas; molaires 6 partout; les deux premiferes fausses molaires
de la machoire suporieure tri's petites, courtes et pointues.

"Pelage bicnlore partout. Jones, cotes du cou et toutes los parties snpdrieures
d'un brun-roussfitre a base des polls noirs ; en dcssous d'un blanc jannatre h la jtoiutc
et brun-fonco a la base * " *.

"Longueur totale 3 ponces 5 lignes, dont la queue prend 1 ponce 4 lignes; enver-
gure 8 ponces 0 lignes; autibrachinm 1 ponce 4 lignes; hauteur de I'oreille depnis le
crane jusqu'an bout 5 lignes; * * *.

"Pairie, L'Am(5rique sci)tentri()nale, dans les environs de Philadelpbie et de New-
York."

'^The original description is as ft)llow8:

"2. Vesp[ertiUo1 carolinensis. Le vespertilion de la Caroline est nioius grand (|ne
Iepr6c6deut [' r. mnrinns' ], mais d'ailieurs il Ini ressemble beaucoup. II a ses oreilles
et oreillons de meme forme et de mcmc dimension relative; son jioil est anssi dedeux
couleurs, cendre-noirati-e d'abord et brnn-marron h la pointe. L'extrcmitt? des poils
est en dessous d'un jaun(^ tirant sur le ventre; enlin les oreilles sont garnies de poils
dans presquo L'l moitie de leui- longnenr. <t l-i qurue a nue petite portion (|ni u'est
)ias euveloppee par la membrane iuterfemorale. Ces considerations reunies a cellea

NOMENCLATURE. 23

liis recent monograph lias applied the uaiue carolinensis to the Georgian
bat {Fipistrellns subjfavus), but there is no reason to doubt that Geof-
froy's animal was the large brown bat. The head and skull are both
figured, the former on PI. I, the latter on PI. II. These are only a trifle
smaller than the head and skull of Ves2)ertilio serotinus figured on the
same plates, and very much larger than the figures of the head and
skull of Pipistrellus iripisireUus^ a species of about the same size as F.
suhjiavus. The teeth are very indistinctly shown in the figure, but in
the two copies which I have examined^ I can find no indication of the
second upper premolar of J'ipistrelhis.

Chrysonotus (Vespertilio). J, A. Allen, Bull. Am. Mus, Nat. Hist.,
N. Y., VIII, p. 240, November 21, 1896. VesperHUo chrysonotus J. A.
Allen, from Kinney Ranch, Wyoming, is a pale example of Myotis
evotis (H. Allen), with mutilated tail. (See p. 80.)

Ciliolabrum (Vespertilio). Merriam, Proc. Biol. Soc. Washington, IV,
p. I, 1886. Vespertilio ciliolabrum, Merriam, is the oidy name based on
the pallid race of Myotis californicus inhabiting the plains of South
Dakota, Kansas, and Texas. The type was taken at Banner, Kansas.

Cinereus (Vespertilio). Beauvois, Catalogue Baisonne du Museum de
Mr. C. W. Peale. Philadelphie, p. 18, 1796. VespcrtU'to cinereus
Beauvois (originally misspelled lincreus) is the first name based on the
hoary bat, Lasiurus cinereus. The description is so detailed and accu-
rate as to leave no doubt as to the animal that Beauvois had in mind.^
The type came from Pennsylvania, somewhere near Philadelphia, where
the species undoubtedly occurs during migrations.

Crassus (Vespertilio). F. Cuvier, Nouv. Ann. Mus. d' Hist, Nat., Paris,
I, p. 18, 1832. I can not identify F. Cuvier's Vespertilio crassus. The

tiroes de la teinte differonte dii pel.age, m'ont paru dtablir avec assez de certitude la
non-ideiitito d'fspf'cc de cc vespertilion avec le 7n>triniis ; c'est ce qii'indiqnent en
outre les pioiioitioiisdii crane. Le chanfrein est ])liis court et plus largo dans Ic ves-
pertilion de la Caroline. Eu voici les dimensions: longueur du corps, (jl millimetres;
lie la (juene, 28; de I'envcrgure, 259.

"Cette espece n'a point encore 6t6 ddcrite: elle m'a etc remise par M. Bosc, (|ui se
Test procurt^e lors de son scjour i\ la Caroline. Ce savant naturaliste a bieu vonlu
ui'informer qu'elle y est excessivement coumiune. On la reconnoitra anx caracteres
suivans: Orcilles ohh>n[/iics, de hi loufjiienr de la trie, reliics en part'ie; oreiUon tn dcml
cu'itr. Pelade d'un bniu niarron en drssiis, jaiindtre oi dcsnoiis."

'In the Harvard College library, Cambridge, Mass., and in the Smithsonian library,
Washington, D. C.

•17. Cliauve-souris grise. Deux premieres <leuts superienres fort petites A: pen
apparentes. Tete blanehatre; oreilles roudes, plates, blanches, le pouitour noir, une
appendice a la base. Polls du corps gris, vers la base ; noirs vers la pointe &, hlaucs
a I'cxtremitt?; de sorte que ranimal a Fair d'etre mouchete, de blanc. Cos polls
s'etendentjusqne sur la membrane qui enveloppe la queue. La membrane ailifonno
est »?galement velue en dessous a la partie anterieure, aiusi qu'au dessus a la base de
I'ongle saillant. Cettc membrane est environ une fois plus graude que dans I'espece
precedente [I'espertillo fiisrus'}. FA\e a de donze a quatorze ])ouces d'envergeure.
Le.s nariues scut emargiuees.

(irey Bat. FespertUio linereus [sic].

Elle ne .se trouve point dccrite diins les auteure. Cette chauve-Souris se trouve
dans la Pensilvanie.

24 NORTH AMERICAN FAUNA.

animal maybe Ni/cHceius humrrali.s^ but tbere is iiotbing in tbe original
description' to indic;ate tliis witb certainty. Fortunately tbe name is
not needed as all tbe species now known to inbabit tbe eastern United
States were already named at tbe time wben it was publisbed.

Creeks (Vespertilio). F. Cuvier, Nouv. Ann. Mus. d'Hist. Nat., Pari-,
I, p. 18, 1832. Vcsjjertilio creels F. Cuvier is anotber unidentifiable
species. Le Conte, bowever, wbo sent tbe type specimen to Cuvier,
states tbat tbe animal is tbe same as Ni/eticea crrpusculari-s Le Conte
{^=^N. Jill mend is llatinesque). IS^otbing in tbe original description- con-
tradicts tbis assertion.

Crepuscularis (Nycticea). Le Conte, MoMurtrie's Cuvier, Animal
Kingdom, I, p. 131, 1831. Tbis bat is tlie Xycticeins humeraUs of liatiu-
esque.

Cubanus (Vesperus). Gundlacb, Monatsber. K. Preuss. Akad. Wiss.,
Berlin, p. 150, 186L The descrii)tion of tbis species indicates a
Nycticeius closely related to N. humeralis. As I bave seen no Cuban
specimens, I am unable to say wbetber tbe animal is specifically distinct
from tbe mainland form (see p. 121).

Cubensis (Scotophilus). Gray, Ann. Nat. Hist., IV, p. 7, 1839. >>coio-
'pliihtH cuhcnsl,s Gray is evidently tbe Cuban VtHpertUio. Tbe original
description is as follows:

Fur blackish brown (in spirits); win>j:8 dark, blackisli; nnrtor.side of the inter-
fenioral membrane whitish, with scattered hairs; feet hirge; heel bono short, taper-
ing; ears moderate, entire; tragus ovate-lauceolate. Body and head 2|; tail V\; fore
arm If. Hab. Cuba.

Tbis is the first name based on tbe animal to wbicb it refers.

Cyanopterus (Vespertilio). Kafines(jue, American Montbly Mag., Ill,
J). 415, 1818. lvafines(iue's Vespertilio eijanopierns can not be identified
witb any known bat. Tbe original description is as follows:

Tail one-third, 2 iucisores above, fi beneath, body dark gray above, bluish gray
beneath, wings of a dark bluish gray, shafts black, ears auriculated, longer than
the head. Length 3 inches, breadth 10.

'Ala tcte des Murinoides, deux fausses molaires auomales de chaque cote des deux
machoires; I'oreille obtuse el Toreillon en couteau.

Tontes les jtarties supi'^rieures du corps sont d'nn brun-marron grisatro, et les ]iar-
ties inforieures blondes ; les polls, a leur origine, sont plus fonces qu'a h-ur extr(5mit.\

Des moTistaches garnissent les cotes de la Itivre supdrieure et rextr(5mit6 de la
machoire infcricure.

Longueur du corps, du bout du muse.au a Torigino de la queue, 2 pouces; de la
queue, 1 ponce 8 lignes; euvergure, S pouces S lignes.

Cette espece est due a M. Lesueur, qui I'a envoyce de New-York, sous le uom (pie
je lui ai conserve.

'5» Le V. Creeks, V. Creels.

A la tete du Serotinoides, point de fausses molaires auomales a la machoire supe-
rienre, et une seule arinferieure; I'orielle est echancree, et I'oreillon en couteau; les
parties supdrieures sont d'un bruu Jauuatre, les parties inferieures d'un gris sale, les
polls de toutes ces parties sout noirs a leur base. Des moustaches garnissent les cotds
du museau et le dessous de I'extremite di; la mAchorio inferieure.

Longuer du corps, du bout du museau a I'origiue de la queue, 2 pouces; de la
queue, 1 pouce 6 lignes; euvergure, 9 pouces.

De G^orgie. Dfi aiix recherches de M. li' major Ijcconte.

NOMENCLATURE. 25

Cynocephalus (Nycticea). Le Coute, McMnrtrie's Cuvier, Animal
Kingdom, I, p. 432, 1831. Tliis is a free-tailed bat, the common Nyc-
tinomus of the southeastern United States.

Domesticus (Vespertilio). Green, Donghty's Cabinet of Natural His-
tory, II, p. 290, 1832. The description refers without much doubt to
Myotis lucifugus Le Conte, named only one year previously. Type
locality a village in western Pennsylvania near a stream which enters
the Ohio a few miles from Pittsburg.

Dutertreus (Vespertilio). Gervais, in Eamon de la Sagra's Tlist. de Tile
de Cuba, Mamm., p. G; Atlas, Tome II, 1840. This is VeapertUio fus-
cus cuhcnsis (Gray), as shown by the number of teeth, 32, and by the
size, forearm 47 mm,

Erythrodactylus (Vespertilio). Temrainck, Monographies de Mamm.,
IT, p. 238 (13me Monogr.), 1835-41. Temminck describes his Vefsper-
tilio erythrodactylus as a bat with short, roundish ears, long tail, inter-
femoral membrane hairy on basal half above, four upper incisors, and
general reddish-brown color.'

This is a combination of characters normally possessed by no known
North American bat. The type is said to have come from the neighbor-
hood of Philadelphia. It is probably Pipistrellus suhfJarus reddened
by alcohol (see p. 8).

Evotis (Vespertilio). H. Allen, Monogr. North Am. Bats, p. 48, 1864.
This is the first name for the large eared Mi/otis of the western United
States.

Exilis (Vespertilio). 11. Allen, Proc. Acad. Nat. Sci. Phila., p. 283,
1866. Vespertilio exilis is a synonym of Myotis californicus. The type
came from Cape St. Lucas.

Frantzii (Atalapha). Peters, Monatsber. K. Preuss. Akad. Wiss.,
Berlin (1870), p. 008, 1871, Peters's Atalapha frantzii from Costa
Rica is the small, scantily furred southern race of Lasiurus horealis.
It had previously been described as Atalapha mexicana by Saussure.

'Taille nioiiulre que la pipisireXle. Tout rantibracliiuiii, la base des doigts ot la
inciiibrane iutcrtligitale da premier doigt ron<;eatre; les autros membranes noires.
Oieilles poiliies depiiis la base jns(iu'a plus de moitie de la loujiueur, petites ovoides ;
tragus cu I'euille do saule; queue tres lougue a graud bout libre; membrane iuter-
ff'inorale en dessus moitio poilue; par dcssous, rayoe de ^eines en losange, d'ou nais-
sent des soies tr.-s oonrtes disposoos a claire-voie. Dents ineisives 4 par paire en
liaut et 6 en bas; uiolaires 5 partout, vsenleuRut line fausse molaire a la macLoire
supi'rieurc.

I't'lage ]f)ng, fin et soyeux; en dessus tricolorc, an dessous bicolore. Toutes les
parties Buporieures d'une teinte brune-rougeatre; mais un pen jaunatre a la tote et
an ecu ; les poils dtaut noirs a la base, puis JauuAtre et le bout brnn-rougeatre ; moitie
do I'iuterfonuirale tr^s poilue; en dessous bruu foucc a la base et brun-roussatre an
bout; membranes des tlancs et iuterfemorale couvertes de poils rares.

liOngueiir totale 2 ponces 10 lignes ou 3 ponces pour maximum, dout la queque
piend 1 ponce 4 lignes; antibrachinni 1 ponce 2 ligues : euvergurc 7 ponces G lignes
on 8 ponces an maximum. * * ■

I'atrie. L'Amerique septentriouale dans les environs de Philadelphie.

2G NORTH AMERICAN FAUNA.

Funebris (Lasiurus), ritziiiger, Sitzmigsber. K. Akad. Wiss., Wien,
Iste Abth., LXII, p. 40, 1870. Lasiurus funebris Fitziiiger, based on
the XycticeJKs norehoracensis of Temuiinck,' from Tennessee and ]\Iis-
souri, is a synonym of Lasiurus horealis (Miiller), as shown by the
reference to the reddish-brown color and wliite shoulder spot.

Fuscata (Atalapha). Eafinesqne, Annals of Nature, p. 2, 1820. IJafi-
nesque's Atalapha fuscata can not be identilied. The original descri[)-
tion is as follows:

Ears loiigers than the head, anriculated and blackish; tail three-sevenths of total
length, jutting only hy an obtuse point; body bro^Ynish above, grayish beneath
shoulders and cheeks dark brown ; hind feet blackish, hairy above; "wings blackish
brown. — Found in the northern j^arts of the state of New York and in Vermont.
Total length three and an half inches. My genus Atalapha (Prec. dec.) contain all
the Bats without fore teeth ; there are 3 or 4 species of them in tiie United States all
blended under the name of VespcrtiJio (or KnctiUo) novehoracensis by the writers.

Fuscus (Vespertilio). Beauvois, Catalogue Eaisonne du Museum de
Mr. C W. Peale. Philadelphie, p. 18, 1796. Vespertilio fuscus 13eanvois
is the first name based on the common brown bat of the eastern United
States.2 The original description is faulty, as it contains a glaring
error with respect to the number of upper incisors, which are said to ho
only two. Nevertheless there can be no doubt as to the animal that
Beauvois intended to describe, since only one brown bat of the size of
Mijotis myotis ('la chauve-souris ordinaire de France') inhabits the
region about Philadelphia.

Georgianus (Vespertilio). F. Cuvier, Nouv. Ann. Mus. d'Hist. Nat.,
Paris, I, p. 1(5, 1832. The specific name //eor///rf^r?;.v long- passed current
for the small ripistrellus inhabiting the^astern United States. In 1893
H. Allen substituted for it the older name carolinensis Geoft'roy. As
already shown, however, there can be no doubt that Geoft'roy's animal
was Vespertilio fuscus. It is equally certain that Cuvier's name can
not be applied to the Georgian bat, since his description probably
refers to a Myotis, while in the same paper Cuvier accurately describes
the Georgian bat as Vrspcrtilio suhfarus. Le Conte, wlio collected the
specimens on which several of Cuvier's siiecies were based, describes
the Georgian bat under the name georgianus," find expressly states that

'Monographles de Manimalogie, II (13me Monogr.), p. 1.58.

-16. Chauve-souris brune. Deux preiuiires dents snperieures, distantos Tunc de
I'autre, & voisines des canines, une fois plus courtes que ces dernii'res: orcilles nues,
noiratres, ov.ales, avec un appondice a leur base; queue prcs(|u'aussi longuequole
corps (la tt'te excepte) memltrune ailiforme noiratre: polls du corps bruns en dessus,
grisatres en dessotis.

Brown bat. Vesprrillio fuscus.

Cette Chauve-souris est la plus commune que Ton tronvcdans les enviroAS deriiil-
adelphie. Ella ressemblc bcaucoup a la chauve-souri.s ordinaire de France, niaiti cu
difftro cssentiellement par )c nombre des dents de la. m.achoire supc^rieure.

^Proc.Acad. Nat. Sci. Thila.. VII (1851-5.-)), p. 131, 185(1.

NOMENCLATURE. 27

this was tlie animal that tlic riciich autlior had in hand. The evi-
dence is so strongly against this view that Le Conte's statement may be
safely disregarded.'

Greenii (ScotopMlus). Gray, List Spec. Mamm. Brit. Mus., p. 30, 1843.
Gray's ScotopMlus greenii is a nomen nudum which refers without much
doubt, however, to Vespertilio fuscus. The name is introduced as fol-
lows: "Green's Bat. Scotophilus Greenii. a In spirits. — N^ortli
America. Presented by Jacob Green, M.D."

Gryphus (Vespertilio). F. Cnvier, Nouv. Ann. Mus. d'Hist. Nat., T, p.
15, 1832. Dr. Harrison Allen has recently used the name '■yesperiiUo''
(jryplius for the ' T.' lucifuyus and 'F.' ftuhuJatus of his first monograph
which he unites as subspecies.'^ The combination of characters; two
premolars in each jaw, light yellow color, aiul hairy lips,^ is not known
in any bat inhabiting the eastern United States. Hence the description
is wholly undetenninable. Le Conte refers the name to Vespertilio fus-
cus,^ but this determination is very doubtfnl.

Henshawii (Vespertilio nitidus). U. Allen, INFonogr. Bats N. Am., p. 103,
1893. VcHpertiJio nitidus hcnshan-ii 11. Allen is a synonym of Myofis
eaUfornicus, based on pale examples of the latter from near Wingate,
N. Mex.

Hesperus (ScotopMlus). H. Allen, INIonogr. N. Am. Bats, ]>. 43, 1864.
This is the first name based on the common Pipistrelliis of the south,
western United States.

Humeralis (Vespertilio). Bafinesque, Americau Monthly Mag., Ill,
p 44."), 1818. While there is nothing absolutely diagnostic in the original

'The ()ri.i:;in.al description of Vespertilio georgianus is as follows:

"Ala t(*te (lea MuriiioTdes; I'oreille est dchancrt^e et Foreillon en ali'iio. Toiitcs
les parties siiporieures du corps sont colorccs par nn melange do iioir et de blond
jauni'itre. Le noir i)aroit, parccqiie la pointe despoils qui est blonde ne recouvre
pas, a cause de sa brevitc, lo reste do la longueur dc ces polls qui est noir. Les par-
ties iuferieures sont grises, niais molauge'es de noir, par la nu'ino cause qui fait
paro'itre cette couleur aux parties supi'rieures. Des moustaches garuissent les cAtrs
des li'vres superieures, ot le dessous de rextremitc do la iiiachoire inferieure.

"Longueur du cori»s, du bout du uiuseau a I'origiue de la queue, 1 pouce G lignes;
dc la queue, 1 pouce 2 lignes; envergure, 7 ponces.

"De Goorgie. Du aux rccherches de M. le major Leconte."

"Monogr. Bats N. Am., j). 75, 1893.

"•The description is as follows:

"A latete des Muriuoidoset deux fansses niolaires anomales fort petites de chaque
cot*^. des deux machoires; I'oreille est dchancrce et roreillou en couteau. Toutes les
parties snp(5rieures du corps sout d'un blond jaunatrc, les parties infcrieures sont
grises, mais les polls des nns et des autres sont noirs a lour extr(^nute inferieure. Les
parties uues sont viohUres. Des moustaches garuissent les cotes de la levre supe-
rieuro et le dessous de rextromitc de la machoiro inferieure. Longueur du corps, de
rextremite du museau a rorigino de la (lueue, 1 pouce 9 lignes; do la queue, 1 jjouce
2 lignes; envergure, 7 ponces 10 lignes.

"Des environs de New York. Du aux vecherches de M. Millifrl-."

M'roc. Acad. Nat. 8ci. Phila. Vll (1854-55), p. 134, 1856.

28 NORTH AMERICAN FAUNA.

description' of this species, its subsequent treatment is such as to leave
no reasonable doubt that Raflnesque liad in mind the bat afterward
named Ni/ctieea crcpuscularlH by Le Conte. In 1819 Raflnesque based
the genus Nycticeius on two of liis species of Vespertilio which differed
from all others known to him in the possession of only two incisors
in the upper jaw. One of these, V. tesselatns, was the red bat, Lasiurus
borealis. The other, V. humeralis, must have been the twilight bat, as
there is nothing" in tlie description that precludes it, and no other small
si)ecies with two ui)per incisors is known in the eastern United States.

Incautus (Vespertilio). J. A.Allen, Bull. Am. Mus. Nat. Hist., VIII,
]). 239, November 21, 1896. Vesperiilio iiieautus J. A. Allen, is a syno-
nym of Myotis velifer (J. A. Allen), based on specimens of the latter
from San Antonio, Tex. (See p. 59.)

Intermedius (Lasiurus). H. Allen, Proc. Acad. Nat. Sci. Phila. (18G2),
p. 14:(), 1863. This is the only specific name based on the bat now
known ns JJasyptcrus intermedius.

Keenii (Vespertilio suhulatus). Merriam, American Naturalist, XXIX,
p. 800, September 1, 1894. W'sjK-rtilio .siibidafiis keenii is the only name
based on the dark form of Myotis snhnlatus occurring on the Queen
Charlotte Islands, Uritish Columbia.

Lanceolatus (Vespertilio). ^^aximilian, Reise in das Innere Nord-
America, I, p. 364, footnote, 1839. The specific name lanceolatus was
proposed Dy Maximilian as a substitute \'ov sn hn lain s, shon\d the animal
which he designated by the latter name i)iove to be different from
Say's.'^ Maximilian's siilmlatus is described at considerable length and
is probably the VespertiUo Ineifnr/ns of Le Conte. The following meas-
urements are given: Total length. 3" V"; extent, 8" 9'"; tail, 1" 3'";
ear, 0'"; tragus, 2.^".

Lasiurus (Vespertilio). Schreber, Siiugthiere, Abth. I, PI. LXII B,
published with Abth. IV, I left 34, 1781.'' The figure of Vespertilio
lasiurus is a good representation of the red bat (Lasiurus horenlis Mid-
ler, 1776). Dobson^ cites this name as dating from 1775, in which case
it would be the earliest for the species. This is, however, a mistake. PI.
LXII appeared with Abth. I in 1774, but PI. LXII B, was not pub-
lished until 1781 with Abth. IV, Heft 34. The species is mentioned in
Abth. I (p. 170) as 'Die nordamerikanische Fledermaus.'

Lasurus (Vespertilio). Boddaert, Elenchus Animalium I, p. 71, 1785.

'Tail tluee-seM!ntli8, upper incisores 2, remote, lower 6, body dark brown above,
slioulders black, ,i;ray beiieatb, wiugs, tail, ears, and snout blackisb, eyes under tlio
bair, ears longer tban tbe bead, elliptical, anriculatod. Lengtb 3 1-2 inches,
breadth 11.

-Diese Fledermaus bescbrieb icb in nieinem Tagebucbe nnter der Benennnng leap.
Janceolaina, sie bat abcr viel Aebnlicbkeit mit Say's V. fiiihiihitns. Zu Betblelieni in
Peuusylvanien erbielt icb zwei Excniplare * * * _

^For date of publication see Sberborn, Proc. Zool. 8oc. London, 1891, p. 589.

■•Catal. Chiroptera Brit. Mus., p. 269, 1878.

NOMENCLATURE. 29

Vcsju'iiilio hisnrus Boddaertis probably a misprint for V. hi,sinri(.s, siuco
reference is made to tSclireber's plate. ^

Lecontii (Plecotus). Cooper, Ann. Jjyceiim Xat. Hist. New York, IV,
}), 72, 1848. Concerning" Plecotus lecontii, Cooper says:

The uauie inacroiis 1 have venturetl to supersede, as being in nowise distinctive of
the species, but in reality derived from a generic cliaracter, which in some species
is more developed than iu tlie present. Tiie ears Ijeing therefore rather miudl for the
genus, this name becomes contradictory ; and no American naturalist will regret the
op])ortunity thus afforded of paying a well merited tribute to the discoverer of so
many rare and remarkable animals of this country.

The name is of course a synonym of macrotis Le Conte.

Leibii (Vespertilio). And. .S: Bach.. Jonrn. Acad. Nat. Sci. Pliila.,
VIII, rt. II, p. 281, 1842. Vespertilio leibii And. & Bach., from Erie
County, Mich, [now Ohio] is probably Myotis lucifuffus Le Conte. The
measurements are as follows: " Length of head and body 1 inch 7 lines;
tail 1 inch 4 lines; spread 7 inches; height of ear posteriorly 2.] lines;
tragus I line."

Longicrus (Vespertilio). True, Science, VIII, oSTo. 20.3, p. ."iS8, Dec. 24,
1886. Vespertilio loiujicrus True, is the only name based on the com-
mon western subspecies of Myotis sithulatKs.

Lucifugus (Vespertilio). Le Conte, MclMurtrie's Cuvier, Animal King-
dom, I, !>. 431, 1831. The original description of Vespertilio lueifwjus
Le Conte is as follows:

Anterior upper fore-teeth bilobate; body above dark brown, beneath cinereous;
nose sub-bilobate; face with a nakedish prominence on each side; ears oblong,
naked, tragus sub-linear, half as long as llie ears; tail projecting a little beyond the
membrane; length to the insertion of the tail two inches and a quarter; tail one
inch and a quarter.

From this alone it would be impossible to identify the animal that
the writer had in mind. Fortunately, Le Conte treated the species in
more detail in a paper published in the Proceedings of the Academy
of Natural Sciences of Philadelphia for 1855 (pp. 431-138). Here he
recognizes three species of ' VespertiHo'' with thirty eight teeth as occur-
irng in the eastern United States. These are V.subulatus, V.lncifuijus,
and Y. (/corf/ianiis. V. (jcorgianHs is clearly Pipisirellns suhflarKs, which
Le Conte placed with the thirty-eiglit-toothed species through an error
in counting the teeth. P. lucifugus and V. snbulatHS of Le Conte are
evidently based on individual variations in the shorter-eared of the two
eastern species of 4///o/ /.v. The only differences in Le Confers descrip-
tions of the two forms are the following: P. subulatns: Ear slightly
emarginate; length 2.0; tail 1.1; extent 9.4; head .9; ears .4; oriUon
.3. V. IncifiKjus: Ears so much emarginated as to appear hooked;
length 3.8; tail l.C; extent 11.7; head .75; ears .45; orillou .2.

' Boddaert's account is as follows:

" Lasurus. 16. V. cauda longissima, rostro oblicjuo truncato, hi Joufjne (Jiieite.
Schreb., tab. 52. B lomjtaUed Bat."

Habitat: "Quare Doct. Krxlebeu, Zimmermauu, Pennant hune notabilem vesper-
tilionem omiserunt, mihi latet."

30 NORTH AMERICAN FAUNA.

Macleayii (Scotophilus). (iray, List Spec. Maiiim. Brit. Mus., p. 80,
1843. i^c'otopliilii.s iii<iclc(ii/ii Gray is a iioineii iiuduiu, probably based
on YesperiUio i'uHvns cubeusis. Gray says merely: "MacLeay's Bat.
SooTorHiLUS MacLeayii a In spirits. Male. Cuba. — Presented by
W. S. MacLeay, Es(i.''

Macropus (Vespertilio). H. Allen, Proc. Acad. Nat. Sci. Pliila,, p. 288.
1866. Vespertilio macropiis H. Allen is a synonym of Myotis yumaneum
(H. Allen). Tbe name is, moreover, preoccupied by Vespertilio macropus
Gould, 1854.'

Macrotis (Plecotus). Le Coute, McMurtrie's Cuvier, Animal King-
dom, 1, !>. 431, 1831. Plecotus macrotis Le Oonte is the first name cer-
tainly applied to tlie bat now known as Corynorhinus macrotis.
llafinesque's Vesiurtilio megalotis may have been the same animal, but
his description is so poor that it is impossible to determine what he
refers to.

Maculatus (Histiotus). .1. A. Allen, Bull. Am. Mus. TsTat. Hist., New
York, 111, }). 1 '.»."), 1891. Histiotus maculatus is the name under which
the bat now known as Eiulerma m<{culatiim Mas first described.

Megalotis (Vespertilio). Patines(|ue, American ^Monthly INIag,"., Ill, p.
•446, 1818. There is nothing- in the original description-' of Bafines(iue's
Vespertilio megalotis by which the si)ecies can be identified. It is pos-
sibly tlie animal afterwards named Plecotus macrotis by Le Conte.

Melanops (Eptesicus). Kafinesque, Annals of Nature, p. 3, 1820.
When Kafinesque transferred his Vespertilio phaiops to the geiuis
Eptesicus^ he clianged the specific name to melanops, thus adding
another to the synonyms of Vespertilio fuscus.

Melanorhinus (Vespertilio). Merriam, North American Fauna, No. 3,
p. 40, September 11, 1890. Vespertilio melanorhinus Merriam is a syn-
onym of Myotis calij'ornicus, based ou a specimen of the latter from San
Francisco Mountain, Arizoiui.

Melanotus (Vespertilio). liafiuesque, American Monthly Mag., Ill, p.
445, 1818. Ivafines(iue's Vespertilio melanotus is hopelessly indetermi-
nable. The original description is:

Tail oue-tliird, brown iibove, gray Ijeueath, body bhuki.sh aljove, whitish beueath,
wing.s dark gray, shafts black, ears aiiriculated, rouuded. Length 4 1-2 inches,
breadth 12 1-2.

Melas (Eptesicus). Le Conte, Proc. Acad. Nat. Sci. Phila.,yil (1854-55),
p. 438, 1856. In a paper on the bats of the United States published in
185(5, Le Conte refers to Eptesic^is melas Rafinesque as an unidentified
species. I have been able to tind no such name in any of llafinesque's
writings and therefore suppose that Eptesicus melas is a misprint for
E. my das, especially as the latter is not mentioned by Le Conte.

'Mammals of Australia, III (fide Dobson).

-Tail three-eighths of total length, body dark gray above, pale gray beneath, ears
very large, duplicated, auricules nearly as long. Length 4 inches, breadth 12 inches.

NOMENCLATURE. 31

Merriami (Vesperugo). Dobson, Ann. & Mag. Nat. ITist., XVIII, p.
124, 1886. Vcspvruyo merriami Dobson, Avas based on a speciiueu of
ripiNtrelhi.s hesperus from I{ed IJhiff, Tehama County, Cal., wrongly
supposed to have been taken at Locust Grove, N. Y.

Mexicana (Atalapha). Saussure, Kevuc et Mag. de Zool., 2e ser., XIII,
J). 1)7, 1861. Atalaplia mexicana Siuissure is the first name based on
the southern race of Lasii(rns horvalis, afterwards descril)ed by Peters
as Aialapha franizH.

Mexicanus (Vespertilio). Saussure, Revue et Mag. de Zool., 2e ser.,
W\. ]). 282, July, 1860. Under the name VcKpertiUo mcxieanns Saus-
sure described the large, dark Mexican form of Myoiis californicns,
which had hitherto received no name.

Miradorensis (Scotophilus). II. Allen, Proc. Acad. Xat. Sci. Phila., p.
287, 1866. ^cotopJtilus miradorenHiH B. Allen is the only name based
ou the large southern form of Vespertilio fu.scKs.

Monachus (Vespertilio). liafinesque, American Monthly Mag., Ill, p.
445, 1818. Theorigiinil description of Ilalinesque's Vcspcrt i I io monachus
leaves no doubt that it refers to Lasinrus borealis (Miiller). It is as
follows:

Tiiil oue-fouitb, liairy Jiljove, friuged hiterally, body jialc. fallow above and Itelow,
head and ueck covei-ed with a longer fur of a dark red fallow, ^viugs dark gray,
shafts red, hiud feet black, uose red, ears concealed in the fur. Length 4 inches,
breadth 12.

Monticola (Vespertilio). And. »S: Bach., Jouru. Acad. Xat. Sci. Phila.,
I, Xo. 7, p. 92, October, 1841. Vespertilio monticola is probably Pipis-
trellus snbfiavns (F. Cuvier), though the description is not wholly per-
tinent to this species. The original account is as follows :

Vespa-tUio monticola (Mountain bat). — V. vespertilione subulata brevior; uuriculns
breviori))UB ; tragus nonexcedentibus, diniidiani longitudinem auiicula' ; colore fulvo.

Mountain Hat. — Smaller than Say's bat ( T. snhnlatns); cars shorter; tragus, less
than half the length of the ear; color, yellowish brown. Upper fore teeth bilobate,
cars moderate, nakeil, erect, rather broad at base; tragus linear, subulate, body
small; wings long; tail i)rojecting a line beyond the interfcmoral membrane, which
is slightly sprinkled with hair above and beneath.

Color. — The nose and chin are black; cars light brown; Aving membranes dark-
brown. The whole of the fur of the body, above and beneath, is from the roots, of
a uniform yellowish-brown color.

The species differs from Say's bat not only in color, but in the much shorter ears and
tragus. The size and sha]ie of the tragus wo have found an invaluable guide in our
American bats; the ears of the j)resent S](ecies, when alive, are always erect; while
those of Say's Bat arc folded backward like those of the long-eared Bats — I'lv-
coliis. * * »

IHmcnsiovs. — Length of head and body, 1 inch 8 lines; length of tail, 1 inch 6
lines; length of spread, 8 inches; height of ear posteriorly,;} lines; height of
tragus, 1:^ lines.

N. B. — The tragus in Say's Bat is four-and-a-half lines in height. Several sjieci-
mens of this Bat were obtained during the summer, on the mountains of \'irginia, at
the Grey Sulphur Springs. They were uniform in size and color.

32 NORTH AMERICAN FAUNA. |

Mydas (Eptesicus). Eafiiiesquo, Annals of Nature, p. '^, 1820. The
description of I'Jpfc.sicns nii/iht.s leaves the species hopelessly indeter-
minable. It is as follows:

Fulvous above, grey beueath ; wings, ears and tail, pale brown, shafts whitissb;
ears double the length of the head; tail naked, slightly inucronate, nearly as long
as the body. — I have observed it in the barrens of Kentucky dying in the houses.
Total length three inches, of which the tail includes live-twelfths. Ears three-
(luartera of an inch long. I mentioned it under the name of J'esp. miidas in my
account of the Bats of the western states, (Am. Mag. v. 3). I have since iustitutcd
two other general with them, Hi/pexodoii and Xycliceiiis (Prodr. 70 N. G. An); the
others are probably Atalaphes. I know already lifteen species of Bats in the
ITnited States, almost iill new ones.

No bat is known to occur in Kentucky that combines the characters
attributed to this animal.

Mystax (Vespertilio). Uafinesiiue, American Monthly Mag., Ill, p.
445, 1818. This species which Kafinesque had already referred to as
Noctilio Dii/stax,^ is described as follows:

Tail two-fifths of total length, upper incisores none, lower 6, 2 warts at the
lower .jaw, body entirely fallow, to]> of the head browniish, oars brown, auricu-
lated, longer than the head. Length 5 inches, breadth 14.

In the diagnosis of the geiuis i/y^>f\rof/o?j, based on this species, some
further characters — such as 'nostrils round, projecting,' and 'lips
whiskered' — are added, which only serve to increase the impossibility
of identifying the aninuil.

Nigricans (Vespertilio). Maximilian, BeitrJige Naturgesch. Brasil., II,
p. 26(>, 1826. Mi/olis ui(/ric<(ns (Maximilian) is a species closely related
to M.ca I {/'(>)•)( icus,\yhi{!,]\ it re[)laces in the tropical fauna from southern
Mexico southward. The name was applied to M. californicns by Dr.
Harrison Allen in his recent monograph (18!>.'>). In the original desciip-
tion Maximilian cites Scliinz ('Thierreich u. s. w. 1>. I. p. 179') as
authority for the name. As I have been unable to verify this reference
I do not know wliether the name was actually ])ublished before 1826.

Nitidus (Vespertilio). H. Allen, Proc. Acad. Nat. Sci. Phila. (1862),
p. 247, 1863. Vespertilio nitidus H. Allen, is the common small brown
bat of the western United States and therefore the name is a synonym
of V. caUfornicus Aud. & Bach., 1842.

Noctivagans (Vespertilio.) Le Conte, McMurtrie's Cuvier, Animal King-
dom, I, p. 431, 1831. This is the first name based on the silver-haired
bat, Lasionycteris noctivagann.

Noveboracensis (Vespertilio). Erxlebcn, tSyst. llegni Anim., I, p. 155,
1777. Erxleben's Vespertilio norehoraceiisis was based on the New
York bat of Pennant (Sym>i). (»)uadr., p. 367), 'Die nordamerikanische
Fledermaus' of Schreber (Siingthiere, I, p. 176), aud 'Der Neujorker' of
Midler (Natursyst. Suppl., p. 20). It is therefore the red bat, Lasturus
borealis. '

Noveboracus (Vespertilio). Boddaert, Elenchus Animalium, I, p. 71,
1785. This is the red bat, TAisinrus iiorealis Miiller. Boddaert meu-

1 American Monthly Mag., Ill, p. 354.

NOiMKNCLATURE. 33

tioiis the white shouhler marks characteii.stic of the species and refers
to Schreber and Pennant.

Obscurus (Vespertilio). H. Allen, Proc. Acad. Xat. Sci. Phila., p. 281,
1866. VespertHio obsourns VI. Alien, is one of the numerous synonyms
of Mijotis califoraicus. The type specimens came from Lower Oaliforuia.

Oregonensis (Vespertilio). II. Allen, Mongr. Bats jST. Am., p. 01, 1864.
The wording of Dr. Allen's account of Vespertilio oreyonensis is so
ambiguous as to leave some doubt as to whether he intended to apply
the uame to specimens from Fort Yuma and Cape St. Lucas (Nos. r)105,
5537, and 5402) or to a skin labeled oref/oitensis by Le Conte. In either
case the name is a synonym of V. californicus Aud. & Bach. Under
V. nitidus he says :

Nos. 5105,5537, and 5102, four specimens in ull, present the following;- peculiarities:
The fur is longer tliau in others of tiio collection. On tlie back the base of the hair
is blackish; upper tliird pale yellow, tuining to a delicate light-yellowish russet
brown ; on the belly the hair is dark brown at the base, with light tips; the hairs on
the interfemoral uienilirane are also of a light color. In other n'spects the charac-
ters arc the same as the other specimens. The diied specimen. No. .5512, labeled by
Dr. I.e Conte J\ ovegonensis, though never described by him, probably belongs to this
variety. If the individuals having the above coloration should be found to consti-
tnte a new species, this name will be reserved for it.

Pallidus (Vespertilio). Le Conte, Proc. Acad. ISIat. Sci. Phila., VII,
(1854-55) p. 4.')7, 1856. Vespertilio pallidus Le Conte is the only name
based on the Eastern form of Antrozous, the- type of the genus. Le
Conte stated that his species came from California, but this is evidently
an error, as pointed out by Baird and Harrison Allen. The type, now
in the United States National Museum, is labeled 'Fort Clark, Texas.'
It agrees in all respects with skins taken in the same region by Dr. E.
A. Mearus.

Parvula (Rhogeessa). II. Allen, Proc. Acad. Nat. Sci. Phila., p. 285,
1866. I\}io(/ei'ssa parrula H. Allen, from the Tres Marias Islands, IVIexico,
is [)robab]y distin(;t from any of the members of the genus that occur
on the mainland. The type is now mislaid or lost.

PfeifFeri (Atalapha). (xundlach, .^lonatsber. K. Preuss. Akad. VViss.,
Berlin, p. 152, 1861. Gundlach's Atalapha p/eifl'eri is the only name
based on the Cuban form of Lasiuru.s borealis.

Phaiops (Vespertilio). Ilaflnesque, American Monthly Mag., Ill, p. 445,
.1818. Under the name Vespertilio phaiops^ Bafinesque gave an accurate
description of Vespertilio fuscus Beau vols. He says:

Tail one-third of total length, naked, mucronate, Ijody dusky bay above, pale
beneath, face, ears aud wings blackish, 4 incisores in the u])per Jaw, 2 on each side,
divided by a large Hat wart, unequal, the outside oiu!S larger aud bilobed, G small
incisores at the lower jaw. Length 41-2 inches, breadth 13.

Priscus (Nyctitherium). Marsh, American Jonrn. Sci. & Arts, od ser,,
IV, p. 128, 1872. Ni/ctitJtcrium priscus Marsh is a name based on a
fragment of a fossil lower jaw from the Eocene or lower Miocene near
Henrys Fork, Wyoming.

Propinqims (Vesperus). Peters, Monatsber. K. Preuss. Akad. Wiss.,
2772— No. 13 3

34 NORTH AMERICAN FAUNA.

Berlin, p. 2(52, 1872. Vespcrns propinquvs Peters from Santa Ysabel,
Guatemala, is the small southern form of YeHpe^'tlUo fusimn, I can find
no other uauie based on this animal.

Pruinosus (Vespertilio). Say, Long's Expedition to the llocky Moun-
tains, I., p. 107, footnote, 1823. Vespertilio pruinosus Say, is the hoary
bat, Lasiurus cineyeus (Beauv.). It was described from a speciuien
taken at l^^ngineer Cantonment, Washington County, Nebraska, .') miles
above the mouth of the Boyer River and not far from Council Bluffs,
Iowa.

Pulverulentus (Vespertilio). Temmiuck, Monogr. de Mamm., II, p. 23.5,
(13*' Monogr.), 1835-1841. Under the name VespertiUo pnJrerulentus
Temminck gives an accurate description of a specimen of Lasionycteris
noctivagans tiiken on the Missouri River.

Rafinesquii (Plecotus). Lesson, Manuel de Mammalogie, p. 06, 1827.
Plecolus rajinesqnii Lesson is a name based on Rafinesque's indetermi-
nable Vespertilio megalotis?

Rubellus (Vespertilio). Beauvois, Catalogue Raisonne du Museum de
Mr. C. W. Peale. Philadelphie, p. 18, 1796. Vespertilio rubellus Beau-
vois is the red bat, Lasiurus boreaUs (Miiller).^

Rubra (Vespertilio). Ord, in Guthrie's Geography, 2d American ed.,
II, p. 291, 1815 (Rhoads' Reprint, 1801). This is another synonym of
Lasiurus horealis (Miiller). The name a^jpears in a nominal list of
North American bats. In a footnote, however, Ord says: "Described
by Mr. Wilson. See American Ornithology, Yol. VI, p. 00." Wilson's
description, as well as his figure on plate 50 (fig. I) of the 1812 edition,
refers unquestionably to the red bat.

Rufus (Vespertilio). Warden, Description des Etats-Unis de I'Amerique
Sei^tentrionale, V, p. 000, 1820. Warden's Vespertilio rufus is another
synonym of Lasiurus boreal is based on Wilson's description and figure.

Salarii (Vespertilio). F. Cuvier, Nouv. Ann. Mus. d'Hist. Nat., Paris,
1, p. 15, 1832. Like most of the species described in the same paper,

' The origiual descriptiou is as follows :

"Pelage (run giis foine on tlessiis, ct d'mi gris pale en tlessous; oreilles tri'S gran-
tles et doubles, poiirvucs d'oreilloiis anssi loDgs qu'elles; uV-st peut-Otre (^u'uiie va-
ridte de uotre oreillard. llabite les Etats-l'iiis."

^18. Chauve-Sonris roiigeatrc Deux premiores deuts plus petites que les canines,
mais appareutes, tote ainsi quo le corps d'une couleur rougeatre uioh'o do quelqiies
polls blancliatres. Oreilles couleur de obair, nues, ropli^es et appeudiouliees a leur
base. Nariuos omarginoos et distantes I'uue de I'autre. Vespertilio rubellus. Eed-
isb bat.

Les polls du corps foruiout quelques Ibis des zones rougeatres et blanches. La
niembrauo aili forme est voluo on d ess us a la partie antorieure, ot couverte de poils
roux dossus ot autour de la queue. L'iudividu (^ue nous dcorivons est d'autant plus
curieux qu'il a etc pris avec trois petits qu'il porlo sur sou ventre. Co qu'il y a de
plus particulier, c'ost ([ue d'eux d'eutr'oux ressemblent parfaitemont a la more pour
la couleur et I'autre est tout a fait roux. La membrane alliforme est couvertes de
raie nn pen trausparentos qui vues au jour, representent des quarr<^s en forme de
Lozaugo. La couleur noiro do cotte mombrano contraste avec los oouleurs du corps
ot oolle des divisions dos pattes de devant, qui sout de couleur de chair, lorsquc I'ani-
nial est en vie.

NOMENCLATUKE. 35

Vespertilio salarii is indetermiuable, No known North Ameiicaii but
combines Iiairy lips, reddisli brown color, and two premolars in each
jaw.'

Seminola (Atalapha borealis). lihoads, Proc. Acad. Nat. Sci. Phila., p.
32, 1895. This is the dull mahogany-brown race of Lasiurus horeaJis
peculiar to the Austroripariau fauna. No other name has been based
on this animal.

Septentrionalis (Vespertilio gryphus). Trouessart, Catalogus Mamma-
lium tam Viventium quam Fossilinm, p. 131, 1897. Trouessart's Ves-
pertilio (jryplius var. septentrionalis is the only name unquestionably
based on the Myotls commonly known as Vespertilio subnlatus Say. It
is merely a latinization of 'northern form of Vespertilio gryphus,'' the
designation applied by Dr. Harrison Allen in his Monograph of 1893
to the V. suhulatus of his first monograph.

Serotinus (Nyctilestes). Marsh, Am. Jouru. Sci. & Arts, 3d ser., IV,
p. 21"), 1872.. The name Nyctilestes serotinus was applied by Marsh to
the fossil jaw of a bat found by him at Grizzly Buttes, Wyoming.

Subflavus (Vespertilio). Cuvier, Nouv. Ann. Mus. d'Hist. Nat., Paris,
I, p. 17, 1832. Vespertilio suhjiavus is one of the few North American
bats named by F. Cuvier that can be identified. It is without doubt
the Georgia bat {Fipistrellus suhjiavus), commonly known as ' Vesperugo
(jcorgianus.'' The peculiar col-oring of this species, unique among the
bats of the eastern United States, is very accurately described.'-^ This
is the first account of an American bat in which this color pattern is
referred to. The mixture of dark and light hues in Cuvier's V. georgi-
anus is due to the shortness of the fur in his specimen, which allows
the dark bases of the hairs to appear irregularly on the surface. This
is not at all the case with the small Fipistrellus of the eastern United
States. In this bat the hairs are tricolored, dark at the bases, yellowish

'The original description is as follows :

"A la tote dcs Muriuoules et deux fausses niolaires de chaque cote des deux uia-
rhoires; Toreille est I'chaiicrc'e et roreillou en coutean. Toutes les parties sup(5ri-
enres du corps sout d'un brim-inanon grisfitre, et les parties iuferieures gris-
l>l:iucliatres. Aux parties brunes les polls soiit plus Ibuces a leur luoitic iufcrieure
•|u'a leur suporieure; ils sont iioias dans cette iuferieure aux parties gris. Les par-
tiis nues sont brunes, des moustaches gainissent les cotes de la levre superieure
et le dcssous de I'extremite de la niachoire iuferieure.

"Longueur du corps, du bout du niuseau a I'origine do la queue, 1 pouce 6 lignes;
de la queue, 1 pouce 7 lignes; envergure, 7 pouces 7 lignes.

''Des environs de Ne\y York. Du aux recherches de M. Milbert."'

-The original description is as follows:

"A. la tete des Muriuoules ; I'oreille est ochancrde, et I'oreillou en demi-cu-ur. Les
parties sup(?rieures du corps sont d'un blond gris clair, legircinent ondulees de
brunatre ; les parties infdrieures d'un blanc jaunatre; les jjoils des parties supdri-
eiues sont noirs a leur base, blancbatres dans la })]us grande partie de hnir longueur,
et brunatres a leur pointe; cenx des parties inferieures sont noirs a leur ufoiti^ in-
fdrieure, et d'un blanc jauiniitre a leur autr(^ luoitid, Des moustaches garnissent les
I cotes do la li-vre superieure, et le dessons de I'extr^mitd de la machoire iufcrieure.

"Longueur du corjis, du bout du museau a I'origine de la queue, 1 pouce G lignes;
I de la queue, 1 pouce .S lignes; envergure, 7 ponces.

"De Georgie. Du aux recherches de M. le major Lecoute."

36' NORTH AMEKICAN FAUNA.

ill tlie middle, and dark at the extreme tips. Tliis is exactly what
Cuvier describes as the character of the fur of his 'Bloiidiu' {F.suhjiavm).
Subulatus ( Vesper tilio). Say, iu Long's Exped. to llocky Mts., II, p. 65
footnote, 1823. The original description of VespertiUo subulatus leaves
the species undeterminable. It is as follows:

Ears longer than broad, nearly as long as the head, hairy on the hasal half, a lUtlo
veutricose on the anterior edge, and extending near to the eye; tragus elongated,
subulate; the hair above blackish at b;ise, tip dull cinereous; the iuterferniorai
membrane hairy at base, the hairs unicolonred, ami a few also scattered over its sur-
face, and along its edge, as well as that of the brachial membrane; hair l)eneath
black, the tip yellowish-white; hind feet rather long, a few set;i' extending over the
nails; only a minute portion of the tail protrudes beyond tije membrane. Total
length, 2i'', inches. Tail, 1-, inches.

While there is nothing in this account that refers unquestionably to
the longer eared of tbe two species of Myotis inhabiting the eastern
United States, the name has jiassed current for this animal so long that,
after careful consideration of all the evidence, I am unwilling to substi-
tute for it Trouessart's nunie scptentrioxdlis, the only one une(|uivocally
based on the species. Say's Vespertilio suhuhitus came from the Arkan-
sas River, near the present town of La Junta, Colorado. The bats of this
region are not well known, but at present Mijotls crotis, M. californivus
ciliolahrum, and M. lucifuyus 1o)if/icrus are the only members of the
genus Myotls which nuiy confidently be expected to occur there. From
the known range of Myotls subulatus to the north and west, however,
its regular occurrence in Colorado is by no means impossible. Appar-
ently Le Coiite was the hrst subsecpient writer to define the name
subulatus, and, as has already been shown, his animal was an individual
variation of the shorter eared of the two eastern si)ecies. If this deter-
mination be taken as final, there can be no (juestion as to the necessity
of adopting the name septcntrioiiaUs for the longer eared animal, but at
present the power of the 'first reviser' is so much in question that too
much should not be staked on it. Harrison Allen, in 1804:, applied the
name subulatus to the longer eared of the two forms, and in this sense
it passed unchallenged until 18i>o, when the same author united the
lucifugus and subulatus of his earlier monograph under the s])e('itic
name yrypfius. This change has not been generally adopted, so that ill
retaining the specific name subulatus I am merely continuing the usage
of the past thirty-four years, not, however, Mithout grave misgivings
that the reasons for so doing are in reality unsound.

Teliotis (Atalapha). H. Allen, Proc. Amer. Philos. Soc, XXIX, p. 1,
February 11, 1891. Atalaphu teliotis II. Allen is the only name based
on the Calilbrnian form of JAisiurus borealis.

Teniiidorsalis (Vespertilio). II. xVlleii, Troc. Acad. Nat. Sci. Phila., p.
283, 1866. This is a synonym of Myotis californicus based on a speci-
men (Xo. 5533, U. S. Xat. Mus.) from Cape St. Lucas, Lower California.

Tesselatus (Vespertilio). Rafinesqne, American Monthly Mag., HI,
p. 445, 1818. Ilafinesque's Vespertilio tesselatus is Lasiurus borealis
(Milller). The original description is as follows:

Tail half of total length, hairy above, upper incisores 2, remote, lower 0, body
fallow above, head pale, dirty fulvous beneath, with a faint fallow collar, with 2

i-

NOMENX'LATURE. 37

hairy white spots above near the thumb, membraue blackish, netted of fulvous inter-
nally and clotted of same externally, shafts fulvous, nose bilobate, oars nearly con-
cealed by the hair. Leno^th 4 inches, breadth 12.

Townsendi (Plecotus). Cooper, Ann. Lyceum ISTat. Hist. New York, IV,
p. 73, 1837. Plecotus townseudi Cooper is tiie only name based on the
form of Corynorhimis inliabiting the northwestern United States.

Tumida (Rhogeessa). H. Allen, Troc. Acad. Nat. Sci. Phila., p. 28G,
1866. h'hoffel'ssa tumida H. Allen is the only name based on the small
Mexican bat to which it is now applied.

Ursinus (Vespertilio). Temminck, Monographies de Mammalogie, II
(13'' Monogr.), p. 235, 1835-41. The description of Temminck's Yesper-
tilio ursinus refers without much (juestion to Vespertilio fuscus lieauv.,
though the statement is made that there is no false molar in the upper
jaw. Color, size, and external characters, however, agree with Y. fuscus.

Velifer (Vespertilio). J. A. Allen, Bull. Am. Mus. Nat. Hist., New
York, in, p. 177, 1890. The name Yespertilio velifer has been applied
by Dr. J. A. Allen to a largo species oi' jUyotis occurring in Mexico and
the south westein United Stales. The animal is closely related to the
Vespertilio albescens of Dobson and may eventually prove to be the same
as T^. albescens (leoff'roy.

Velox (Nyctitherium). Marsh, Am. Journ. Sci. cS; Arts, 3d ser., lY,
p. 127, 1872. Nyetitherium velox is a fossil bat from the Eocene or
lower Miocene near Henry Fork, Wyoming.

Veraecrucis (Vesperugo). AVard, American Naturalist, XXY, p. 745,
August, 1891. Ycsperugo verwerueis Ward is the only name based on
a form o^ Plpistrellus occurring in southern Mexico.

Virginiauus (Vespertilio). And. & Bach., Journ. Acad. Nat. Sci. Phila.,
I, No. 7, J). 1)3, October, 1841. Yespertilio rir(/inianns can not be
identified with any degree of certainty, though it is without much
doubt one of the small species of Myotis. The original description is
as follows :

Vi'sperlUlo riyi/iiiianiis (Virginian bat). — V. vespertiliono monticola pnulnlnm lon-
gior, anririiluH itaiiluliuii lougioribus inagis<iu(' acutis; dcntibus i»rinioril)ns maxiliie
sii])eri()rissiin])licil)us; iuteri'eiiiorali meiiilirana niida ; corporesuprai'uligineo-fusco;
Hubtus cinereo-l'iiscato.

flrfiinian hat. — A little larger than the Monntain Bat; ears a little longer and more
pointed ; upper fore teeth simple ; interfemoral membrane naked ; sooty brown above,
ash brown beneatli.

r> '^-'^ 1-1

Dentition. — Incisors "^ - . Canines .
(J 1-1

In size tiiis species is intermediate between V. caroltnensis and V. sninlatns. The eai"
is naked, less rounded, and more pointed tlian eitiier of the other closely allied species.
Tho tragus is very narrow, linear, and less than half the length of the car. The tail
18 inclosed in the interfemoral membrane, except the penultimate joint, Avhich is free.
The anterior upper fore teeth, instead of being sub simple, as in the J'. caroUnensis,
or bilobate, as in V. snhnlatus and T. montanns, are simple.

Color. — Tlie nose, upper lip and upper Jaw are black; wings dark brown. Tlie
back is sooty brown; on each slioiilder, at the insertion of the wing, there is a cir-
cular black spot about 1 lines in diameter; on the under surface ciuerious brown.

38

NORTH AMERICAN FAUNA.

Dimensions. — Length of head and hody, 2 inches 5 lines; length of tail, 1 inch;
length of spread, 8 inches 8 lines; height of ear posteriorly, 4 lines; height of
tragus, IJ lines.

^Hal). — Mountains of Virginia.

Volans (Vespertilio). H. Allen, Proc. Acad. Nat. Sci. Thila., p. 282,
1866. Vespertilio volans H. Allen is another of the numerous syno-
nyms of Myofis vcdifornieus. The name was based on a specimen from
Cape St. Lucas, Lower Oalifornisx.

Yumanensis (Vespertilio). H. Allen, Monogr. N. Am. Bats, p. 58, 1864.
j\lyotis yumanensis of U. Allen is the small, lar.ye-footed bat, to which
the same author a few years later applied the name macropus, and
finally in his second Monogra]>h regarded as identical with Myotis
alheseens (Geoftroy).

In a paper published in the Proceedings of the Philadelphia Acad-
emy of Natural Sciences for 18GG, Dr. Allen gives a revised description
of M. yumanensis, based on a Fort Yuma specimen not mentioned in
the original account of the species. This specimen was Tlf. ealifornicus,
as shown by the very small hind foot which measured only two lines,
or 4.2 mm., about half as much as the foot of M. yumanensis.

LISTS OF NORTH AMERICAN VESPERTILIONIDiE.

Forty-six species and subspecies of Vesj)€rtilionida^ are here recog-
nized as occurring in America north of Panama and in the West Indies.
This numl )er will probably be materially increased when the West Indian
and Central American species are better known, and when adequate
series of skins from the nm inland ])ermit the definition of certain geo-
graphic races which doubtless exist but wliose characters can not be
determined from the material now in collections. Tlie North American
forms now known, with the names used for them by Harrison Allen in
1804, Dobson iu 1878 and Harrison Allen in 1803, are as follows:

Co7)ipi(ratirc taMe of names used for Xorlh American Tcspcrtilionida'.

H. AHen, 1864.

Names used in the present
paper.

Antrozoii8pallidna(LeCniite) Aiilrozoiis pallidns
(part).

Antrozons pallidus jiacificns Antrozons pallidas

]VIeri'iaiii.

Eiidcrnia niacnlatum (J. A.
Allen).

Corynorliinus macroti.s (Lo
Coiite).

Corynorliinus macrotis jial-
lescens subsp.nov.

Corynorliinus macrotis to \vn-
sendi (Cooper).

Myotis velifer (J. A. Allen) . .

ilyotis lucifugus (Le Conte).

(part).

Synotus luacrolis..
SynotiKs townsendi

Vespertilio lucifugus.

Dobson, 1878.

Antrozons iiallidus .

IMecotns macrotis .

H. Allen, 1893.

Antrozons pallidus

(part).
Antrozons pallidus

(part).

Euderni.a maculata.

Corynorliinus macrotis.

Coryiiorliiinis town-
sendi.

Vespertilio albcscen.s
velifer (part).

Vespertilio gryplms
^^ ..,. ,.. I) lucifugus.

^ espertiho carolii .... Vespertilio albescens

1 1, atbnis.

TABLE OF NAMES. 39

Comparative table of names used for North American VespertHionidce — Coutiuued.

Names used in the present
paper.

Myotis lucifusfus longicrus
(True).

jrv<)ti.s lucifugus alascensis
subsp. nov.

RfyotisyunianensisCH. Allen)

Myoli.s yumanensla satura-
tu.s. subsp. nov.

Mvotis californicus (Au<l. &
Bach.).

Myoti.s californicus ciliola-
brum (Merriaui).

Myotis californicus caurinus
subsp. uov.

yotia calilbniicus niexi-
caiius (Saussure).

Myolis nigricans (^faxiuiil-

i:iii).

Myotis snbulatus (Say)

Myotis snbulatus keenii
(Mcrriani).

Myotis evotis (H. Allen)

H. Allen, 1864.

Vespertilio yunianen-
sis.

Vespertilio nitidus.

Myotis thy sanodea sp. nov.

Lasionycteris noctivagans
(Le Coute).

Vespertilio suljiilatus

Vespertilio evotis.

Scotophihis noctiva-
gans.

Pijiistiellus hesperus (H. Scotophihis hesperus
Allen). i

Pipistrellus hesperus austra- '

lis subsp. nov.

I'ipistrellus sul)tlavus (F. Scotophihis georgi-
Cuvier). anus.

I'ipistrellus suhtlavns ob-

scurus subsp. nov.

ripistrellus v e r a' c r r c i s I

(Ward).

Vespertilio fuscus Beauvois., .Scotoi)hihis fuscns...

Vespertilio fuscus niirado-
rensis (H. Allen).

Vespertilio fuscus projiin-
quus (Peters).

Vespertilio fuscus baliamen-
sis subsp. nov.

Vespertilio fuscus cubensis
(Gray).

Vespertilio albigularis (Pe-
ters).

Lasiurus borealis (Miiller) . . .

Lasiurus borealis seminolus
(Rhoads).

Lasiurus borealis pfeitferi
((iundlach).

I-:isiurus borealis teliotis (H.
Allen).

Lasiurus novebora-
censis.

Dobson, 1878.

H. Allen, 1893.

Vespertilio nitidus ...

Vespertilio nitidus
longicrus.

(■Vespertilio albescens.

Vespertilio n i t i <1 u s
! aiacropus.

Vespertilio nitidus (pe-
[ domorphic variety).

Vespertilio nitidus.
Vespertilio n i t i d u s
henshawi.

i Vespertilio albescens
luelanorliinus.
Vespertilio niiiricans
(part).

Vespertilio nitidus cil-
iolabrum.

Vespertilio nigricans . I Vespertilio nigricans
j (part).

Vespertilio snbulatus. j Vesi)ertilio grypbus
(northern form).

Vespertilio evotis

Vesperugo noctiva-
gans.

Vespertilio albescens
evotis.

Vespertilio albescens
velifer (part).

Lasionycteris noctiva-
gans.

Vesperugo hesperus.

Vesperugogeorgianus Vesperugo caridinen
i sis.

Vesperugo serotinus Adelonycteris fuscus.
var.Ve.sperus fuscus.

Vesperugo propinquus

Vesjierugo albigularis

Atalapha novebora- i Atalapbanoveboracen-

Atalapha novebora-
censls var. pfeiti'eri.

Atalapha teliotis.

40 NORTH AMERICAN FAUNA.

Comparatire table of names used for North American Vespertilionida' — Contiiinod.

Names used in the present
paper.

H. Allen, 1864.

Dobson, 1878.

H. Allen, 1893.

Atalapha novebora-
censis var. frantzii.

(Saussure).

Ltiwiurus oinerons (Beauvois)

Lasinrn.s cincreus

Atalaplia cinerca

Atalapha cinerea.

Da^^vl>te^us interinodins H.
Allen.

Lasiurnn intermedins.

Atalapha inte; media . .

Da.sypterus interiiio-
diuH.

Nv(ticeiu.s Iniiiierali.s IJafl
iicsf]iie.

Nyctice.jus crepuscn-
iaris.

Nycticejus crcpuscu-
laris.

Nycticejus hiinierali.s.

mis (Guudlacli).

Yo.speiMii;o parvulus..

Kliogeessa parvula H. Allen.

Ubogeijssa alleui Thomas

1

1

List of Xortli American Vrsperliliovidn', urith tiipe localities.

Name of species Tyjie locality.

A)itro:oHs pallidus (Lo Conti-) El Paso, Toxms.

AnIrozoHs paJlidus pacificns Meniam Old FortTejou, Canada dc las Uvas, Cali-
fornia.

Eiulerma macuhitum (.T. A. Allen) Near Pirn, Ventura County, California.

Coriinorhinus macrolis (Le Conte) Georgia (prob.ibly near Rieeboro).

Cortinorhinnsmacrotis pallescensH\x\iH\}.\\o\ AKeam Canon, Navajo Couuty, Arizona.
('(ivjinorliiniis maerotis toirnsendi (Cooper).. Columbia River, Oregon.

ilijolis velifer (.T. A. Allen) Santa Cruz del Valle, near Guadalajara,

Jalisco, Mexico.

Afnotis liicifiif/iis (he Conte) Georgia (probably near Rieeboro).

]\Ii/olis hicifiu/iis ahisccnsin sulisp. nov Sitka, Alaska.

Alyolis litcifitgiis lonf/icriis (True) Pnget Sound.

Mi/otis i/iimaneiisis (II. Allen) Old Fort Yuma, California.

Mjiotis yumancnsis saturatiis snbsp. nov llamiltou, Washington.

Mi/oiis cali f or nicn s {A.nd. & Racli.) California.

Mi/otis californiciis caiirinns subsp. nov Massett, Queen Charlotte Islands, British

C<dunibia.
J^ljioiis calif ornicus ciliolahriim (Merriara).. Trego County, Kansas.
il///o/is c«///o/'«('cMs JHPXiCrtHM.s (Sanssure). ..Mexico (probably Vera Cruz, Puebla, or

Oaxaca).

jMi/otis nic/ricans (Maximilian) Fazenda de Aga, near Iritiba River, Brazil.

]\f!l<)tis NiihHhitiis{Sa.y) Arkansas River, near La Junta, Colorado.

]\fijotis siihuhiliis l:eenii (Mcrriam) Massett, Queen Charlotte Islands, Britislj

Columbia.

Myotis erotis (H. Allen) Monterey, Caliibrnia.

Mi/otis thysaiiodcs sp. nov Old Fort Tejon, California.

Lasionycleris vociirayans {he Conte) Eastern United States (exact locality un

known).

ripistrcUus licsperns (11. Allen) Old Fort Yuma, California.

ripistrellus licsperns a«s/m7is subsp. uov. ..Barranca Ibarra, Jalisco, Mexico.

ripistrcUns snbflarus (F. Cuvier) Georgia (probably near Rieeboro) .

ripistrcllnssnhjlavits ohscurus subsp. nov ..Lake George, New York.

ripistrellns vera-crucis ( Ward) Las Vegas, Jalapa, Vera Crnz, Mexico.

Vespertiliofuscns Beauvois Philadelphia, Pennsylvania.

Vesperiilio fiiscus miradorensis ( H. Allen) . . Mirador, Vera Cruz, Mexico. •?.

VesperiiUo fascns propinquns (Peters) Santa Ysabel, Guatemala. ■ ;

FAMILY VESPERTII.IONID^. 41

List of North American Vespertiliouidw, ivith type localities— Continued.

Name of species. Tyi e Ideality.

Vespertilio fiiscus iahamcnsis snbsp. nov. ..Nassau, New Providence, Baliainis.

Vespcviiliofuseus cuiensis (Gray) Cuba.

YvsperMio aUngniaris (Peters) Mexico.

Lasinrus horeaUs (Miiller) New York.

Lasiiirus horenlis seminohis (Rlioads) Tarpon Springs, Mniida.

Lasiurits horealis pfeifferi (Gundlac^li ) Cuba.

Lasiurus horealis teliotis (H. Allen) (California.

Ladiivus horealis mexicaniis (Sans^nre) Mexico (probably Vei'a Cruz, Puebla, or

Oaxaca).

Lasinrus cinercus (Beauvois) Pliiladelpbia, Pennsylvanin.

Busupier us intermedins ]i Allen Matamoras, Tamaulipas, Mexico.

Nycticeins hnmeralis Raiines(|uo Kentucky.

Nycticeiiis humeral is CH/)rt(/»s((iuudlach). ..Cuba.

L'hoyei'ssa tumida H. Allen Mirador, Vera Cruz, ]\Ioxico.

Bhofiei'ssa parvula H. Allen Tres Marias Islands, Mexico.

lihofiel'ssa firarilis sp. nov Piaxtla, Puebla, Mexico.

Ehogeissa alleni Thomas Santa Kosalia.nearAutlan,. Jalisco, Mexico.

DESCRIPTIONS.

FAMILY VESPERTILIONID.^.

Characters. — Bats with turbinal l)Oiie.s folded, bony ])alate defective
anteriorly owing to the absence of pahital processes to the i)renuixillie
(fig. L'/>); molars with cons])icnous W-

shaped cusps; tail included nearly to p^'^'^g^SkSk /V^ ?\:=iv

til) in large iutei'fenioral membrane; f'^^C^Sl M Km3 \
muzzle and nostrils variable, but Mjr^f^iM § A 1/

former never provided with distinct '/nrx' /«^ "or^

noseleaf. (^^nt^ (^^^ )v^

Rcmarls. — 'Mie family as thus de- (^Mw^ Lij C ^

fined is represented in North Amer- / •filiilllla /^illlilllf 'lltei /
ica by tliree well-marked subordinate f,„. 2._A,,tenor p.nrt of nostrum of .paces
groups, each of which may be ranked ot (a) Ph!/Uvf:toinati<i<r mid (ht ] mpn-tu-
as a subfamily. Specimens from the '°'"'^"'- ' ^'•

region in (juestion may be referred to their proper groups by tiic fol-
lowing wholly artificial key.

KFA' TO Tni-; SUBFAMILIES OF NOltTU AIMEKICAN VESrKRTILlOKID.E.

Lower incisors 4 Autrozoinw (p. 41)

Lower incisors G.

Ears Joined at anterior base riccoliud' (p. 46)

Ears separate Vesperliliouino' (p. 54)

Subfamily ANTROZOIN.^.

This subfamily is represented by the genus Antrozons peculiar to
southwestern North America. Its members may therefore be recog-
nized by their generic characters.

42

NORTH AMERICAN FAUNA.

Genus ANTROZOUS H. Allen.

1862. Antrozous H. Allen, Proc. Acad. Nat. Sci. Phila., p. 247.
ISGi. Antrozous H. Allen, Mouogr. N. Am. Bats, p. 67.
1>^1H. Antrozous Dobson, Catal. Chiioptera IJrit. Mns., p. 170.
1893. Antrozous IT. Allen, IMonogr. liats N. Am., p. 64.

Type species. — Antrozous paUidus (Le Conte).

Geof/raphic ilisirihniton. — Austral zones from Texas to the Pacific,
and from tlie Columbia lliver to (^iieretaro on the tableland of Mexico.

1-1

Generic cliar<(cte>s. — Dental formula: i, ,^ ,>; c, -i ~,\

3-3 ^,

Fig. 3 Muzzle of A7itrozous paUidus (< 2).

mammje 2; muzzle truncate;
nostrils surrounded by a
horseshoe-shaped ridge (fig,
3); lower lip free in front.

Remarls. — The genus An-
trozons differs so widely
from all others occurring in
America that it needs no
comparison with any of
these. In many ways, how-
ever, it resembles Xycfoph-
ilus of the Old World.'
While adult Auirozons in-
variably has only two lower
incisors in each mandible,
an immature individual from
Silver City, N. Mex., has a
third on the right side (fig.
4). Tlie outer lower incisor
is probably normally i)resent
in the young, though very
early crowded out by the
growth of the others.

One species and one sub-
species are known, both of
which occur in the United
States.

KEY TO TUE SUBSPECIES OF AXTKO/OITS.

Forearm 18 mm. to 'f?> mm. ; color, whitish drab gray paUulus (p. 43)

Forearm 56 mm. to 6(> mm. ; color, pale yellowish, drab-brown pacijicus (p. 45)

.Sec 11. All. Ml, :\Ionogr. Bats N. Am. (1893), p. 65.

ANTROZOUS PALLIDUS.

43

ANTROZOUS PALLIDUS (Lo Coiitc). Pale Bat.

1856. J'es2>eftirto palUdns Le Confce, Proc. Acad. Nat. Sci. Phila., VII (185-4-1855) p. -J37.

1862. Antrozoits jialUdus H. Allen, Proc. Acad. Nat. Sci. Phila., p. 247.

1864. Antrozoun paUidus H. Alleu, Monoyr. N. Am. Bats, p. 68 (part).

1878. Antrozons pallidus Dobson, Catal. Chiroptera Brit. Mus., p. 171 (part).

1893. AntrozoHS paUidm H. Allen, Monogr. Bats N. Ana., p. 66 (part).

Tjipe locaUUi.—m Paso, Texas. (Type Xo. 54G7, U. f^. National
Museum.)

Geographic distribution, — Lower Austral zone tluouuliont the desert
regioii of eastern California, i^evada, Arizona, New Mexico, and western
Texas.

General characters. — Size large (average length of forearm about 50
mm.) ; ears large, reaching 20 mm. beyond tip of nose when laid forAvard ;
color very pale drab-gray.

Ears. — The ears (PI. I, fig. 10) are larger than in any other North
American Vespcrtilionda' except the species of
riccotina'. Laid forward they extend about 20 mm.
beyond the tip of the nose. The anterior bases are
rather close together, but separate. In form the
ear is so simple as to call for no very detailed
description. Anterior l)order strongly convex im-
mediately above well-marked anterior lobe, then
almost straight to narrowly rounded-ort" tip. Pos-
terior border slightly concave immediately below
tip, then gently convex to base. Posterior basal
lobe very slightly developed. A transverse ridge
4 mm. in length extends obli<iuely upward and for-
ward from near posterior base of tragus.

Tragus long, straight, and slender. Anterior
border nearly straight to narrow tii). Posterior
border at first almost i)arallel with anterior bor-
der, then slightly convex to notch above well-developed basal lobe.
Whole posterior margin of tragus faintly crennlate.

McmJn'ioies. — Tlie membranes are tiiick and leathery, much more so
than in any of the North American Vcspertilioniiuc which approach this
species in size. Wing membranes attached at base of toes ; interfemoral
membrane at base of terminal caudal vertebra. Free border of inter-
femoral membrane considerably hmger than calcar.

Feet. — The feet are broad and strong, about half as long as tibia.
Toes armed with large claws and sprinkled with a few short hairs on
dorsum of i>halanges.

Fur and color. — The fur is sparse and short, that on middle of back
only about 8 mm. in length. It is closely confined to the body, and
extends on ears and membranes in a narrow border along extreme
base only.

Fig. 4. — Abnormal front
teeth oi All trozoiis pit l-
luliia, sliowiii}! tlirco
incisoiH on riglit side.
No. 61)119 I'min Silver
City, N, Mex. ( 10).

44

NORTH AMERICAN FAUNA.

On the back the fur is pale drab gray, most of the hairs Avitli faintly
dusky tips. Belly grayish wliite, tinged with drab on sides.

Shull. — The skull of typical Antrozoiis pallldus (fig. 5) varies in greatest
length from 18 mm. to 20 mm., and in zygomatic breadth from 11 mm. to
12.5 mm. P>rain case, rostrum, and palate broad. Length of bony
l)iilate behind molars (exclusive of median spine) usually less tiian
width at base of median spine.

Teeth. — Tlie teeth (fig. (> r() are large and strong. Upper premolar
transversely long and narrow. First lower premolar small and closely
wedged between canine and second i^remolar.

Measurements. — See table, page 16.

Flfi. Ti. — Slcnlls of (n) Aiitrozoiai palUdus and (6) Antrozous jmllidus pacificus (X 2).

Specimens examined. — Total number, 61, from the following localities:

Arizona: iVlontli of Colorado River, 1; Yuma, 5.

California: Old Fort Yuma, 1; Owens Valley, 2; Panamiut. Valley, 1; Walker

Basin, 2.
Nevada: Amargosa Desert, 1; Timpahute Mountains, 2.
New Mexico: Silver City, 4 (skins).
Texas: Conistock, 6; Devils River, 8; El Paso, 1 (skin, type); Fort Hancock,

20 (3 skins); Painted Cave, 3; Paisano, 1; Sycamore Creek. 6.

Oeneral remarlcs. — In the original description of Antrozous paJlidus
the animal is said to be a native of California, but both Baird ' iuid
Harrison Allen ^ have shown that the type specimen came from El Paso,

1 Rept. Mcx. Pound. Siirv., l\, p. 5, 1859.
^Mouogr. N. Am. Bats, p. 69, 1864.

ANTROZOUS PALLIDUS PACIFICUS.

45

Texas. Tlie type, uow iu the United States National Musenm, is in good
preservation and clearly referable to the Eastern form.

Typical Antro.zous paUidus is readily distiiignishable from .1. j>.
pacijicus by its smaller size, j)aler color, shorter, broader skull, and
narrower upper premolar.

ANTROZOUS PALLIDII.S TACIFICUS Meniam.

1864. Anirozous paUulns 11. Allcu, Monogr. N. Am. Bats, p. 68 (part).
1878. Antrozoiis pitUidiis Dobsoii, C'atiil. Chiioptera 13rit. Mus., p. 171 (part).
1893. Jntrozoiis 2>(tfl>(liis H. Alk'ii, Monogr. Bats. N. Am., p. G'o (part).
1897. Jnlrozoim jxdUdu-i pacificus Merriam, Proc. Biol. Hoc. Wasliiugtou, XI, p. 180,
July 1, 1897.

Type locality. — Old Fort Tejon, Cahada de las Uvas, California.

Geoijraphic distribution. — Austral zones in the United States west of
the Ivocky Mountains, south to Lower California and (^ueretaro.

Gencvid (7/<n-rtrfer.s'.— Slightly larger than typical Aiitrozoiis pallidus
(total length about 120 mm,, average length of forearm about 54 mm.);
color, yellowish drab brown.

Ears^ membranes, (Did distribution of
fur. — Essentially as in typical j>^f//?V//fS.

Color. — Color uniform yellowish drab
throughont to base of liairs; under
parts clear and unmixed with darker;
back strongly but irregularly shaded
by the dusky tips of the hairs.

81'uU. — Tlie skull of A ntrozous palli-
dus x>aei_ficuM{Q.g. 5 b) varies in greatest
length from 20 mm. to 22 mm., and in
zygomatic breadth from 13 mm. to 14
nun. r>rain case, rostrum, and bony
palate considerably narrower than in
typical p((llidus. Supraoccipital region
more pointed and overhanging than in tyi)ical pallidus. Length of bony
palate behind molars (exclusive of median spine) usually eipial to or
greater than width at base of median spine.

Teeth. — Teetli (fig. (Jb) essentially as in true pallidus except that all
are larger and the up])er premolar is conspicuously broader and shorter.

Measurements. — See table, page 46.

Specimens examined. — Total number, 59, from the following localities:

Calilornia: Alhambra, 1; licar Valley, 8; Berkeley, 1; Diilzura, 6; Fort
Crook,!; Fresno, 3; OldFortTejou, 6; Peso Creek, 1 (skin); Santa Barbara,
3; Santa Ysabf 1, 4 (3 skins); Witch Creek, 2.

Lower California: Cape St. Lucas, 3; Comondu, .5 (skins) ; San Fernando, 5
(Miller coll.).

Oregon: Fort Dalles, 1 (skin) ; Twelve Mile Creek, 1.

Qneretaro : .lalpan, 7.

Utah: St. Thomas, 1.

Fig. 6. — Teeth of (a) Antrozo'us pallidus and

(h) Antrozous pallidus ^jnci/i'cws ( \ 5).

46

NORTH AMERICAN FAUNA.

(ieneral rcDKo-Ks. — Antrozoun jyallidus pacifiens needs no comparison
with typical palliflns t'lutlier than that already given under the latter.

Arerai/e measureineids of suhxpecies of Antrozous paJVidns.

Subspecies.

jHlllidus

pacificus

Locality.

Texas : El Paso

Uomatock

Fort Hancoc-.k

Sycamore Creek

IJucretaro : Jalpau

■Califoruia : Santa T sabel

Dulzura

Oregou : Fort Ualles

Twelve Mile Creek ,

1 '

4?9
6??

6??

4?9
1 ?
1 1

O CS

8

^ .i

20.5

105 44. 2 21
11546 ^20
109 44.6 20.6:10
114!47.8'22.610.

10

J....I22
118 46.7 22.2
21
20

Eh

49

51

49

51

54.5

54.5

53.5

55

56

9

8.5
8.3
8.9
9.4
9.5
9.5
9
10

28 18.4111. 6 '

30 |20 |l4
28. 518. G 13
30 119 14

' Typo.

Subfamily PLECOTIN.^.

This subfamily is represented in I^orth America by two genera, Cory-
norhinus and Eaderma which may be recognized among the other
Vespertilionidw of the region by their huge ears, joined together across
the forehead. None of the North American Yes])ertilionincv show this
peculiarity.

Genus EUDERMA H. Allen.

1891. HistiotKn J. A. Alleu, Bull. Am. Miis. Nat. Hist., N. Y., HI, p. 19.5 (not Oervais

IS-).-)).

1892. Eudcrma H. Allen, Pioc. Acad. Nat. Sci. I'liila., 1891, p. 467, Jan. 12, 1892.

1893. Eiidcnna II. Allen. Mouogr. Bats N. Am., p. 00.

Type speclen. — Eudcrma macidatum (J. A. Allen).
G('0(jra])liic distrihidion. — The genus Eaderma is at present kiiowu
from one specimen taken in Ventura County, California.

Generic eharaeters. — Dental formula : i.

1-1

;imi,

3-3

,>^,^=31.

2-2
^'3 3' ^'1-1'^^'"' 2:2
Ears (PI. I, fig. 11) even larger than in Corl|norl^^nus,yn\^(i([ together
across forehead and with posterior base of tragus united with external
basal lobe. Face without evident glandular swellings.

liemarli's. — The genus Euderma resembles Corynorhinns more closely
than any other American bat, but differs in the presence of two less
premolars, in the simple nostrils, and in the more complicated structure
of the ear. Only one species is known.

EUDERMA MACULA rUM (J. A. Allen).

1891. RlsUoius maculafus J. A. Allen, Bull. Am. Mus. Nat. Hist., 111. p. 19.5, February

20, 1891.
1893. Euderma maculata H. Allen, Mouogr. Bats N. Am., p. 61.

EUDERMA MAC U LATUM. 47

Type localiUj. — N^ear Pirn, Veatura Coiiuty, California (probably at
mouth of Oastac Creek'). Type iu American Museum of Natural His-
tory (No. trifiif f.) Skull now lost.

Geo(jniphic distribution. — Emlerma nKtculatiim is known from the type
locality only.

(reneral cJiaracters. — Size large; ears about three -fourths as long as
forearm; color blackish blotched with white.

Ears. — Ears very large (PI. I, fig. 11), fully three- fourths as long
as forearm, joined together across forehead by a low band of mem-
brane; auterior basal lobe continuous with keel wbich extends upward
from anterior base of tragus and fades into substance of ear at about
terminal part of lower fourth, beyond which it continues to tip as a
well-detiued line; anterior border of ear nearly straight through lower
half, then gently convex to broadly rounded off tip; posterior border
slightly con'cave immediately below tip, then convex to base; posterior
basal lobe joined to base of tragus by a low band, below which a distinct
pocket is formed; back of this band a conspicuous ridge extending
inward toward meatus; ear membrane marked by about fifteen trans-
verse ridges; anterior margin of ear sprinkled with wliitish hairs.

Tragus nearly straight on anterior border, convex on posterior border
except at posterior base, where it is straight for a distance of 2 mm. (the
resulting form strongly suggests a table knife with short blade).

Membrancfi. — The membranes are broad and ample, the wing mem-
brane (PI. Ill, fig. 3) attached at base of toes, the iuterfemoral mem-
brane at base of terminal caudal vertebra. Free border of iuterfemoral
membrane apparently longer than calcar.

Feet. — The feet are moderately large, a little less than half as long
as tibia. Toes sparsely sprinkled with short bristly hairs on dorsal
surface and armed with strong claws. Calcar indistinct in the dried
skin, but apparently short and without keel or terminal lobule.

Fur and color. — The fur is full and soft, about 12 mm. in length on
middle of back. It extends on extreme base of ear, and on membranes
forms a very narrow border close to body.

Back very dark sepia, almost black; occiput and fore part of neck
distinctly less dark; hairs on sides and on middle of back faintly anuu-
lated with gray near tips ; patch at base of tail and on each shoulder
pure white; whole ventral surface of body white, the blackish bases of
the hairs showing through irregularly; fur everywhere blackish at
base; ears and membranes light brown.

^Icull and teeth. — As the skull of Eudcrnia maculatum is lost, I quote
the descriptions published by Dr. J. A. Allen and Dr. Harrison Allen.
The former writes :

SkuU and f?ei(/i7(o?i.— Basilar length, 16.5 inm. (0.65 in.); total length, 19 mm. (0.75
iu.); zygomatic width, 10.9 mm. (0.43 in.); height, 7.6 mm. (0.30 in.); length of

'This information was given to members of a Biological Survey party by the col-
lector of the type specimen.

48 NORTH AMERICAN FAUNA.

lower j;i\v, 12.7 iiim. (()..50 in.) ; licij^lit at coiidyh!, o.3 iimi. (().V.> in.) ; iKiii^lit at coi-o-

noid process, ,S.<S mm. (0.15 in.); len,utU of upper tooth row, G.8() mm. (0.27 in.);

lengtli of lower tooth row, 7.6 mm. (0.30 iu.).

" , , ,. 1 • • 2-2 . 1-1 1-1 3-3 14

Dental lormiila: mnsors, ^ ; canines, j_j ; premolars, .,_.;; molars, .^_^ = ^^ = 32.

The skull is thin and jiajiery, heinj^ evidently that of a young animal. The facial
portion is narrow and pointed; the brain case is (juadrato, llatttimul above, but rises
abruptly at the frontal border, the forehead being suddenly depressed.

The lower border of the zygomatic arch is curved upward; the iqjper border is
greatly expanded vertically, the upper border of the malar forming a high angular
process at the middle of arcli ; the zygomatic process of the 8(iuamosal is short, and,
with the malar, passes forward in a line nearly parallel with the axis of the skull,
with only a very slight outward curvature. The tympanic bulhe are enormously
expanded, having an autero-iiosterior length of 5.81 mm. (0.23 in.), and a transverse
breadth of 3.3 mm. (0.13 in.), their length fully ecinaling one-third of the length of
the skull. In other respects Ihe ventral aspect of thoskull ))resentsuotliing peculiar.
The lower jaw is narrow, the coronoid process small, rising luit little above the con-
dyle; the angle is well developed.

The dentition is weak, the incisors and canines being very small, relatively to the
molar series. The outer upper incisor is about one-half the size of the inner; both
have a small outer cusp at the base. The upper canine is about equal in size to the
anterior half of the upper premolar. The molars iiresent nothing distimtive. The
lower incisors are sliglitly double-notched (trifid) ; the lower canines are very small ;
the tirst premolar is about half the size of the second.

Ill liis original accouut of tlie genus Euderma^ Dr. Ihinison xillen
describes the skull and teeth as follows:

Skull. — Hraiu-case low, <|uadrate, the height one-half the bimastoid diameter.
The metencephalon as long as mesencephalon and pro-encephalon. Sagittal crest
rudimentary, does not extend beyond a line answering to the middle of the zygoma —
the remaining portions of the j)0^terior temporal crest Avidely separated — the ante-
rior not detined. Dorsum of face-vertex with a shallow concavity whicli is not
sharply defined; orbit with inflated inner wall and rugose elevated upper border;
lachrymal tubercle marked. Infra-orbital canal short; the foramen on line with
interval between second premolar and lirst molar. Line of the upper margin of the
anterior nasal aperture if produced would intersect the second premolar; tympanic
boue apparently incomplete above.

The paroccipital process bold, tren<-liant; sterno-mastoid impression deeply con-
cave; mastoid composed entirely of the S(|Uaniosal elenjent. Zygoma- (juite as iu
Cori/iiorltinns — the squamosal part twice as wide as maxillary ; sphenopalatine fora-
men present, of large size. Occipital crest trenchant. Tympanic bone greatly
inflated, e(iuals one-third the length of the skull, not touching basi-occipital, or
basi-spheuoid; excavate anteriorly. It extends to a line which answers to the
middle of the glenoid cavity. The mesopterygoid fossa as long as one-third the
distance from the posterior palatal border to the incisors. The sphenoidal foramen
is at the bottom of a deeiJ recess. The coronoid process is round, small, raised
scarcely one-third the height of the ascending ramus ; lower border of the horizontal
ramus near the angle slightly concave. The angle is raised from the plane on which
the mandible rests * " *

Uppir tectli. — Incisors contiguous, slightly inclined toward the median line, but
the lateral tooth separated from the canine by a moderate interval. Central incisor
cuspidate, with a small cuspule in-ojected midway on the posterior surface ; a distinct
cuspule also arises from the ciuguluui posteriorly. Lateral incisor oiiehalf the size
of the central, and cuspidate, with a small cuspule arising from the ciugulum on
the anterior and a .second on the posterior portion. Canine not larger than the

CORYNORHINUS. 49

secoud ])rem(>lar, the buccal surface is abruptly convex. The first premolar is small,
not wedged in, with coiu])lete cingulum. The space between it and canine narrower
tban that between it and second i)remolar. The second premolar as long as the
canine and slightly tinted. Molars as in Corynorhiniis.

Lower teeth. — Incisors crowded, trifid, i. o., the main cusp possesses a well-devel-
oped cusi)ule on each side of the base, the cingulum on the posterior side being-
large. The first and second teeth overlap for a distance eqmiling one-half of their
diameters. The third incisor retains a posterior cuspule which is larger than the
anterior and separated from the main cusj) by a wide interval. The canine is small
and jirojects but a slight degree above the incisors. It exhibits a marked cuspule
on the cingulum anteriorly.

Measurements (from skin). — Total length, 110; tail vertebra', 50;
tibia, 21; foot, 0; forearm, 50; thumb, 6.8; longest finger, 91; ear from
meatus, 31; width of ear, 22; tragus, 13; greatest width of tragus, 5.

Specimens examimd. — One, the type (Am. Mus. Nat. Hist., New York).

General remarJcs. —JEuderma maculatum differs so widely from all
other known bats that no comparison with any is necessary. Its pecul-
iar color at first suggests albinism, but since the fur is everywhere dark
at base, even in the white areas, the i)atteru is probably normal. It is
useless to hazard any conjecture as to its probable geographic range
or exact faunal x>osition.

The following note on this bat is kindly furnished by Dr. C. Hart
Merriam :

The type of this remarkable genus and species, recently described by Dr. J. A.
Allen, is believed to have been olitaiued at the mouth of Castac Creek in the Santa
Clara Valley, near San Fernando, Cal. The type specimen remains the only one thus
far collected, but the s^tecies probably ranges over much of the Low^er Souoran
Desert region in summer. While in Vegas Valley, Nev., I was told by the Stuarts,
the owners of X'^egas Ranch, that a very large bat ''with ears like a jackass and a
white stripe on each shoulder" is abundant at that place in the summer, liut docs not
occur in spring or fall. They stated that it had not yet arrived at the date of our
visit, May 1, 1891.

Genus CORYNORHINUS H.Allen.

1831. riccottis Le Conte, McMurtrie's Cuvier, Animal Kingdom, I, Appendix, p. 431

(not I'lrcotus Ceoffroy 1818).
1861. Synotns H. Allen, Monogr. N. Am. Bats, ji. 62 fnot Synotm Keyserling &. Blasius

1839).
1865. Cory iiorhin 118 H. Allen, Proc. Acad. Nat. Sci., Phila., p. 173.
1878. Pleeotnii Dobson, Catal. Chiroptera lirit. Mus., i». 177 (part).
1893. Corynorliinus H. Allen, Monogr. Rats N. Am., p. 53.

Type species. — Corynorhiuus macrotis (Le Oonte).

Geof/raphic distribution. — Austral zones throughoutthe United States,
and in Mexico south at least to Vera Cruz. Limits of range imper-
fectly known.

Generic characters. — Dental fornuila:

S-3' ^'1=1' ^^'*'3-3'^"'3-3 = ^^-
Skull (fig. 8, p. 52) slender and highly arched, the rostral portion relatively
smaller and weaker than in any other North American geuus of the
2772— No. 13 1

50

NORTH AMERICAN FAUNA.

family (with the possible exception of Uuderma, the only known skull
of which is now lost). Ears very long-, (PI. I, fig. 9), joined together
across forehead, and with tragus free from external basal lobe. Kegion
between eye and nostril occupied by a prominent thickened ridge which
terminates in a conspicuous club-shaped enlargement (fig. 7). First
phalanx of third finger shorter than second (PI. Ill, fig. 2).

General remarks. — Corynorliinus differs widely from the other known
American genera of Ve.spertilioiiida', but somewhat closely resembles
the Old World riecotus. From the latter it is separated by its ditfer-

FlG. 7. — Muzzles of (a) Plecolrig awl (h) Corynorhinvs (y2).

ently formed nostrils, conspicuously glandular muzzle, and differently
proportioned fingers (fig. 7, and PI. Ill, figs. 1 and 2). The genus is
represented by one species which may be divided into three subspecies,
differing from each other chiefly in color. All of these occur in the
United States. The material by which this genus is now represented
in collections is very unsatisfactory.

KEY TO THE SUBSPECIES OF CORTNORHINUS.

Fur everywhere distinctly bicolor macrotis (p. 51)

Fur uot (listiuctly bicolor.

Color yellowish gray lyaUescens (p. 52)

Color blackish browu townsendii (p. 53)

CORYNORHINUS MACROTIS. 51

CORYNORHINUS MACROTIS (Le Coute). Big-eared Bat.

1S31. riceotiis miurotis Le Coute, McMiirtrie's Cuvier, Animal Kiugdoni, I, Ai)pi]Klix,

p. 431 ((ieorgia).
1837. riecotus htonlii Cooper, Aiiu. Lye. Nat. Hist. N. Y., IV, p. 72. (Name proposed

as substitute for murrotis.)
18G1. Syiiofus macrotis H. Allen, Monogr. N. Am. Bats, \k (i3.
1865. Corijnorhinus macrotis H. Allen, Proc. Acad. Nat, Sci. PLila., p. 173.
1893. Conjnorhinus macroti'i H. Allen, Moiiogr. Bats N. Am., p. 55.

Type locality. — Georgia (see Proc. Acad. Nat. Sci. I'liila., 1S55, p. 434),
probably near the Le Conte IMantation, 5 miles south of liiceboro.

Geoyraphw distribution. — Lower Austral zoue in the eastern United
States.

General characters. — Size large (forearm 41 mm, to 42 mm., ear about
32); fur everywhere distinctly bicolor; general color yellowish brown.

Ears. — The ears of typical Gorynorhinus macrotis do not appreciably
difter from those of the other subspecies. They are so different from
those of all other North American bats as scarcely to recpiire detailed
description. They may be instantly recognized by their lengtli, much
more than half that of forearm, and by the form of the tragus. This
is simple, with a large basal lobe, and wholly free from the auricle. In
Euderma maculatum, the only other JSTorth American bat with ears
approaching those of Gorynorhinus in size, the tragus is joined to the
external basal lobe.

Membranes. — The membranes are broad and ample. In texture they
are remarkably thin and delicate for so large a bat. Wings (PI. Ill,
tig. 2) attached at side of metatarsus just below base of toes. Uro-
patagium extending to extreme tip of tail.

Feet. — The feet are slender, less than half as long as tibia', and armed
with strong claws. Calcar a little shorter than tibia, and about equal
to free border of interfemoral membrane. It is witliout vestige of keel
oil posterior border. The termination is obscure and without lobule.
Back of toes sprinkled with long bristly hairs.

Fur and color. — The fur is soft and silky, that on middle of back
averaging about 12 mm. in length. In distribution it shows no striking
peculiarities. It extends a short distance on the dorsal base of the ear,
but scarcely reaches the wings or interfemoral membrane.

Back uniform yellowish brown, much as in Myotis Incifuyus; the
hairs everywhere sepia through a little more than the basal half and
with very indistinctly pale tips. Belly grayish white; throat and chest
darker and more tinged with yellowish. Throughout the ventral sur-
face the fur is very sharply bicolor, the dark bases of the hairs consider-
ably darker than on the back. The light tips are too short wholly to
conceal the dark under fur. Ears and membranes light brown,

81cull and teeth. — The skull (fig. 8) and teeth (fig. 9) have been suffi-
ciently described under generic characters. An adult skull from
Houma, La. ( $ No, 45894, United States National Museum, Biological
Survey collection), measures: Greatest length, 16; zygomatic breadth,

52

NORTH AMERICAN FAUNA.

U; breadth of lostrum at posterior border of large premolar, 5; man-
dible, 10.(5; upper tootb row (exclusive of iucisors), 5.1'; lower tooth
row, (5.8.
Measurements. — !See table, page 54.

Specimens examined. — Total number, 9,
from the following localities:

Alabama: Grceusboro, 1 (skin, MLTriam coll.).

Kentucky: Bowling Green, 1.

Louisiana: Houma, 4 (skins).

Mississippi: Bay St. Louis, 1.

South Carolina: Hardteville, 1 (skin, Miller

coll.).
Virginia: Dismal Swamp, 1 (skin).

General remarks. — Tyi^ical Gorynorhinus
macrotis, like the western subspecies, is dis-
tinguishable at a glance from all other North
American bats by its generic characters.
Among the forms of Gorynorhinns it maybe
distinguished by its conspicuously bicolored
iur. The liniits of this animal's range are
not well understood, but it is probably a
characteristic sjiecies of the Austroriparian
fauna.

CORYNORHINUS MACROTIS PALLESCENS
STibsp. nov.

1864. Synotus iown-seniJi It. Allen, Monogr. N. Am.

Bats, p. 65 (not Plevotus totvnsvndi Cooper, 1837),

1893. Coryuorhiuus townsendi II. Allen, Monogr. Bats N. Am., ji. 58.

Ty2)e from Keam Canon, Navajo County, Arizona. Adult 9 (skin), No. 65534, U. S.

Nati<mal Museum (Biological Survey t-ollection). Collected August 3, 1894.

by Dr. A. K. Fisher. Collector's No., 1715.

Qeo(jrapliic distribution. — Probably through-
out the Austral zones from California, Colorado,
and western Texas to southern Mexico.

General characters. — Similar to typical ma-
crotis, but much paler in color; fur nowhere
distinctly bicolor.

Golor. — Back yellowish drab gray, becoming
paler about head, the hairs with faintly defined
light ijlumbeous bases and faintly darker tii)s.
Belly slightly i^aler than back, but hairs with
out distinctly lighter tips and with darker
basal area so ill defined as to be scarcely visible
light brown.

Measurements. — See table, page 54.

Specimens examined. — Total number, 30, from the following localities

Arizona: Fort Huachuca, 2; Keam Canyon, 1 (skin).
California: Dulzura, 1; Owens Lake, 1; Owens Valley, 1.

Fig. 8 — Skulls of (a) Oorynorh^nus
to%vnsendii and (b) C. macrotis { 2).

Km. 9.— Teeth of (a) Gorynorhi-
nus toumsenilii and (b) C.)na-
crotis ( X 5).

Ears and membranes

OORYNORHINUS MACROTIS TOWNSENDII. 53

Colorado: Larimer County, 1 (skiu, Miller coll.).

Guanajuato: Santa Rosa, 17.

Miclioacan: Patzcuaro, 1.

Oaxaca: Oaxaca, 1.

San Luis Potosi : Hacienda La Parada, 1.

Texas : East Painted Cave, 1.

Utah: 1.

Vera Cruz : Jico, 1.

General remarls. — Corynorhinvs macrofis pallescens differs from tine
macrotu in its iiuicli ])aU»r, more uniforni coloration. vVliile this form
is represented in the National Musenm by numerous specimens in alco-
hol, the series of skins is very incomplete. Until this series can be
o;reatly increased the limits of distribution of the subspecies must
remain ])urely a matter of conjecture.

The differences in form of the inner upper incisor which have been
suppot-ed to distinguish this race from true macrotis^ appear to be
inconstant. While the few specimens of typical macrotis that 1 have
seen have this tooth without exception bicuspidate, western specimens
vary greatly. In a series from
Santa Kosa, Guanajuato, both ex-
tremes are represented, while sev-
eral specimens are with difficulty
referred to one or the other

(tig. 10). FlQ. 10. — Left upper incisors of Corynorhiniis ma-

This is the bat to which Dr croUs pnlleaccns fmm Santa Ro.sa, Guanajuato,

TT • n 1 !• 1 1 Mexico.

Harrison Allen has applied the

name foivtisendi. True fonmsendi, however, the type of which came

fro'.n the Columbia River, is the diirk northwest coast form.

CORYNORHINUS MACROTIS TOWNSENDII (Coo])(n).

1837. Plccotiis townsendii Cooper, Ann. Lye. Nat. Hist., N. Y., IV, ]». 73. (Columbia

River.)
1878. Plecotns macroiis Dobson, Catal. Chiroptera Brit. Mus., p. ISO (not Pleeotus

macroth Le Conte, 1831).

Type locolity. — Columbia River, Oregon.

Geoyraphh- disfribiitiou. — Humid coast district of Oregon, Washing-
ton, and southern British Columbia.

General char act ers. — Similar to typical macrotis^ but much darker ; fur
nowhere distinctly bicolor.

Color. — Back uniform dark brown, the hairs indistinctly light plum-
beous at base, and very faintly tipped with yellowisli. Belly dark
brown, the hairs light plumbeous at base. Ears and membranes
blackish.

Measurements. — See table, page 54.

Spechnens examined. — Total number, 3, from the following localities:

British Columbia: Comox, 1 (skin).
Oregon: Creswell, 1; Gold Beach, 1.

'H. Allen, Monogr. Bats N. Am., p. 58, 1893.

54

NORTH AMERICAN FAUNA.

General remarlis. — GorynorMnus macroUs toumsendii is the dark north-
west coast representative of the species. In coloration it bears a close
resemblance to the other bats ijeculiar to the same region. It is so
different in general appearance from macroUs and pallescens that it
needs no special comparison with them. More material is necessary
before the exact relationship of this form to the others can be determined.

While this is the true toirnsendii of Cooper, it is not the bat to which
Harrison Allen has applied the name. The latter is G. macrotis pal-
lescens. It was on a specimen of this form from Vancouver Island that
Dobson based his descri])tion of '■Flecotns^ macroUs in the Catalogue of
Chiroptera in the British Museum.

Average measurements of subspecies of Corynorhinus.

Siibspecies.

pallescens . . .

I^ocality.

Soutli Carolina : Harrteeville

Kentucliy : Bowling Green

Louisiana : Houma

Arizona : Keam Canyon

Fort Huacliuca

Coloi ado : Larimer County

Texas: East Painted Cave

Guanajuato : Santa Rosa

British Columbia : Comox

Oregon : Gold Beach

Creswell

1?

IcT

4??

19'

1,/

1?

1?

10

1?
1?
1?

105 52

106 !48. 5 21.

105
95

90 |45
59.3 48.1

105
100

Ph

41
43.4
41.5
42. G

44

41

41.9

42

44

73

77

73

76

71

77

73 33

77 33.9

76..

80 33

77 34

20
23.7

14.6
12.7

' Type.

Subfamily VESPERTILIONIN.^.
This subfamily contains the great majority of North American Yes-
pertilionidw. Those of its members that occur in the region in ques-
tion may be known by their simple muzzles and nostrils, separate ears,
ample interfemoral membranes, and six incisors in the lower jaw.

KEY TO GENERA OF VESPERTlLIONINiE OCCURRING IN AMERICA NORTH OF

PANAMA.

[Based on dental characters only.]
Upper incisoTS 2.

Upper premolars 2.

Mandilmliir tooth-row more than 8 mm Dasypiiriis (p. 115)

Mau(lil)ular tooth-roAv les.s than 7 mm.

Third lower incisor much smaller than second or i\vst . . Iihof/ei'ssa (p. 122)

Third lower incisor abont 0(iual to second or first Xycticeius (p. 118)

Upper premolars 4 Lasimnis (p. 105)

Upper incisors 4.

Upper premolars 2 Vespertilio (p. 95)

Upper premolars more tlian 2.
Upper premolars 4.

Lower premolars 4 Fipistrellus (p. 87)

Lower jiremolars 6 Lasioni/rteris (p. 85)

Upper jiremolais 6 Myotia (p. 55)

MYOTIS.

55

Fig. 11. — Skulls of (a) Myotig myotis, (h) M. thysanodes, and (c)
M. ni(jricans ( > 2).

Genus MYOTIS Kaup.

1829. Myotis Kaup, Skizzirte Entw.-Gescli. u. Natiirl. Syst. tl. Enrop. Thierw., I, p.

106. Tj'pe VespertiUo murUms Schreber (not V. mnrinus Liuu.).
1829. Nyatactes Kaup, Skizzirte Eutw.-Gesch. u. Natiirl. Syst. d. Europ. Tliierw., I, p.
108. Type Vesiieriilio
hechsteinii Leisler.
1839. VespertiUo Keyserling »fe
Blasius, Wiegmann's
Archiv f. Naturgescb.,
5ter Jahrg., Bd. 1, p.
306 (not VespertiUo
Linn:eus, 1758).
1841. Selysius Bonaparte, Icon-
ografia Fauna Italica,
I, Introduzione []>. 3].
Type VespertiUo mysta-
einus Leisler.
18.56. lirachyotKS Kolenati, All-
gem. Deutscli. Natur-
hist. Zeitg., Dresden,
Neue Folge, II, pp.
131, 171-177. Based on the species mystaciniis, daiihentonii, and dasycneme.
1856. Isotns Kolenati, Allgeni. Deutsch. Naturhist. Zeitg., Dresden, Neue F<dge, II,

pp. 131, 177-179. Included the species nattereri and emarginatus.
1864. Vespei'tiUo H. Allen, Monogr. N. Am. Bats, p. 46. (Not VespertiUo Linn., 1758.)
1870. Aeorestes Fitzinger, Sitzungsber. Math. -Nat. CI. K. Akad. Wiss., Wien, LXII,

Abth. I, pp. 427-436. Included the
species viUosissimus, aUicsccns, and
iiigricans.
1870. Comastes Fitzinger, Sitzungsber.
Math. -Nat. CI. K. Akad. Wiss.,
Wieu, LXII, Abth. I, pp. 565-.579.
(Included caparcinii, mcyopoilius,
(hisycneme and liitniophilit'i.)
1878. VespertiUo I)ol)Soii, (!atal. Chiroptcra
Brit. Mus., p. 284 (not I'espertiUo
Linnieus 1758).
1893. I'espertiUo H. Allen, Monogr. Bats N.
Am., p. 70. (Not VespertiUo Linn.,
18.58).

Type species. — Myotis myotis (Bech-
steiu)= VespertiUo murines Sclireber
et Auct., uec Linn.

Geogyaphic distribution. — Temiier-
ate and tropical parts of both liemis
pberes.

Generic characters. — Dental form-
ula :

Fifi. 12. — Skulls of (a.) Myotis nigricans, (6) M.
tliysanodes, and (c) M. myotin (X2).

2-2

1-1

m.

3-3

=38.

3-3
7 3_3; ^7 i_i5 P'"i 3_3J ""J 3_3-

Remarks. — The North American members of the genus Myotis are all
small, delicately formed bats, whicb, aside from their dental formula,

56 NORTH AMERICAN FAUNA.

shared hy no other American genus of Vespertilionidw, are usually
recognizable by their slender forms, long tails, hairy faces, narrow ears,
and tapering, straight, or recurved tragi. As I have had no opportu-
nity to study any of the Old World species except M. myotis, I can
make no attempt to define the genus in detail, but the dental formula,
coupled with the characters of the family, is sufficient to distinguish
the genus among American bats. The species of Myotis differ greatly
among themselves, especially in size (see figs. 11 and 32). It may
eventually prove necessary to divide the genus into two or more sub-
genera.

KEY TO NORTH AMERICAN FORMS OF MYOTIS.

Free border of nropatagium conspicuously fringed ilnjsanodcs (p. 80)

Free border of nropataginm not conspicnonsly iVinged.
Forearm more tban 40 mm.

Ear wben laid forward extending barely beyond nostril relifer (p. 5fi)

Ear wben laid forward extending 7 to 10 mm. beyond nostril erotis (p. 77)

Forearm less tban 40 mm.

Ear wben laid forward extending 7 to 10 mm. beyond nostril erotis (p. 77)

Ear wben laid forward extending less tban 6 mm. beyond nostril.

Fnr on back not distinctly bicidor nUjricans {]>. 74)

Fur on back distinctly bicolor.
(ieneral color wbitisb gray.

Foot 8 to 10 mm yiimanensis (p. 06)

Foot ."> to 7 mm ciliolahnnn (p. 72)

General color never wbitish gray.
Forearm 30 to 35 mm.

C<dor ligbt yellow isb gray colifoniiciifi (p. 69)

Color yellowisb brown.

Forearm 31 to 33 mm cauriniis ( ]>. 72)

Forearm 32 to 36 mm merican us (p. 73)

Forearm 34 to 40 mm.

Tibia less tban 15 mm ..saiuralns (p. 68)

Tibia 15 to 19 nnn.

Ear and tragus slender, tbe latter 9 mm. or more, in length.

Color ligbt brown siihulatus (p. 75)

Color blackish leeiiii (ii.77)

Ear and tragus short and broad, the latter 8 nnn. or less in length.

Tibia, 17.5 to 20 nun lonrficnts (p. 64)

Tibia 15 to 17 nnn.

Ear 12 to 14 mm liicifiifius (p. 59)

Ear 14 to 16 mm ahtscoisis (p. 63)

MYOTIS VELIFER (J. A. Allen).

1890. Vespcrniio relifer J. A. Allen, Bull. Am. Mns. Nat. Hist., N. Y., Ill, p. 177, D<'C. 10,

1890.
1896. Vespn-tiVio incantns J. A. Allen, Bull. Am. Mns. Nat. Hist., N. Y., VIII, p. 239, Nov.

21, 1896. (Texas.)

Type locality. — Santa Cruz del Valle, near Guadalajara, Jalisco,
Mexico.

Geoyrapliic (Ustrihution. — Near border line between upper and lower
Sonoran zones from Missouri and Indian Territory south to Hidalgo,
northern Michoacan, and the City of Mexico.

MYOTIS VELIFER. 57

General characters. — Largest species of Myoti.s known to occur in
Mexico or the United Sttites. Lengtli 90 to lOo, forearm 40 to 47.
Calcar slender, without well-developed lobe. Free border of uropata-
gium naked. Ears short, reaching tip of nose. Wings from metatarsus.

Ears. — The ears are short and pointed ; laid forward they reach to or
just beyond nostril. Auricle concave on both sides iiiinudiately below
the narrowly rounded oif tiip (most strongly so on the posterior border).
Anterior l^order straight or very slightly convex through the basal two-
thirds of its lengtli, then concave or almost straight to tip; posterior
border strongly concave directly below tip, then still more strongly con-
vex to basal notch, tlie widest part of the ear at about mid height. Basal
notch well defined, isolating a prominent basal lobe, which is slightly
notched on the lower side and joins the side of the face in a line which
if continued would coincide with the margin of the upper lip.

Tragus moderately long and broad, the anterior < dge straight or very
slightly convex toward the tip; posterior border with a well developed
lobe at base; just above the lobe the tragus attains its greatest width;
the two borders are usually parallel for a short distance, after which
the posterior border bends rapidly forward to the tip, below which it
may be either straight or very slightly concave.

Mnnhrancs. — Tlie membranes are, for an American Myotis, rather
thick and opaque. The uropatagium is sparsely haired both dor-
sally and ventrally on its proximal fourth; the free border, which is
distinctly shorter than the calcar, wholly naked. Wing from point
between aidde and base of toes, but nearer the latter. When drawn
away perpendicularly from the leg, the wing appears to be attached to
the ankle.

Feet. — Feet (PI. II, fig, 0) large and strong, half as long as tibiae Toes
(without claws) slightly longer than sole, scarcely united by memlmme
at extreme base, all sprinkled with long, stift' hairs. Caltar long, slen-
der, usually terminating indistinctly, but sometimes with a more or less
well defined lobule at tip.

Ffir and color. — There is nothing i)eculiar about the distribution of
the fur to distinguish this bat from other American members of the
genus.

In color the fur is dull sepia throughout, paler on the belly, the hairs
everywhere dusky slate at base. Individual variation is trifling and
is mostly confined to the belly. This is usually dull broccoli brown,
but in some specimens by admixture of yellow it becomes more nearly
isabella color.

8J:uU. — Skull stronger and more heavily built than in any other
Myntis found in Mexico or the United States, but not actually larger
than that of M. thysanodes. Greatest length (5 specimens) 10 to 10.4;
zygomatic breadth, 10 to 11; interorbital constriction, 4 to 4.2; width
of rostrum at anterior root of mj^, 0 to 0.2; length of mandible, 12 to
13. When viewed from above, the brain case is subcircular in outline

58

NORTH AMERICAN FAUNA.

but truncate posteriorly and slightly longer than broad. Forehead
moderately elevated above muzzle; occiput high and compressed, with
well-developed ridges. Distance from posterior border of last upper
molar to tip of hamular equal to or less than distance between alveoli
of posterior molars.

Teeth. — Upper incisiors diverging at tips, the inner tooth of each pair
much the larger. First and second premolars in the tooth row or
second displaced slightly inward and partly concealed by the anterior
edge of the third, the relative size of the two teeth variable, but first
always the larger. The second premolar is always much shorter than
the first, but the cross section of the crown is sometimes nearly equal
in the two teeth. On the other hand, in rare cases the area of cross sec-
tion in the first is nearly double that of the second. Third upper jire-
raolar triangular in outline, posterior margin longest, the outer margin
abruptly convex anteriorly, the anterior and posterior borders slightly
concave; inner apex of triangle bluntly rounded and not reaching to
level of inner margins of molars. Crowns of first and second upper
molars trapeziform in outline, the anterior edge longest, and the i^oste-
rior, outer, and inner edges successively shorter. Anterior edge slightly
convex, posterior edge slightly concave. Crown of first molar consid-
erably shorter in proportion to its width than second (fig. 14 d, j). 6L).
Inner mandibular incisors smallest, their crowns compressed and trifid;
middle incisors similar but larger; outer incisors about as large as the
two others together, their crowns irregularly terete, and with four indis-
tinctly developed tubercles, one of which is much larger than the others.
First and second premolars perfectly in line, the first considerably larger
than the second, though not much wider in cross section. Third pre-
molar as broad as long, trapeziform, the posterior margin longest.

Mcafiurenienfs. — In the following table average measurements are
given of 20 specimens of MyoUs velifer from six localities.

Average inea>iurements of 20 specimens of Myotis relifer from 6 locaHties.

Locality.

o ^

g

o
H

> ^

'5
H

3

1

£

o

a

§
1-)

C3

a

CS

i

o

3

5

3

IcT

4

1'

4

;!

99

97

94

98.8

95.2

93.3

93.6

44.8

42

40

42

44.4

41.3

39.6

18.5

18.3

17.0

16.5

18

18.1

17.9

9.5

8.9

8.6

9

8.4

9.3

9.8

45

43.6

42

41.5

43

43

42.6

7.2

7.2

6.4

6.4

7

6.9

7

73.4

71.3

71

69

62

67

69.3

16

16.1

15.4

10.6
10.3
11

9

8.5

Mexico : Ixtapalajiii

9

7.8

Missouri : Marble Cave

15.6

10.1

8.1

'Type of 'incautus.^

MYOTIS LUCIFUGUS. 59

Specimens examined. — Total number 46, from the following localities :

Arizona: San Bernardino Rancli (Monument 77, Mex. Bound. Line), 5 (skins).

Hidalgo: Tulancingo, 4 (1 skin).

Indian Territory : Fort Reno, 3.

Jalisco: Guadalajara, 2 (skins, Merriam Coll.) ; Santa Cruz del Valle, 3 (skins,

Am. Mns. Nat. Hist.) ; Hacienda San Marcos, Tonila, 1 (skin, Am. Mus.

Nat. Hist.).
Mexico: Lerma, 1; Ixtapalapa, 1; City of Mexico, 1 (skin).
Michoacan: Patzcuaro, 11 (3 skins).
Mi^<sou^i : Marble Cave, Stono County, 3.
San Luis Potosi: Abualulco, 1.
Texas: Mouth of Pecos Eiver, 4; New Braunfels, 1; San Antonio, 4 (skins,

Am. Mus. Nat. Hist., including type of ^incaHtus').
Vera Cruz: Las Vigas, 1.

General remarlcs. — Through the kindness of Dr. J. A. Allen, I have
been able to examine two of the original specimens of Myotis relifer
from Santa Cruz del Valle, Guadalajara, Jalisco, and four specimens
(iucludiiig the type) of ' VesperUUo ineautus'' from San Antonio, Texas.
After comparing the specimens of '■ineautus'' with seven M. relifer
from various parts of Mexico, I can find no characters to separate the
two even subspecifically. In size as well as in cranial and dental
characters they agree i^erfectly, while the difference in color is too
sliglit to be described by words. As the specimens from Mexico were
all taken in midsummer and those from Texas were killed in October
the variation in color is probably seasonal.

The large size of this bat distinguishes it at a glance from all other
Mexican or United States species except M. tkysanodes. From the latter
the darker color, slender ciilcar, naked free border of interfemoral mem
brane, and shorter ears and tragus separate it without difficulty.

Myotis velifer, while totally different from all other bats found in
Mexico or the United States, is doubtfully distinct from the South
American ]\[. albescens. Mr. Oldfield Thomas, who has compared for
me specimens of the former with the albeseens in the British Museum
so named by Dobson after examination of the type, writes that M. vel-
ifer and jIA. albescens are iiractically identical. It is best, however, to
retain the name velifer for the bat occurring in Mexico and the United
States until the South American species has been positively i<lenti tied.

Dr. Harrison Allen refers to this bat in his recent monograph as
Vespertilio albescens velifer. Under the same name he mentions a
specimen of 71/. thysanodes from 'Dalyura' (=Dulzura), Oal. ; while the
Texan specimens of velifer in the Biological Survey collection he has
labeled ' V. albescens f '

MYOTIS LUCIFUGUS (Le Conte). Little Brown Bat.

1831. Vespertilio hicifwius LeConte, McMurtrie's Cuvier, Animal Kingdom, T, Append.,

p. 431. (Southern Georgia.)
1856. Vespertilio snbitlatus Le Conte, Proc. Acad. Nat. Sci. Phila. (18.54-55), p. 435.
1864. Vespertilio nffinis H. Allen, Monogr. N. Am. Bats, p. 53.
1864. Vespertilio lucifugus H. Allen, Monogr. N. Am. Bats, p. 55.

60

NORTH AMERICAN FAUNA.

1S78. Vespertilio carolii Do1)Sou, Catal. Chiroptera Brit. Mns., p. 325.

1893. VespcrtiUo gnjphiis Var. (a) Vespertilio gryphus lucifufjus H. Allen, Monogr

Bats N. Am., p. 78.
1893. VespertiUu albescens affinis H. Allen, Monogr. Bats N. Am., p. 93.
1897. Vespertilio Incifugus austroriparim Khoads, Proc. Acad. Nat. Sci. Phila., p. 227,

May 1897. (Tarpon Springs, Florida.)

Type locality. — Georgia, probably soutliern Liberty County.
Geogrftphic distribution. — The wliole of North America north of the
southern boundary of the United States, except in the Rocky Moun-
tains and on the Pacific coast of ('alifornia, Oregon, Wasliingtou
British Columbia, and southern Alaska.

General characters. — Size medium; length 80 to 90; forearm 36 to
40; tibia 14.6 to 16.6. Calcar vslender, indistinct, about equal in length
to free border of uropatagium, usually terminating in a iaintly indicated

lobule; keel very slightly developed, if
at all. Free border of uropatagium
naked. Ears short, laid forward they
reach about to nostril.

Ears. — The ears (fig. 13, c) are short
and ])()int('<], reaching when laidforward
barely to tip of nose. The anterior bor-
der is straight from base through lower
third, then for a short distance strongly
convex, and finally straight to narrowly
rounded off tip. Posterior border gently
con<;ave from just below tip to about
middle, where it becomes convex and
continues so to basal notch. Basal
notch moderately developed, isolating
a broad but not conspicuous basal lobe.
Tragus short, blunt, bent slightly
forward. Anterior border straight or
slightly concave from base to tip. Pos-
terior border straight or slightly con-
vex immediately below tip, then more
strongly convex to notch above large and prominent basal lobe. Greatest
width of tragus through basal lobe or at about middle height, accord-
ing to convexity of posterior border.

Membranes. — Membranes rather thick and leathery, entirely naked
except where fur of body extends in a narrow line at the base of the
wings and uroi^ataginm. On the latter the fur occupies about the
basal fourth on the dorsal side, rather less ventrally. The wings are
attached at the base of the toes.

Feet. — The feet are large and strong, slightly more than half length
of tibite. Toes longer than sole, joined by membrane at base to a
point Slightly beyond middle of proximal phalanges. The membrane
extends farther on first digit than on fifth.

FiQ. 13.— Ear of (a) Myotis subulatns, (h)
M. keenii, (c) M. liieifmjns, .and (rf) M.
alascensin ( ' '_').

MYOTI8 LUCIFUGUS.

61

Fur and color. — The distribution of the fur in Myotis lucifugus is in
no way peculiar. Tlie Irairs are everywhere dusky slate at base.
General color dull brown with a distinct gloss in certain lights, the
ventral surface paler and more yellowisli. The exact shades are vari-
able. Thus in three specimens taken at Washington, 1). C, in June,
the color ot the back is respectively wood brown, raw umber, and
sepia, the belly in each pale wood brown tinged to a varying degree
with gray. In the majority of individuals the color tends toward
sepia. Seven skins from Elk Eiver, Minn., and three from Kadiak
Island, Alaska, are indistinguishable in color from those taken at
Washington. Ears and membranes light brown.

SJcuU. — The skull of Myotis lucifugus is characterized by the broad

Fig. 14. — Teeth of (a) Myotis yumanensit, (h) M. lucifugus, (c) M. lucifugji.s lonijknis, aud
(d) M. vcUfcr (X 5).

muzzle and palate and gradually sloping forehead. In most specimens
the face line begins to rise almost from the tip of the muzzle; iu others,
however, there is a short flat area back of the nasal opening. The
l)rain case is broad and inflated at the back, less so in front, produc-
ing in many individuals a wedge shaped outline. Distance from pos-
terior molar to tip of hamular less than distance between j)osterior
molars.

The skull of Myotis lucifugus differs from that of M, suhulatus in its
slightly smaller size, broader palate and muzzle, and less abruptly
elevated face line.

Tee/Zi. —Upper incisors diverging at tips (fig. 14 i). Crown of first
bicuspidate, and, when viewed from below, nearlj^ rectangular and

62 NORTH AMERICAN FAUNA.

iibout twice as long as broad. Larger cusp i)laced at extreme anterior
end; tbe smaller one on tbe inner edge near posterior border. Crown
of second incisor subterete, nnicuspidate. Crown of first premolar
longer than second when viewed from tbe side, sligbtly larger, or iu
rare cases very mucli larger tbaii tlie latter in cross section; tbe two
teetb in line, or second sligbtly dis^daced inward. There is nothing
characteristic in the form of tbe third premolar or of the molars.
JVIaxillary incisors as nsnal in tbe genus, tbe middle pair and tbe next
compressed and tritid, tbe outer much larger and subterete. First
maxillary premolar with crown longer than the second when viewed
irom tbe side, and one fourth to one-third larger in cross section; the
two teetb perfectly iu line, or second slightly displaced inward. Third
premolar subquadrate, nearly as broad as long. The lower molars
show no distinctive characters.

As compared with M. siilmlatus, the species with which it is most
likely to be confused, Myoiis lucifuyus shows numerous differences in
dental characters. The tooth row, as a whole, is shorter, and the indi-
vidual teetb relatively smaller. Tbe first and second premolars in each
jaw are actually smaller, and in most specimens more nearly ec^ual iu
size in cross section, in M. suhulatua tbe premolars are so large that the
second is often crowded inward from tbe tooth row, a condition rarely
seen in M. luci/ugns. The form of tbe third lower premolar is very
difi'erent in the two species. When viewed from tbe side, this tooth
is conspicuously broader in proportion to its heigh in 71/. snhulatu.s.
When viewed from above, the tooth is much larger in M. subiilidus, and
distinctly longer than broad, whde in M. luci/iujus it is nearly as broad
as long.

Measurements. — See table, page (J5.

JSpecimens examined. — Total number 5615, from tbe following localities:

Alabama: Greeusboro, 1 (skiu, Merriain coll.)-

Alaska: Kadiak Lslaiul, 9.

Conuecticnt: 1 (Merriani coll.).

District of Columbia: Wasbiugton, 20 (majority in Merriam coll.).

Florida: Tarpon Springs, 7 (two skins, Rboads coll., type and topotypea of

* aiistroriparius') .
Illinois: Warsaw, 141; West Nortb field, 2.
Kentucky : Mammotb Cave, 218.
Maine: Eastport, 1.
Maryland: Seneca River, 1.
Massachusetts: Woods Hole, 1.
Minnesota: Elk River, 7; Fort Snelling, 4.
Newfoundland: Bay St. George, 4 (skins. Bangs coll.).
New York : Adiroudacks, 1 (Merriam coll.) ; Big Moose Lake, 1 (Merriam coll.) ;

Catskill Mountains, 2; Howes Cave, 2.5 (Merriam coll.); Lake George, 1;

LocustGrove, 9 (Merriam coll.); Lyons Falls, 4 (Merriam coll.) ; Oneida Lake,

63 (Miller coll.); Peterboro, 1 (Merriam coll.); Sing Sing, 1 (Merriam coll.);

AVest Point, 1.
Nortb Carolina: Roan Mountain, 1.
Nova Scotia : Halifax, 1 .

MYOTIS LUCIFUGUS ALASCENSIS. 63

Ontario: Gravenliuist, 1 (Miller coll.); James Bay, 2; North Hay, Lake Nipis-

siiig, 1 (Miller coll.).
Peunsylvania: Bradford, 1; Center County, 19.
Qnel>ec: Godbout, I (Merriani coll.); Ottawa, 2 (Merriani coll.).
South Carolina: Beaufort, 3.
Virginia: Riverton, 1.

General rcmarlis. — Myotis lucifugns resembles M. velifcr more closely
tluui it does any other North American species. From the latter it is,
liowever, readily distinguishable by its much smaller size. From M.
siibulatus, the only species of the genus with which it is associated in
the eastern United States, it may be at once recognized by its shorter
ear and shorter, less acuminate tragus.

This bat is the Vespertilio gryiilms lucifugus of Dr. Harrison Allen's
recent monograph. Dr. Allen's 'northern form of Vespertilio (jryphns''
is M. suhulatuK.

Through the kindness of Mr. S. N. Ilhoads I have examined the type
and six topotypes of Vespertilio lucifugus austroriparius from Tarpon
Springs, Fla, 1 can find no characters by which these specimens may
be distinguished from those taken at other parts of the range of Mijotis
lucifugus. The two skins, one of which is the type, are those of partly
grown individuals whose immaturity is clearly indicated by the soft,
pai)ery skulls in which the nasal sutures are still clearly visible, and
by the imperfectly formed joints of the fingers (see fig. 1, p. 9). These
specimens differ from northern adults in smaller size, shorter fur, and
duller, browner color. Three adult topotypes in alcohol show only one
of these peculiarities — the shortness of fur — and in the fourth this also
is absent. The fifth alcoholic specimen is immature. That the adult
specimens of My ot is from Tarpon Springs are not smaller than Af. luci-
fugus from other localities is clearly shown by the table of measurements
on page 05. The short fur of three of the adults is evidently a seasonal
character, since all showing this peculiarity are in worn, ragged coat,
while the only one in fresh pelage (killed September 12) has fur of
the ordinary length. The fur of all these specimens, after nearly five
years immersion in alcohol, has lost the warm, glossy appearance char-
acteristic of freshly killed individuals. It can be perfectly matched,
however, among the series of alcoholic specimens collected in Center
County, Pa., during the winter of 1893.

MYOTIS LUCIFUGUS ALASCENSIS subsp. nov.

Tfipe from Sitka, Alaska. Adult 9 (in alcohol), No. 77416, U. S. National Museum
(Biological Survey collection). Collected August 5, 1895, by C. 1'. Streator.
Collector's number, 4754.

Geographic (listrihution. — Humid coast district of southern Alaska
and northern British Columbia.

General characters. — More like typical lucifugus than like longicrus,
but darker in color and with longer ears.

Ears. — As shown in the table of measureuients on page G5 the ears
of this form average distinctly larger than those of the typical sub-

64 NORTH AMERICAN FAUNA.

species. 1 cau uot see, liowever, tliat they differ appreciably in form
(fig-. i3d).

Fur and color. — The fur is distributed as in true hicifuyuH. In color
it is evidently darker thau that of the typical form, but the exact
differences can not be determined from specimens in alcohol. Ears and
membranes blackish.

Measurements. — See table, page 05.

Specimens examined. — Total number IG, from the following localities:

Alaska: Fort Wrangel, 1 (skiu, Merriaui coll.); Loriug, 4 (1 skin); Sitka, 8

(3 skins).
British Columbia : Massett, Queen Charlotte Islands, 3.

General remarls. — Myotis lucij'iujus alascensis is distinguishable from
both typical M. hicifiKjus and M. lucijuijus lon<jicrus by its longer ears
and darker color. From il/. luci/ugus longicrus it differs further in its
much shorter tibia.

MYOTIS LUCIFUCUS LONGICRUS (True).

188G. I'espertUio louf/icrus True, Science, VIII, p. 588, Dec. 24, 1886.

1893. J'esjK'rlilio nitldus loni/icru.s II. Alien, ^louogr. Data N. Am., p. 103.

1893. VeispertUio alhtsceiis (melauic phase) H. Allen, Monogr. Bats N. Am., p. 92 (part).

Ty2)e locality. — Puget Sound.

Geographic distribution. — Boreal and Transition zones from Puget
Sound east to Wyoming; south at least to Arizona and southern Cali-
fornia, and probably much farther.

General characters. — Similar to typical Myotis luci/ugus, hat larger
(length, 94: to 103; forearm, 37 to -10; tibia, 17.8 to 19), and with longer
tibia and proi)ortioually shorter ear and forearm.

Ears. — The ears are more rounded and proportionally slightly sliorter
than in typical M. luci/ugas, the inner side of the conch usually more
hairy. Tragus as in M. luci/ugus.

The membranes and feet differ in no way from those of the true
31. luci/ugus^ except that the feet appear shorter in proportion to the
tibia'.

Fur and color. — The fur shows no peculiarities in distribution. In
color it is darker and duller than in the tyj)ical subspecies (especially
in specimens from northern California), but the difference is apparently
never very striking, while two skins from Arizona are indistinguishable
from specimens of luci/ugus taken at Washington, D. C.

Sicull. — The skull of Myotis luci/ugus longicrus does not differ appre-
ciably in size or form fiom that of true luci/ugus.

Teeth. — In dental characters Myotis luci/ugus longicrus agrees closely
with typical 31. luci/ugus. While there appear to be no constant and
imj)ortant differences between the teeth of the two forms, the third
upper premolar averages slightly larger in longicrus, and there are
usually trilling differences in the relative sizes of the lower premolars
(fig. 14c).

Measurements. — See table, on page 05.

MYOTIS LUCIFUGUS LONGICRUS.

65

Specimens examined. — Total number 97, from the following localities:

Arizona: San Francisco Mt., 2; Cliiricaliua Mts., 1 (skiuj.

California: Nevada City, 4; Js'icasio, 72; Owens Lake, 1; Point Keyes, 1; San

Eniigdio, 1 ; Walker Pass, 2.
Chihnalma: San Francisco Water Canyon, San Luis Mts., 1.
Colorado: Grand Junction, 1.

Nevada: Cottonwood Range, 1 (skin); I'anaca, 1.
New Mexico: Santa Fe, 1.
Oregon: East base Cascade Mts., near Mt. Tbielson, 1 (skin); Beaverton, 2

(Miller Coll.).
Washington : Cape Flattery, 1 ; Colville, 1 ; Geyser Basin, 1 ; Fort Townsend, I.
Wyoming: Lake Fork, 1.

General remarks. — MyoUs Ivcifi<gvs longicrus is a well-marked geo-
graphical race of ,1/. lucifunn.s, replacing the typical form of the latter
ill tlie western United States throughout the region west of the Great
Plains. The northern and southern limits of its range can not at pres-
ent be determined.

A single skin from Cofre de Perote, Vera Cruz, is probably referable
to .1/. lucifiKjns longicrus. The tibia, however, is slightly shorter than
in true longierns, and more material from southern localities may show
the necessity of recognizing another geographic race.

.^fyotis lucifugns longicrus is the bat to which the 'melanic form of
Vespcrtilio albescens-^ of Dr. Allen's recent monograph for the most part
lefers. Under this name, however, Dr. Allen also included dark-
colored specimens of il/. californicus.

Measurements of subspecies of Myotis lucifugus.

Subspecies.

Ijricalit.y.

o aJ

si,

3 ft

.a
a

3

o

i

a
'3

o

o

a

£

o

.a
H

■-fl

u

a
o

il

$

o

3

S
H

lucifugus

4

89.5

40

16

8 7

38 8

fi ">

63 7

13 7

<» 4

7 1

District of Ciihinil)ia: Washing-

2??

86.5

37.5

9

38.9

6.7

12.2

9.4

7.5

ton.

Peiiiisylvauia : Centre County

6

85.3

:i7.6

15.5

7. 3 37. 6

5.6

60.2

13.3

9.7

7.2

10

4??

6

1?'

86.3
87.7

38.4
37 ''

16.5
15 5

8.7 38
9 36.7

6.6
6 1

61.5
61 '

13.2

9.1

8

Newfountllaml : liny St. Goorgo. . .

88.6
91

38.3
39

16.1
16

9. 1 36. 1
9 38

6.4

60

13.6
16

8.5
10.6

7

alancensin

Sitka

8

Sitka

8??

86.3

38.6

16

8. 4 36. 6

6.2

59

15.2

9.5

7.6

3
3

80
85.6

35
37.6

15
16.9

8
8.9

34.6
37

6.2
6.2

57.6
60

14.8
14.9

8.9
9.9

7

British Cnlunibia: Massett

7

lonijicriig

Washington: Puget Sound

1?

97

44

19

8

38.5

6

65

12

10

7

California : Nioasio

10

95.3

43.6

18.7

7.7

37.6

5.5

65.3

12

9.8

7.3

1?
1?

102

46

19 6

8

39 6

6

71

13

9.8

7.4

Owens Lake

96

45

17.8

8.4

6

69

12.6

9

7.4

Walker Pass

2??

97

44.6

18.5

7.4

39.6

6

69

12.6

10

7.4

19
Id-
ler

2

100
97

45
4'i

18.8
19

7.6
8

40

38

5

7

69
71

13.4
1'^.6

9.6
9

7.4

7.4

94
93.5

43

41

18.4
17.3

8
7.7

39.6
38.5

5.6

68
68

12
13.5

9
9.5

7.4

Arizona: San Francisco Mountain

8.2

2772— No. 13-

' Type.

66 NORTH AMERICAN FAUNA.

MYOTIS YUMANENSIS (H. Allen).

1864. }'ei<2)erlilio i/iimaiiensix II. AUoii, Mouogr. N. Am. Bats, p. 58.

1860. Vespertilio mavropns II. Allen, I'roc. Ac;i<l. Nat. Sci. Pliila., p. 2X8, (nee. Gould,

1854).
1893. I'espertilio nitidiis (ixdouiorjiliic variety) H. Allen, Monogr. Bats N. Am.,

pp. 72, 73.
1893. Vespertilio albescens II. Allen, IVIonoi^r. liats N. Am., p. S7, (part, ne(- fieofif.,

1805).
1893. Vespertilio nitidus macropiis H. Allen, Mouoyr. Bats N. Am., p. 100.

Ty2)€ locality. — Old Fort Yuma, California.

Geof/rapJiic (listrihution, — Austral zones and lower edge of Transi-
tion zone from the southwestern United States to San Luis Potosi and
Michoacan, Mexico.

General characters. — Size small; length 74-88; forearm 32-37 ; calcar
distinct, considerably longer than free border of interfemoral mem-
brane, terminating in a well-marked lobule; free border of uropatagium
naked; ears moderate; wings from base of toes, but on account of
extent of web l)etween toes api)arently from side of metatarsus; feet
very large and strong as compared with otlier small American species.

Uars. — The ears (PI. I, fig. 2) are moderately long; laid forward they
reach just beyond the tip of the nose. The anterior border is straight
for a short distance at base, then strongly convex, and finally straight or
even slightly concave just below tip. The tip is narrow and abruptly
rounded off. The posterior border is concave from the tip to the
widest ])art of the auricle, just below mid height, then strongly convex
to basal notch, which isolates a well-marked rounded lobe.

Tragus slender, acutely pointed. Anterior border slightly concave
at base, then straight or very faintly concave to tip. Posterior border
crenulate, straight or slightly concave from tip to broadest i)oint at
about lower third. A very large lobe at base; this lobe so large that
the greatest breadth of the tragus is often through it.

Membranes. — The membranes, especially the urojiatagium, are, for so
small a bat, thick and leatliery. The interfemoral membrane (PI. II,
fig. 2) is furred at the base, both dorsally and ventrally,but otherwise is
naked excei)t for a sprinkling of short hairs along the veins. Wings
from base of toes, but on account of the cousincuous webbing of
the latter the membrane appears to be attacihed to the side of the
metatarsus.

Feet. — The feet (PI. II, fig. 2) are, for so small a bat, very large, broad,
and strong, more than half as long as the short tibite, the whole leg
and foot suggesting a small NycUceiiis rather than a Myotis. Toes
(without claws) as long as sole, united by membrane at base to distal
fourth of proximal phalanges. Calcar strong and distinct, much longer
than free border of uropatagium, usually terminating in a distinct
lobule. Keel on posterior edge very slightly developed.

Fur and color. — The fur shows no peculiarities in distribution. On
the middle of the back it averages about 0 mm. in length.

MYOTIS YUMANENSIS. 67

Color i^ale wood biowii, varying to broccoli brown ; belly dirty whitish ;
the fur everywhere light plumbeous at base; ears and membrane very
light brown ; the uropatagium and wing membranes edged with whitish.

Specimens from the type locality are the palest that I have seen.
Those from Fort Yerde, Arizona, and apparently also alcoholic specimens
from Tulare and other localities in southern California, are slightly
(hirkcr, but still very different from M. yumanensiti saturatus.

SLidl. — The skull of Myotis yumani'n.si.s resembles that of 31. lucifufjus
in form, but is distinguished from the latter by its smaller size, and
shorter, broader palate. The brain case is broader and flatter than in
31. li(cifi((ii(.s. From the skull of 3r. colifornicns that of 3[. yiinKinensi.s
is readily distinguished by its slightly larger size and very much
broader, more robust form, the rostrum in i^articular being noticeably
broader.

Teeth. — The teeth of 31yotis yumancnsis (fig. 14 a) more closely resem-
ble those of ilf. Incifugus than any other species. They are, however,
smaller, and the crowns of the molars are longer in pro])<ntion to their
width. The crown of the third lower premolar is only slightly longer
than broad, thus resembling the corresjjouding tooth in 31, Incifugus,
and differing from 31. valiJhrnicKS, which, like 31. .snbuhdns, has the
crown of this tooth very distinctly longer than broad.

31e(i.surements. — See table, ])age 69.

Specimens examined. — Total number 142, from the following localities:

Arizona: Fort Verdi-, 6 (skins); White Monutains, 1 (skin, Am. Mus. Nat.

Hist.).
California: Fort Iveadiuy, 1; Fort Ynma, 5 (skins); Fresno, 8; Horse Shoe

Bend, Colorado Kivia-, 1; Keeler, 7; Lone Pine, 2; Mount Whitney, 1;

Nt'vada City, 2; Nicasio, 1; Owens Lake, 5; Owens Valley, 1; Old Fort

Tejon, 13; San Lnis Key, 8; Tulare, 45; Walker Pass, Kern County, 1.
Michoacau : Patzcuaro, 13.
Nevada: Pyramid Lnke, 1.

San Luis Poto.si : .Icsus Maria, 7; Hda. La Parada, 3; Ahualulco, 9.
Utah: Provo City, 1.

General remaHs. — Myotis yumanensis needs comparison with ilf. cali-
fornicvN only. From the latter it is readily distinguished by its much
larger foot and longer calcar. More detailed comparison of the two
will be found under 71/. calif oruicus.

This is the species to which Dr. Harrison Allen's recent account of
VespcfUlio albescens for the most part refers. His so-called larger
melanic form of albescens is 3Iyotis hicifuqus longicrus (True). Speci-
mens of 31yoiis californicus, 3f. tltysanodes, and 31. velifer in the Biologi-
cal Survey collection have been labeled by Dr. Allen 1". albescens, the
last two, however, with a query. This bat is ahso the Vesjiertilio
macropus and Y. nitidus macropus of Dr. Allen. The name macropns,
however, is preoccupied by Vespertilio macropus Gould, 1854.'

3[yotis yumanensis is a much smaller bat than 31. albescens, and does

1 Mammals of Australia {Jide Dobson).

68 NORTH AMERICAN FAUNA.

not agree with the descriptions of the latter given by GeofiFroy or Dob-
son. Mr. Oldfiekl Thomas, after ('X)m[)aring specimens of jI/^. yuni«nen-
sis with the M. albescens identified by Dobson in the British Museum,
writes me that the two are in no way closely related.

Lack of an ade(piate series of skins prevents any determination of
the extent of individual color variation in this species. Specimens
taken at Fort Verde, Arizona, in May and August, are slightly darker
than those killed at the type locality in April, while two July skins
from Tatzcuaro, Miclioacan, are nearly as dark as the lightest examijles
of M. yumanensis satnratus.

MYOTIS YUMANENSIS SATURATUS subsp. uov.

Type from Hamilton, Wasliingtou. Adult ^ (skin), No. iJloi'? U- S. National Mnseum
(Biological Survey collection). Collected September 13, 1S89, by T. S. Palmer.
Collector's number, 392.

Geographic distribution. — Transition zone in Oregon, Washington,
and British Columbia.

General characters. — Similar to typi(;al Myotis ymnanensis, but fur
longer and color much darker.

Fxr and color. — Fur distributed as in the typical subspecies. On the
middle of the back it averages about D mm. in length. Back dark glossy
yellowish brown (the exa(;t shade usually between the 'sepia' and
'mummy brown' of liidgway's Nomenclature of Colors, PI. Ill); belly
isabella color; chin, throat, and sides darker than belly; fur every,
where deep blackish plumbeous at base; ears and membranes blackish.

Measurements. — See table, page 01).

Specimens examined. — Total number 19, from the following localities:

British Columbia: Kamloops, 1 (skin) ; Kultus Lake (near Chilliwack), 1 (skin,
Miller coll.); Mount Lehman, 1 (skin, Aiu. Mus. Nat. Hist.); Port Moody, 1
(skin); Shuswap, 1 (skin); Sumas, 3 (skins. Miller coll.).

Oregon : Crooked River, 1' ; Lone Rock, 2' ; Twelve Mile Creek, 1 '.

Washington: Chelan, 3'; Hamilton, 2 (skins); Lake Cushman, 1 (skin) ; Neah
Bay, 1.

General remarlcs. — Myotis ynmanensis saturatus is readily distinguish-
able from true ynmanensis by its much darker color. In this character,
however, it closely api)roaches the typical form of ]\[. lncifugns,fvom
whi(;h it differs chiefly in smaller general size and much smaller skull.
From j1/. lucifugus longicrus it differs very noticeably in its shorter tibia
as well as in other characters.

'These specimens may be nearer true yitmanethtis than safin-ntns. In alcohol, liow-
ever, their color appears much too dark for the typical subspecies.

MYOTIS CALIF0RNICU8.
Measurements of subspecies of Myotis jiududkhsis.

69

Subspecies.

Locality.

H

yumanennu

California : Fort Yuma

Tulare

Arizona : Fort Yerilc

San Luis Potoai: Jesus Maria

Miclioacan : Patzcuaro

British Columbia: Sumas

Washington: Chelan

Hamilton

4
10
4

5??
8
3
3

83. 7 35. 5
80. 5 36. 6

79.2
81

80.6
82.3

77
77

36.7

34.9

36.3

36.5

35

30

14.1
15

15.7

15.5

14.6

14

15

15

8.7|32.7. 5.2
8.734.2' 6
8.8 33.9..
7.934.4J 6.5
8. 3 34. 1' 5. 7
8.6 34.6J 5.3
8. 3 35. 3! 6
8.6 33 [5

56.5. ...I...
56. 4 14. 2 9
55. 5

57.8
7
57
59.3

14. 4 '.). 1
14.2 9

14.3 8.9

7.6
7.6

' Type.
MYOTIS CALIFORNICU8 (Aud. & Hach.).

1842. Vespiriilio californicux Ami. & Bach.. .lomn. Acad. Nat. Sci. Pliila., p. 280

(California).
1862. VesprrWio nitidu^ H. Allen, Proc. Acad. Nat. Sci. Pbila., p. 247 (Monterey,

California).
1864. Vesperlilio nitidiis H. Allen, Mono.i;r. N. Am. liats, p. 60.
1864. Vespertilio oregonensis H. Allen, Monogr. N.Am. Bata, p. 61 (Cape St. Lucas and

Fort Tejon).
1866. Vesperlilio ohscurua H. Allen, Proc. Acad. Nat. Sci. Phila., p. 281 (Lower

California).
1866. Vespcrfilio volans H. Allen, Proc. Acad. Nat. Sci. Phila., p. 282 (Cape St. Lucas).
1866. VespertUio exilis H. Allen, Proc. Acad. Nat. Sci. Phila., p. 283 (Cape St. Lucas).

I'espert'dio tenuidorsalis H.Allen, Proc. Acad. Nat. Sci. Phila., p. 283 (Cape St.

Lucas).
Vesperlilio yumanensis H. Allen, Proc. Acad. Nat. Sci. Phila., p. 283 (nee H. Allen,

1864).
VespertUio nifidus Dobson, Catal. Chiroptera Brit. Mns., p. 318.
Vesperlilio melanorhintis Merriam, North American Fauna, No. 3, ]>. 46, Sept. 11,

1890 (San Francisco Mt., Arizona).
1893. Vespertilio albescens melaiiorhivus H. Allen, Monogr. Bats N. Am., p. 91.
1893. Vespertilio nifidus H. Allen, Monogr. Bat.s N. Am., p. 94.
1893. Vespertilio nitidus henshawi H. Allen, Monogr. Bats N. Am., p. 103 (Wingate, N.

Mexico).
1893. Vesperlilio niurieans H. Allen, Monogr. Bats N. Am., p. 97 footnote (nee Maxi-
milian 1826).

Type locality — <■ California.'

Geographic distribution. — Austral zones and lower part of Transition
zone throughout the western United States and Lower California, east
to Wyoming and Texas. South limit of range not known.

General characters. — Smallest species of Myotis known to occur in the
United States. Length, 70 to 87; forearm, 30 to 36. Calcar about as
long as free border of uropatagium, very slender but distinct and with
a more or less well developed lobule at tip, outer edge with a distinct
keel. Legs slender, the small feet reaching when extended backward
to within about 5 mm. of tip of tail. Free border of uropatagium naked.

1866.

1866.

1878
1890

70 NORTH AMERICAN FAUNA.

Ears moderate, reaching just beyoud tip of nose. Wings from base of
toes. Fur on back distinctly darker at base than at tip.

Ears. — The ears are moderately long (Pi. I, fig. 2), reaching when laid
forward 1 to 3 mm. beyond tip of nose. The anterior border of the
auricle is straight or slightly convex at the base, then strongly convex
to a i)oint somewhat beyond the middle, after which it is straight or
even a little concave to the narrowly rounded otf tip. Posterior border
concave from tip to point slightly below the middle, after which it is
convex to basal notch. Basal lobe strongly developed and notchetl on
its lower border.

Tragus varying much in shape, but with anterior border usually
straight, or nearly so, and posterior border strongly convex and with
small basal lobe.

Membranes. — The membranes are thin and delicate. Uropatagium
(PI. II, tig. 1) furred on basal third, otherwise naked, except for a few
hairs along the veins. Wings from base of toes, naked, except for a
narrow strip along side of body.

Feet. — The feet are small and weak (PI. II, fig. 1), distinctly less than
half as long as tibia. Calcar slender but distinct, shorter than free
border of uropatagium, usually terminating in a distinct lobule. The
posterior border is provided with a keel beginning abruptly about 2
mm. from the base and fading away gradually at about middle of calcar.
This keel is supported by 1 to 3 cartilaginous outgrowths from the
calcar.

Fur and color. — The fnr is soft, full, and long, that on middle of back
averaging about 8 mm. in length.

Color light yellowish gray, paler on the belly, the fur everywhere
dark plumbeous at base. Membranes, ears, lips, and muzzle blackish.

ShttU. — The skull of Myotis californicns is smaller and more lightly
built than that of any other North American Myotis. The brain case
is moderately rounded, and the long narrow muzzle fades gradually
into the gently sloping forehead. The skull is thus very different from
tlvAt of M. yumanensis J the only species with which M. californicns is
likely to be confused. In form it resembles the skulls of M. evotis
and M. thysanodes, but the latter are among the largest of the species
found in the region inhabited by 31. californiciis.

Teeth. — The teeth of Myotis caUfornieus (fig. 15, a) are, like the skull,
small and delicate. In general they closely resemble the teeth of ]\f.
suhvlatiis, and differ from those of M. yumancnsis in numennis details,
as in the shape of the third upper molar and tliird lower premolar, the
former being distinctly narrower and the latter longer in proportion to
its width than in M. yumanensis.

Measurements. — See table, page 74.

Specimens examined. — Total number 152, from the following localities:

Arizona: Camp Grant, 2; Oracle, 5; Prescott, 1 (skiu. Am. Mua. Nat. Hist.);
SaataCat.iliua Mountains, 1 (skiu); Tinujas Altas, Yuuia County, 3; White
Mountains, 1 (skin, Am. Mas. Nat. Hist.).

MYOTIS CALIFORNICUS.

71

Califoruiii : Amargosa River, Moliave Desert, 1; Banning, 1; Calito, 1; Colo-
rado Desert, 1 ; Death Valley, 14 (including Bennett Wells, 1 ; Funeral
Mountains, 1; Saratoga! Springs, 6); Dulzura, 15 (6 skins, Miller coll.); East
Fork Kaweah River, 3; Fort Crook, 1; Fort Tejon, 11; Old Fort Yuma, 1;
Jacumba, San Diego County, 1; Kern River (25 miles above Kernville), 1;
Monterey,!; Mount Shasta, 1; Nicasio, 7; Petaluma, 1 ; Point Reyes, 5; San
Clemeute Island, 3; Santa Barbara, 1; Santa Ysabel, San Diego County, 23;
Tfjon Pass, 1; Tres Pinos, 1 ; Twin Oaks, San Diego County, 1; Witch Creek,
San Diego County, 7.

Chihuahua: East side of San Luis Mountains, 3.

Lower California: Cape St. Lucas, 2 (' T. (A)8curu>i^ H. Allen) ; San Fernando, 3
(Miller coll.).

Nevada: Colorado River, 2; Cottonwood Range, 4 ; Gold Mountain, Esmeralda
County, 2; Pahrump Valley, 1; Panaca, Lincoln County, 1; Vegas Valley,
Lincoln County, 1.

New Mexico: Fort Defiance, 1; Fort Wingate, 2; Silver Citj' 1 (skin).

Oregon: Elgin, 1; John Day River, 3; Twelve Mile Creek, 2.

Texas: Paisano, 1.

Washington: Almota, Whitman County, 1; Blue Creek, 1; Chelan, 1.

Wyoming: Bitter Creek, Sweetwater County, 2 (skins. Am. Mus. Nat. Hist.);
isullLake, 1.

^^^^A^S^^msgs^J^

Flo. 1.5.— Teetli of (a) Myotit califdrnicu.i, (h) M. siibvlahts, (c) if. evoflfi, ami (</) M. thysanodefi ( X 5.)

General remarls. — Typical Myotis californicus varie.s considerably in
<'-ol()r, size, and proportions, bnt may always be recognized among-
North American and Mexican species by its small size, slender form,
delicate membranes, long tail and legs, small feet, and pale yellowish
color. il/?/o^/.s'if/?f»j^nje«.s/.s',theonlyotlier species of eqnally small size, has
conspicuously shorter legs, larger feet, shorter tad, and thicker mem

72 NORTH AMERICAN FAUNA.

braiies. The species resembling M. caUfornicns in form are all conspic-
uously larger.

In his recent monograph Dr. Harrison Allen uses the name Vesper-
tilio nitidus for Myotis ealifornicns. As subspecies of californicits he
includes M. yumanensis (=' Yespertilio nitiflus maeropu-s^), M. ealifor-
nicns ciliolabrum, and M. lucifiu/us longieriis {=^ Vespertilio nititlus
longicrus^). Vespertilio melaHorhinns,n synonym of 71/. ealifornicns, he,
however, refers to ' V. albescens^ {=]\[. yumanensis) as a subspecies,
* Vespertilio albescens melanorhinusJ

MYOTIS CALIFORNICU8 CAURINUS snbsp. nov.

Type from Masaett, Qiieeu Charlotte Islands, liritisli Columbia. Atliilt, ^ (in alco-
hol), No. 72219., U. S. National Museum (P.iological Survey collection). Collected
iu 1895 by .J. H. Keeu.

Geographic distribution. — The humid coast district of British Colum-
bia, Washington, and Oregon (possibly ;ilso of northern California).

General characters. — Similar to typical M. c«i(/brnicws, but very much
darker in color.

Ears, membranes, feet, andfnr. — As in typical calif ornicus.

Color. — Very deep, frequently almost blackish sepia throughout,
slightly yellowish on belly, the fur everywhere blackish plumbeous at
base.

Measuremeiits. — iSee table, page 74.

Specimens examined. — Total number, 14, from the following localities:

British Columbia: Port Moody, 1; Massett, 9.

Oregon: Marmot, 1 (skin).

Washington: Fort Steilacoom, 1; Pnget Sound, I; Tenino, 1.

General remarks. — In color Myotis ealifornicns caurimis closely re-
sembles dark specimens of }[. californicus mexicanns. It is readily
distingnishable from the latter, however, by its much smaller size.

MYOTIS CALIFORNICUS CILIOLABRUM (Merriam).

1886. Vespertilio ciliolahrum Merriam, Proc. Biol. Soc. Washington, IV, p. 1.
1893. Vespertilio nitidus ciliohthnim H. Allen, Monogr. Bats N. Am., p. 101 (i)art).

Type locality. — Trego County, Kansas.

Geographic distribution. — Trego County, Kansas, and central South
Dakota. Limits of range wholly unknown.

General remarks. — Similar to typical M. californicus, but very much
paler in color.

Uars. — In form the ears of Myotis californicus ciliolabrum are as in
typical M. ealifornicns. They average, however, slightly larger.

Membranes. — The membranes are thin and translucent. Wings from
base of toes, and entirely naked except a narrow line close to the body.
Uro])atagium thinly haired on proximal fifth ventrally and on proximal
half dorsally, otherwise naked, but with a few hairs along the veins.

Feet. — The feet are moderately large, about half the length of the tibia,

MYOTIS CALIFORNICUS MEXICANUS. 73

tlie toes longer tlian the sole. A distinct wart at lieel. Calcar slender,
distinct, about e(|ual to freeborder of interfenioral membrane, terminat-
ing in a small but distinct lobule and noticeably keeled along posterior
edge. The keel is supported by 1 to 3 cartilaginous processes.

Fur and color. — Except for its unusual extension on the back of the
uropatagium the fur shows no peculiarities in distribution.

In color tlie fur is ]>ale yellowish white throughout. Tlie ears, muzzle,
and chin are dark brown in strong contrast. Membranes light brown
with pale edges.

Measurements. — See tal)le, page 74.

Specimens e.r a mined. — Total number L">, from the following localities:

Knnsas: Trego County, <i.

South Dakota: Carroll Draw, I'ine Ridnc Indian Keservation. 7 (skins Am.
Mus. Nat. Hist.).
(JeneraJ remarks. — Myotis californicns ciUoJahrnm is a i)ale, whitish,
race of M. cali/ornicit.s, presenting the opposite extreme from M.c. can-
rinus. Except in coh)r, 1 can not find that it differs in any constant
characters from typical californ tens. The specimens from Gran t Conn ty ,
N. Mex., referred to in the original description of j\I. c. ciliolahrum are
undoubtedly true californicns, as are those from Death Valley referred
to this subspecies by Dr. Harrison Allen.

MYOTIS CALIFORNICUS MEXICANUS (Saussnre).

18(j0. Vcspertilio mexicatnis Saussnre, Revue et Mag. de Zool., 2e ser., XII, p. 2S2.
1X66. ]'espertino (iffiiis H. Allen, Proc. Acad. Nat. Sci. Pliila., p. 282 (Mirador, Max.).

Type locality. — Unknown, but probably Vera Cruz, Puebla, or
Oaxaca.

(ieographic distribution. — Austral and Transition zones in central
and southern Mexico (San Ijuis Potosi, Michoacan, and Oaxaca).
Limits of range not known.

(ieneral characters. — Slightly larger than typical Myotis californ inis,
and averaging somewhat darker and yelh)wer in color.

Ears, membranes, feet, and fur. — As in typical californicns.

Co/or.— Dull yellowish brown, slightly i)aler on the belly. IMem-
branes and ears in dry skins blackish. Two immature specimens fiom
Reyes, Oaxaca, are considerably darker than any of the adults, but other
wise a series of thirteen skins sliows very little individual variation.

Skull and teeth. — As in typical californicns.

Measurements. — See table, page 74.

Specimens examined. — Total number 51, from tlie following localities:

Michoacan: Patzcnaro, 44 (8 skins).
Oaxaca: Cuicatlan, 1; Reyes, 5 (.skins).
San Luis Potosi: Hacienda La Parada, 1.

74 NORTH AMERICAN FAUNA.

Average measurements of subspecies of Myotis californicus.

Subspecies.

californicus.. .

caunnus.

ciliolahrym .

inexicamis.

Locality.

California: Saratoga Springs

Death Valley

Arizona : Oracle

British Columbia: Massett..
Ma.ssett. .

Kan.sas : Trego County

Trego County

Miclioacan : Patzcuaro

■oQ

P p.

ScTcT
5

Id'
9

1?'

5

10

80

77.8

82

75

77

75

80.2

81.5

> Si

H

39

38.8

39

34

34.5

34

36.';

38.:

14.2

13.9

14.4

14

13.8

l.j

14 G

5.4 31.3 4
5.7 31 I 4.1

4.2 5.9

32 I 4.

32 5
32. 1 4. 3

33 I 6
32.8 5.4
34.1 4.4

a

0

a

a

a

0

p s

p

0

58

14.2

9.9

55.4

12.9

9.9

57

14

10

54

12.8

8

54.1

13.2

Q.r>

59

14

9

57

14.6

10

60

14

9.6

8
8.3

7.1
8.«

' Typo.

MYOTIS NIGRICANS (Maximilian).

1826. VesperHlio nigricans ''Schinz, Tliierreicli 11. s. w., B. J, p. 170" Maximilian,

Beitriige 7,ur Naturgesch. v. Biasilien, II, p. 266.
1878. Vespcrtdio nigricans Dobson, Catal. Cliiroptera Brit. Mns., p. 310.
1893. VespertiJio nigricans H. Alleu, Monogr. Bats N. Am., p. 06.

Type locality. — Fnzenda de Aga, near the Iritiba River, southeastern
Brazil.

Geof/rapliic distrlhution. — Tropical America, north to extreme south-
ern Mexico (Chiapas). Limits of range not known.

General ehararters. — About the size of typical Myotis calif ormcus, but
with slightly larger foot and smaller ears; fur on back not distinctly
darker at base than at tip.

Ettr.s. — The ears are slightly smaller than in 3T. californicus, but not
different in form.

Membranes. — As in 71/. californicus.

Feet. — The feet are relatively larger than in M. californicus, but smaller
than in ]\[. yumancnsis. Calcar about as long as free border of uropa-
tagiura, terminating in a small but distinct lob6; keel obsolete.

Fur and color. — Fur short, that on middle of back averaging a little
less than 0 mm. in length, nearly unicolor on back but distinctly bicolor
on belly. Back clove brown (lighter than No. 2 on PI. Ill of Ridgway's
Nomenclature of Colors), the hairs just perceptibly darker at base and
with glossy tips, which in certain lights produce a slightly grizzled
appearance. Belly light broccoli brown, the basal half of the hairs
deep plumbeous. Ears and membranes blackish in dry skins. A series
of ten skins from Iluehuetan, Chiapas, shows no variation in color,

Skull. — The skull of Myotis niyricans (figs, lie and V2a) is slightly
smaller than that of M. californicus, and has a shorter rostrum and
less frontal concavity in the dorsal outline. The differences are slight,
but very evident when series are compared.

Teeth. — The teeth of Myotis nigricans do not differ appreciably from
those of M. californicus.

MYOTIS SUBULATUS.

75

Measurements.— TliQ average measurements of 10 specimens of Myotis
igricans from Huehuetau, Chiapas are given iu the following table:

Arerage measurements of 10 sjfecimeus of Myotis nigricans.

o a

.4

o

fl

o

!S

a

bo

-^

«

a

a>

Locality.

■ 2
3 3

s

^5

C3

4J

g

g

.2 3

o

3

^^

H

H

H

o
6.9

O

H

o
1-1

^

Iiiapas, Huehuetan

10??

76.7

35

1.3.6

33.5

4.4

57

12.7

8.6

6.6

Specimens examined. — Total number, 34; all from Huehuetan, Chiapas
iltitude about 300 feet).

General re)nark-s. — Jiryotis nigricans differs from JA, ealifornicus in
nmerous characters, each of which is trivial in it<elf but which with
he others goes to mahe up a sum quite different from that shown by
ny other ^orth American bat. The constancy in color of tlie 10 skins
y which this species is represented in the Biological Survey collection
^ very remarkable.

MYOTIS SUBULATUS (S.iy). Say's Bat.

■^23. ? Vespertilio siibulaius Say, Long's Exped. to Rocky Mts., II, p. 65, footnote

(Arkansas River, near La Junta, Colorado).
•<64. T espertilio subnlatus H. Allen, Monogr. N. Am. Bats, ]>. 51.
■!7S. Vespertilio subnlatus Dobson, Catal. Cliiroj^tcra Brit. Mus., p. 324.
■!93. VespertiJio griiphns var. (b). Northern fonii of Vespertilio gryphns, H. Allen,

Monogr. Bats N. Am., p. 80.
^97. Vespertilio gryphns var. sepfenlrionalis Tronessart, Catal. Mamni. t. \'ivent. q.

Fi>8s., p. 131. (Only name undoulitedly based on this animal.)

Type l(>c<(Jity. — Arkansas liiver, near La Junta, Colorado.

Geographical (listributioii. — North America east of the Rocky Moun-
ains.

General characters. — Size medium; length SO to 90; forearm 34 to
7. (Jalcar slender, slightly btiiger than fre<' border of uropatagium,
Piniinating indistinctly or with a slightly developed lobule; keel rudi-
lentary or absent. Free border of uropatagium naked. Ears long,
t-aching 2 to 5 mm. beyond tip of nose. • Wings from base of toes.

Ears. — The ears (fig. 13 «) are long and slender, reaching when laid
()rward,2 to 5 mm. beyond tip of nose. Anterior border straight fiom
•ase to near middle, then for a varying distance moderately convex^
inally straigiit to narrowly rounded off tip. I'osterior border concave
loin point immediately below tip to about middle, where it becomes
onvex and continues so to basal notch. Basal notch strongly marked,
solating a narrow and very conspicuous lobe.

Tragus slender, straight, or slightly bent backward. Anterior bor-
ler straight throughout or slightly convex near tip. Posterior border
!traight or evenly and slightly concave from tip to widest i)oint, which
s opposite or slightly above level of anterior base. Basal lobe small.

76 NORTH AMERICAN FAUNA.

width of tragus throiigli lobe always iinicli less than width at base o
anterior edge.

Membranes. — The membranes are thin and translucent, naked except
for a narrow line close to the body. On the uropatagium the furrec
region occupies the basal fourth dorsally, rather less ventrally, other
wise the membrane is naked except for scattered hairs along the veins
Wings from base of toes.

Feet. — The foot is moderately large, about half as long as tibia. Toe;
longer than sole, united by membrane at base to a little beyond niiddl*
of proximal phalanges, and sprinkled with coarse hairs on dorsal sur
face. Calcar slender, equal to or slightly longer than free edge of inter
femoral membrane, terminating indistinctly or with an ill-detiued lobe
Keel rudimentary or absent.

Fur and color. — The fur is full and soft, but shows no peculiaritie
in distribution. In color it apparently does not differ from typical M
lucifugns; but too few skins are now available to determine the limit
of variation.

*9/.w?/.— The skull of Myotis subulaUis resembles that of Af. evotia s
closely that it is impossible to distinguish with certainty between th
two. In M. suhidatiis the skull is very slightly smaller, but the diffei
ence is trilling and intangible. The skull of M. snbuhttHS does nc
closely resemble that of M. Incifngus.

Teeth. — The teeth of Myotis subulatus (fig. 15 ^>) agree in form aii
relative size with those of M. evotis, and I am unable to find any difi'ei
ences by which to separate them. They differ, however, in man
details from the teeth of A[. lucifufins.

Measurements. — See table on page 77.

Specimens examined. — Total number 53, from the following localities

Alberta: Near Red Deer, 1 (skin, Miller coll.).

Illinois: Cliii-ano, 1.

Indiana: Brookville, 1; Wheatland,!.

Kentucky: P]ubanks, 2.

Maine: Eastport, 2.

Maryland: Forest Glen, Montgomery Connty, 2 (Miller coll.).

Massachnsetts : Woods Hole, 1.

Minnesota: Elk River, 2.

Missouri: Marble Cave, 9.

New York: Hanimondville, 12 (Merriam coll.); Hemlock Lake, 1; Hijjblan,

Falls, 1; Lake George, 3; Peterboro, 2 (Miller coll.).
Ontario: Mount Forest, 1 (skin. Miller coll.); North Bay, 1 (Miller coll.).
Pennsylvania: Meadville, 1.

Quebec: Godbont, 1 (Merriam coll.); Ottawa, 3 (Merriam coll.).
Tennessee: Bellamys Cave, 1.
Virginia: Alexandria, 1.

West Virginia: Aurora, 2 (Merriam coll.). '

Wisconsin: Bayfield, 1.

General remarls. — Myotis snbnlafns may be distinguished from M
lucifugns, the only s])ecies with which it is likely to be confused, by it]
narrower skull, longer ears, and longer, more sharply pointed tragus

MYOTIS EVOTIS.

MYOTIS SUBULATUS KEENII (Meniaiu).

77

1895. respertilio nubulatus keeiiii Merriaiu, Americau Naturalist, XXIX, p. 860, 'Sep-
tember 1, 18115.

Type locality. — Massett, Queen Charlotte Islands, Britisli Columbia.
Type in U. B. National Museum (Biological Survey collection). Adult
9, No. 7i'922 (in alcohol).

Geographic ditstribution. — Myofis suhiilatiis Ixcenii is at present known
Ironi the type locality only. It doubtless occurs throughout most of
:he humid northwest coast district.

General characters. — About the size of typical Myotis suhulatus, but
vvith longer tail and ears; color much darker than in true .subnlatus.

Uars. — The ears of ilf. stihiilatu.s licenii (fig. 13 h) average distinctly
oiiger than those of typical suhulatus from the eastern United States,
jut do not difl'er in form.

Fur and color. — The fur appears to be longer than in true srbulatus,
lud considerably darker in color, but with alcoholic specimens only for
■oniparison it is impossible to determine the degree of difference
jetween the two forms. IVIembranes and ears blackish.

Measurements. — See table below.

Specimens examined. — Total number o, all from the type locality.

General remarks. — Myotis suhulatus Iceenii is a well-marked race, char-
icterized, like the other bats of the humid northwest coast district, by
larkuess of color. In addition to its color differences it has longer ears
han its eastern lepreseutative, in this respect showing much the same
variation as M. lucifugus alascensis.

MeasuremcntH of suhspecies of Myotis suhtilaiiis.

Subspecies.

■mhulatus .

keenii.

Locality.

New Tiirli : Haniniomlville . . .

Quebec : (iodbout

Mi.ssouri : Marble Cave

IJritisb C'olumbia: Mas.sett ..
Massett ..

o ^

rd

ei

a

3

a
u 2
o s

II

o

H

il
>

'a

3

o
o

i

s

13
.a

H

■■a

a
o

61

16.3

o
10.2

10

85.6

38.8

17.2

7.5

35.7

6.3

'■idd

80

36.8

16.4

7. 5 33. 8

6.5

58

15.6

0.8

8

85.3

.38.5

17.1

8 35.3

6.2

61

16.7

10.7

!?■

86

41

16.4

8.8 36

6

60

17.4

11

3

84.3

40.6

16.3

8. 3 35. 3

6

58

17.8

10.8

9.6
9.8

1 Type.

MYOTIS EVOTIS (H. Alleu). Long-eared Bat.

1H64. VespcrHlio evotis H. Allen, Monogr. Bats N. Am., p. 48.
1878. VespcrtUio erotis Dobsou, Catal. Cliiroptcra Brit. Mus., p. 324.
1893. VeapertUio albescens eroiis H. Alleu, Monogr. N. Am. Bats, p. 89.
1896. VesperHUo chrysonotus J. A. Allen, Bull. Am. Mus. Nat. Hist., VIII, p. 240, No-
vember 21, 1896. Kinney Ranch, Sweetwater County, Wyoming.

Type locality. — Not stated, and no type designated. In the original
description specimens are mentioned from the upper Missouri Eiver,

78 NORTH AMERICAN FAUNA.

and the Pacific coast from Paget Sound to Cape St. Lucas. Monterey,
Cal. (one of the localities given), may be selected as the type locality.

Geographic (list rihnt ion. — Austral and Transition zones from the
Pacific Coast to the eastern edge of the Rocky Mountains; south to
Vera Cruz.

General characters. — Size large; length 85 to 02; forearm 3() to 43.
Calcar longer than free border of uropatagium, slender, distinct, and
with a more or less well-develo[)ed lobule at the tip. Free border of
uroijatagium naked or very indistinctly ciliate. Ears very long, reach-
ing 7 to 10 mm. bej^ond tip of nose. Wing from base of toes.

Ears. — The ears (PL 1, tig. 0) are long and slender; laid forward they
reach considerably (7 to 10 mm.) beyond tip of nose. Anterior l)order
of auricle regularly convex from base to a point slightly beyond mid-
dle, thence straight or nearly so to the tip. Posterior border slightly
concave immediately below tip of ear, then gradually and moderately
convex to base, liasal lobe strongly developed, and notched on the
lower border. The auricle is usually marked with three or four distinct
cross ridges.

Tragus long, slender, and iwinted. The anterior border straight or
slightly concave from base to about mid height, then moderately con-
vex, the terminal third or fourth usually straight. Posterior border
with a small but distinct lobe at base. Above this lobe the nmrgiu
bends abruptly outward for a varying distance, sometimes forming a
sharp and conspicuous angle with the lower end of the concavity which
extends downward from the tip of the tragus, in other cases sei)arated
from the latter by a region of varying extent in which the posterior and
anterior borders are parallel. These variations bring about striking
contrasts in the form of the lower part of the tragus in different indi-
viduals, and suggest the existence of more than one species or race.
Si)ecimens from approximately the same region, however, show both
extremes and intermediate conditions.

Membranes. — The membranes are thin and light. Uropatagium hairy
on basal fifth, otherwise naked except for a few hairs along the nerves
and on the free border. Wing from base of toes (PI. II, figs. 3 and 4)-

Feet. — The feet are moderately large, slightly less than half as long
as tibia\ Toes (without claws) distinctly longer than sole and united
by membrane through basal third of proximal phalanges. Whole dor-
sal surface of foot sprinkled with stiff hairs. Calcar distinct, ecpial to
or longer than free border of uropatagium, terminating in a lobule of
varying distinctness. Posterior border never distinctly keeled.

Fur and color. — The fur is full, soft, and not peculiar in distribution.
It is light yellowish brown, paler ventrally, the hairs everywhere dusky
slate at base. The absence of a series of skins of this bat makes it
impossible to describe the color accurately or compare it in detail with
that of its allies, M. thysanodes and M. subulatus. A skin from Shuswap,
British Columbia, has the fur of the back dull, pale raw umber, the

MYOTIS EVOTLS.

79

dusky bases of the hairs showing through along the sides. The belly-
is hght broccoli brown. In front of tlie shoulder and Just below it is
a small tawny olive area which contrasts strongly with the color of the
belly. In another specimen (No. 1382, collection of Dr. C. Hart Mer-
riaiu, San Bernardino Mountains, California, August 14, 1885, F.
Stephens) the color is similar but a shade paler and yellower through-
out, the dark bases of the hairs nowhere showing through. No dark
shade in front of shoulder. This specimen is i»ractically indistingnish-
able in color from the palest examples of M. titysanodes, but the fur is
nmch darker at base and the general color is slightly clearer yellow,
with the tips of the hairs more glossy. Other skins are duller and
less yellow.

Skull. — The skull of Myotis ecotis equals that of 71/. thysanodes in
length and mastoid breadth, but is narrower across zygomata and has
the occiput less elevated. The occipital outline is rounded as in M.
thysanodes, and the occipital crest is very sliglitly developed. Fore-
head rising above the muzzle gradually, in tliis respect also resem-
bling M. thysanodes. Rostrum more sk'nder than in M. velifer or M.
thysanodes. Pterygoids and posterior part of i)a]atines as in 71/.
thysanodes.

The skull of Myotis evotis is easily distinguished from that of all other
North American species except 71/. suhulatus. From the latter, how-
ever, it ditfers merely in very slightly larger size.

Teeth. — In dental characters Myotis evotis does not differ essentially
from 71/. thysanodes. The premolars apparently show less tendency to
crowding, but I can find no tangible difference in form or relative size
(fig. 15 c).

Measurements. — The measurements of 8 specimens of Myotis evotis
from eight localities are given in the following table:

Measurements

of S specimens of Myo

is evotis from 8 localit

ies.

S

S

a

t3

fee

^

■-a

a

©

Locality.

a

?

c4

a

a

4^ t^
11 (D

c s

3

o

o

CS

^

m

H

H

H

f^

Ph

H

I-]

W

F=

H

Washington: Easton

cT

85

41

19
18

8
8.6

36
40

7
7

62
64

19.4

11.8

10.8

Wyoming: Kinney Ranch

California: Inyo Mountains ...

89

43

20

7

38

8

63

23

14.6

13

Owens Lake

?

91

43

17.6

8

38

6.4

63

22

13.4

12.4

San Joaquin Kiver.

cf

92

43

19

8

38

6

62

22

13.6

12

Twin Oaks

9

90
85

42
40

18.6
18

8
7.6

37
36.6

6.4

7

62
62

21
22

13
14.6

11

Nevada : Pahranagat Valley . . .

12

Vera Cruz : Perote

cT

91

42

20

9

40.4

6

67

20

12

11

'Tj

rpe of

Vesper

tllio ch

rysonc

itus J.

A. All

en.

80 NORTH AMERICAN FAUNA.

Specimens examined. — Total number 3li, from the followiug localities:

Arizona: Chiricahna Monutains, 1 (skin, Am. Mus. Nat. Hist.); San Francisco
Mountain, 1; Si)ringervilli', 2 (skins); Whitt; Mountains, 2 (skins. Am. Mu8.
Nat. fiist. ).

British Columbia: Shuswap, 1 (skin).

California: Dulzura, 3 (2 skins. Miller coll., 1, Am. Mus. Nat. Hist.); Inyo
Mountains, 1; Owens Lake, 1; San licrnardino Mountains, 1 (skin. Mcrriam
coll.); North Fork San Joaquin Kivor, 1; Twin Oaks, San Diego County,!.

Chi-hnahua: San Lnis Mts., 1 (skin).

Colorado: Loveland, 4 (2 skins, Miller coll.).

Montana: Hot Springs, 1.

Nevada: Cottonwood K'ange, 1; Paliranagat Valley, 1.

New Mexico: Vermejo River, 1 (skin).

Oregon: Blue Creek, 1; Harney, 1; Twelve Mile Creek, 1.

Vera Cruz: Perote, 1.

Washington: Easton, 1; Colville, 1.

Wyoming: Bull Lake (east of Fremont Peak), 1; Kinney Ranch, Sweetwater
County, 1 (skin. Am. Mus, Nat. Hist., type of F. rliri/aonotns J. A. Allen).

General remarlis. — Myotis evotis is so totally distinct from all other
bats occurring- in Mexico or the United States tliat no detailed com-
parison with any is needed. The only species Avith which it could be
contused are M. thysanodes and M. sKhuIaiti.s. Tlie ears, however, are
much larger than m either of these, and the free border of the uropata-
giun) is never densely haired, as in M. tliysanodes.

Through the kindness of I)i'. J. A. Allen, I have before me the type
of YespcrtUio ehrysonoius from Kinney IJanch, Wyoming. I am unable
to find that it differs in any way from Myotis erotis. The tail is muti-
lated so that it gives no characters. The forearm is only 2 mm. longe
than in the largest evotis from the United States that I have seen, a
difference too trivial to be taken into account. In color the type of
chrysonotus is a barely perceptible shade yellower than skins of evotis
from the San Bernardino Mountains, California, and Vermejo IJiver,
New Mexico, but the difference is wholly inconsequential.

MYOTIS THYSANODES sp. nov. Fringed Bat.

1893. VespertUio albescens relifer (variety) H. Allen, Monogr. Bats N. Am., p. 93. Dul-
zura, California.
1893. VespertUio albescens erotis H. Allen, Monogr. Bats N. Am., p. 90 (part, specimen

No. 29827, from old Fort Tejon, California).
Type from Old Fort Tejon, California. Adult 9 (in alcohol). No. 29827, U. S.
National Museum (Biological Survey collection). C(dlectod July 5, 1891, by T. S.
Palmer. Original number, 235.

Geographic distribution. — Lower Sonoran zone from near the southern
border of the Western United States to San Luis Potosi and Michoacau.

General characters. — In size nearly equal to Myotis velifer. Length,
<S5 to 95; forearm, 40 to 4G. Calcar thick and distinct, usually termi-
nating in a well-marked pointed projection. Free border of uropata-
gium thickened and densely haired. Ears moderately long; laid for.
ward they reach 3 to 5 mm.l)eyond nostril. Wings from point between
i'likle and base of toes, but nearer latter.

MYOTIS THYSANODES. 81

Uars. — The ears (PI. J, fig. 5) are moderately long and obtusely
pointed; laid forward they reach 3 to ."> mm. beyond the tij) of the nose.
Anterior border of auricle straight or slightly convex through basal half,
then more convex for a short distance, after which it is nearly straight
to the rounded tip; posterior border at hrst straight or slightly (con-
cave, sloping rapidly backward to the widest point at about mid-height,
below which tbe border becomes convex and continues so to the well-
marked basal notch. Basal lobe distinct and moderately large.

Tragus long and slender, the anterior border straight or slightly con-
cave at base, then straight or slightly convex to near the tip, just below
which the border is always convex. Posterior border with a well-
developed lobe at base, widest part of tragus through this lobe or
immediately above it. A more or less developed notch above the lobe.
Beyond this notch the border is at first strongly convex, then slightly
concave below the tip, which is thus always bent backward. Posterior
border indistinctly crenulate.

Memhriuies. — The membranes are moderately thick and dark colored.
Uropatiigium noticeably more leathery than wing membranes, distinctly
thickened at free edge, sparsely haired on proximal fourth both above
and below, the rest of the membrane with a few scattered hairs, which
become more abundant toward the free border, where they form a con-
spicuous fringe both above and below (PI. II, fig. 5). Wing from side
of foot, just below base of toes.

Feet. — Feet (PI. II, tig. 5) large and strong, half as long as tibia?.
Toes (without claws) slightly longer than sole, scarcely united by mem-
brane at extreme base; all sparsely haired. Calcar di.stinct and thick,
considerably longer than free border of interfemoral membrane, termi-
nating distinctly, but usually without well-developed lobule.

Fur and color. — There is nothing peculiar in the distribution of the
fur in this species, except the thickly haired border of the uropatagium.

In color the fur is everywhere light, dull, yellowish brown, distinctly
paler ventrally, the hairs everywhere dusky slate at base. The color
is subject to considerable individual variation in shade. The palest
specimens are yellowish wood brown inclining to clay color; the darkest
specimens dull raw umber. The belly varies from clear gray scarcely
tinged with yellow to a strong yellowish gray, and in other specimens to
dull brownish gray. The exact shades are very variable and impossible
to describe accurately.

SlcuU. — Skull (fig. 11 ft, and fig. 12 h) large, exactly the same size as
that of i¥. i-elifer, but more lightly built. Brain case oval in outline,
abruptly rounded posteriorly, occipital region inflated and lacking well-
formed ridges. Forehead moderately elevated above muzzle. Distance
from posterior border of last upper molar to tip of hamular greater
than width between alveoli of posterior molars.

Although the skull of this species and that of M. velifcr are equal in
size, that of the former is easily distinguished by its more inflated brain
2772— No. 13 G

82

NORTH AMERICAN FAUNA.

Fig. 16.— Maxillary teeth of four speci-
mens of Mifotis thysanodes, showing in-
dividual variation in form ;uirt position
of premolars: a, specimen from Patzcu-
aro, Michoacan ; b, c, and d, from Ha-
cienda La Parada, San LuisPotosi( 5).

case, forehead more abruptly elevated above muzzle and rounder less
angular occiput. When viewed from above, the posterior margin of
the brain case is rounded in M. thysanodes, truncate in M. velifer. When
viewed from behind, the brain case in M. thysanodes is broader in pro-
portion to its height than in M. velifer and lacks the conspicuous
occipital crest of the latter. The posterior part of the palate, from the
last molars to the tips of the hamulars, is shorter in proportion to

the distance between the hindermost
molars in 31. velifer than in M. thysanodes.
Teeth. — Upper incisors diverging at the
tips. First and second upper premolars
very variable in relative position and
size. The first is always much longer
than the second and has the crown at
least one-fourth larger in cross section.
The cross section of the first may, how-
ever, be nearly twice that of the second.
In some specimens these two small pre-
molars are perfectly in the line of the
tooth row, the first in contact with the
canine, the second touching the first, but
separated from the third by a distinct
space. In others the second premolar
while perfectly in line is in contact with the third as well as with the
first. Rarely the second premolar lies slightly external to the tooth
row, while very commonly it is displaced to a varying degree inward,
so much so in some cases as to be almost hidden from the outer side by
the close approximation of the first premolar and the anterior edge of
the third. These variations are independent of age and sex. The
extremes with intermediates of all degrees occur among a dozen of the
females collected by Mr. Nelson at Hacienda La Parada, San Luis
Potosi, August 16, 1892; while specimens with teeth much worn or
Avholly unworn may have the premolars in-
differently greatly crowded and displaced or
wholly in the tooth row (fig. 15 d, fig. 16, and
fig. 17.)

Third premolar triangular in outline, the
outer border abruptly convex in front, and
equal to posterior border; anterior and poste-
rior borders concave; inner apex rounded, not extending back to level
of inner margins of molars. First and second molars trapeziform, the an
terior edge longest, the posterior outer and inner margins successively
shorter. Anterior border straight to near inner edge, where it is bent
abruptly backward, posterior border very slightly concave. First
molar shorter and broader than second, and with anterior border nearly
straight.

Flu. 17. — Abnormal premolar of
Miiotis thysanodes (No. 52228) : a,
crown ; h, side ( X 20) .

MYOTIS THYSANODES.

83

Central lower incisors with crowns compressed and trifid, the next
pair similar but larger, the outer incisors still larger and with crowns
indistinctly terete and quadrituberculate. First and second mandibu-
lar premolars variable in position and in relative size, the first always
the larger. The second is shorter than the first, but in some specimens
its crown has a cross section nearly equal to that of the latter. The
first is always in contact with the canine and usually with the second
premolar also, but may be separated from the latter by a narrow space.
The second premolar is either wholly in the line of the tooth row and
not touching the third, in line and touching the latter, or more or less
displaced inward. Third i)remolar trapeziform, slightly broader than
long.

In dentition Myotis thysanodes shows many points of difference from
M. velifer. One of the most striking of these is the great variability
in the size and position of the first and second upper premolars (figs.
16 and 17), which in M. velifer are comparatively constant. Other dif-
ferences may be seen in the form of the third upper and third lower
premolars. The crowns of the upijer molars are proportionally broader
in M. velifer than in .1/. thysanodes.

Measurements. — Average measurements of 23 specimens of Myotis
thysanodes from four localities are given in the following table:

Average measurements of £3 specimcnH of Myotis thysanodes from 4 localities.

Locality.

a

11

"Si

a

1

Tail vertebra;.
Tibia.

i

o

a
u

1

s

H

1^

a

o

a

Vl

o

1

t

California : Old Fort Tejon

01(1 Fort Tejon

'1

10
.3
10

87
87
89
90.4

36 18

37 17. 6
37 i 16.7
36.9 17.4

8
8

8.9
8.9

41 1 6
41.2 6.3
41.8 1 6.7

69

69.2
71.5
73.6

18

17.6
18.3
18.5

12
11.8
11.9
12.2

11

10.5

10.8

San Luis Potosi: Hda.LaParada

42.7

6.7

10.5

'Type.

Specimens examined. — Total number 88, from the following localities :

California: Dulzura, 1 (skin, Miller coll.); Old Fort Tejon, 16.
Chihuahua: East side San Luis Mountains, 2 (skins).
San Luis Potosi : Hacienda La Parada, 62 (6 skins).
Michoacan: Patzcuaro, 5 (2 skins).

Jalisco : La Laguna, Sierra de Juauacatlau, 1 (skin) ; Sierra Nevada do Colima,
1 (skin, Am.Mus. Nat. Hist.).

General remarks. — Myotis thysanodes needs no close comparison with
any other species occurring in Mexico or the United States. Its large
size separates it from all others but M. velifer, while from the latter the
ciliated free border of the uropatagium, peculiar thickened calcar,
larger ears and paler color together with the cranial and dental
characters readily distinguish it.

84 NORTH AMERICAN FAUNA.

lu certain respects M. fhysanodes resembles M. evotis. The color is
very much the same, while the ears in tliese two species reach their
maximum (leveloi)ment amon^- the species of this genus found in Nortli
America. The free border of tlie interfemoral membrane in J/, evotis
shows a sligiit tendency to the ciliation so conspicuous in M. thysanofles.
Myotis crotis is, however, a smaller animal and lias ears proportionally
longer than in M, thysanodes, while the free border of the interfemoral
membrane is never distinctly ciliate.

That this species is the same as the South American Myotis albescens
is exceedingly unlikely. Dobson, who has seen the type of the latter,
gives for it the following characters, which do not in the least apply to
the present species: "Ears shorter than the head; laid forward, the
tips do not reach to the end of the muzzle; calcaneum feeble, termina-
tion indistinct; above dark brown." Moreover, Mr. Oldfleld Thomas,
who has compared specimens of Myotis thysanodes v^'ith the albescens in
the British Museum, writes me that the two do not in the least resem-
ble each other, and that M. albescens is allied rather to M. vclifer. For
further discussion of the question, see under the latter species.

In Dr. Harrison Allen's recent monograph (p. 93) a specimen of this
species in my collection, taken at Dulzura, [misspelled Dalyura], Cal.,
is recorded as a variety of 'F. albescens velifer.'' A specimen from Old
Fort Tejon, Calilornia, in the Biological Survey collection, is labeled by
Dr. Allen ' V. snbnlatusj' while fifteen others of the same species from the
same locality are marked ' V. albescens?.^ One of the latter (Xo. 20827),
however, is recorded as ' V. albescens evotis^ (p. 90).

Dr. T, S. Palmer has kindly furnished me with the following account
of the (jolony from which the type of Myotis thysanodes was taken:

In July, 1<S9I, -while one of the parties of the Death Valley Expedition was collect-
ing at Old Fort Tejon, California, several species of hats were ohserved. The most
ahundant was a small VenpertUlo [^^Mijotis'], which could be seen at dusk flying about
the oak trees near the old barracks in great numbers, and passing in and out of the
ruined buildings. A long two-story adobe building, with the roof still intact, seemed
to be the center of attraction, and about sundown bats could be seen streaming forth
from a window in one of the gables. On the morning of July 5 an examination was
made of the attic of this building, and the bats were found clinging to the ridgepole
and the rafteis, literally by thousands. Individuals of all ages, from recently born
young to adults, were hanging together in bunches as big as a bushel basket. Others
found concealment in cracks and crevices, but very few were flying about. Evidently
the colony had occiipied the attic for several years, but it was too dark to see whether
more than one species was present.

A sack was carried along under the ridgepole and specimens swei)t into it from sev-
eral of the larger bunches. In this way more than a hundred bats were collected in
a few minutes. As soon as they were disturbed they uttered a peculiar 8<iueakiug
note and flew about in a confused manner in their eft'orts to escape. The sack was
carried out under one of the oak trees and the specimens examined; 160 had been
captured, and of these 25 were preserved' and the remainder allowed to escape.
Some of the bats which had been given their liberty attempted to fly back to their
retreat, but da/ed by the sunlight took refuge in the branches of the nearest tree;

' Sixteen proved to be Myotis thysanodes; the others were M. yumanensis.

LASIONYCTERIS.

85

others made no attempt to escape, except to crawl up the trunks of tlio trees, wlicre
they remained until dark. Some of the young ones failed to find their way back to
the building, and remained about the spot for several days.

Genus LASIONYCTERIS Peters.

1864. ScoiopMlus H. Allen, Monogr. N. Am . Bats, p. 27 (part, not Seotophilus Leach, 1821 ) .

1865. Lasionycteris Peters, Monatsber. K. Akad. Wiss.

Berliu, p. 648. Type WnpertUio noctiragans

Le Coute.
1870. Cnephaiophihis Fitzinger, Sitzungsber. K. Akad.

Wissensch., Wieu, LXII, Abth., I, p. 8 (part).
1875. Vespcrides Coucs in Cones' and Yarrow's Zool-
ogy of Wheeler's Exped., p. 83. Type T'es-

perfilio noctivagans Le Conte.
187N. Vesperugo Dobsou, Catal. Chiroptera Brit. Mus.,

p. 183 (part).
1893. Lasiomjcleris H. Allen, Monogr. Bats N. Am., p.

104.

Fig. 18. — Skull of Lasionycteris noc-

tivarians (>:2.)

»», o Q — 36.

Type species. — Lasionycteris noctivagans (Le
Coiite).

Geographic distnhntion. — Tlie range of tlie
genus Lasionycteris is tlie same as that of the type and only known
species.

Generic characters. — Dental formula : «', q^ ; c, ^pr ; pm, '^t

Skull (fig. IS), flattened; rostrum very broad in pioportion to brain
case, strongly concave on each side back of the nasal aperture; dorsal
profile of skull nearly straight and sloping gradually from external
nares to occiput, whicli is scarcelj- angular, and always without sagittal
crest. Ears short, nearly as broad as long ; when laid forward, reaching
barely to nostril; basal lobe very large. Tragus short, straight, and
bluntly rounded at tip, width much more than half length of anterior

margin. Back of iu-
terfemoral mem-
brane furred on basal
half. Mamma', 2.

General remarJis. —
Among the Ameri-
can YespertiUonida'
the genus Lasionyc-
teris is readily distin-
guished by its dental formula, combined with its short, broad ears,
broad tragus, and partially furred uropatagium.

The genus Lasionycteris is peculiar to North America, where it is
represented by one widely distributed species whose characters are
remarkably constant throughout its range.

Fig. 19. — Teeth of Lasinmjcfens noctivagans (X5).

86

NORTH AMERICAN FAUNA.

LASIONYCTERIS N0CTIVAGAN8 (Le Coute). Silver-liaired Bat.

1831. Vesperfilio iiociivuf/anfi Le Conte, McMmtrie's Cuvier's Animal Kingdom, I, p. 31,
June, 1831.

1831. VespertUio auduhoni Harlan, Monthly Amer. .Journ. Geol. and Nat. Hist., I, p.
220, PI. II, November, 1831 (Philadelphia, Pa.;.

1835. respertiUo jmlverulenlus Temmiuck, Monogr. de Mammalogie, II, p. 325 (Mis-
souri River).

1864. Scotopliihis noctiva{/ans H. Allen, Monogr. N. Am. Bats, p. 39.

1865. Lasiunycteris noctivu(ians Peters, Monatsber. K. Preuss. Akad. Wiss., Berlin,

p. 648.
1878. Vespenigo noetivagans Dobson, Catal. Chiroptera Brit. Mas., p. 238.
1893. Lasionycteris noctii-aganx H. Allen, Mouogr. Bats N. Am., p. 105.

Type locality. — Eastern United States.

Geographic (listrihution. — North America, from Atlantic to Pacific;
probably not breeding south of the Transition Zone.

General characters. — See generic characters given on page 85.

Color. — The fur is deep, blackish, chocolate brown throughout, many
of the hairs on the back, belly, and furred part of interfemoral mem-
brane tipped with silvery white. The white tips are most numerous
on middle of back. They are absent, or nearly so, from face, crown, and
throat.

Skidl and teeth. — The cranial and dental characters of Lasionycteris
7ioctivagans have been sufficiently described in the diagnosis of the
genus.

Measurements. — The average measurements of 21 specimens of Lasi-
onycteris noctivagans from eight localities are given in the following
table :

Average measurements of 21 specimens of Lasionycteris nociivaqans from S localities.

,a

S

p

a

S

fcr

XI

t(3

e

©

Locality.

>

.2

.a

e

s

II

3

09

a

O

o

.a

o

IT"

^"^

H

H

H

fe

1^

H

hi

Nl

ts

H

New York: Sing Sing

10

10.5. 8

42.4

17.1

7.9

41.1

5.3

73.4

15.9

14.1

6.7

Montana : Elatbead Lake

2??

100.5

41

16.2

7.5j

42

4.5

75

15.6

14.1

6.2

IcT
2??

97
95.5

38
32.5

16
16

8
8.4,

39

40

4.6
6.2

68

16

15.8

12
11.8

6

Nevada : Badger

6

Oregon: Blue Mountains

Id

97

39

16.4

8

41

6

15.4

11

.5.6

Ci'ooked Kiver

IcT

95

41

17

7

41.4

5

16

12

6

ElKiu

3

101
98

43
44

16.3
15

8.9^

8.6

43
41

4.3
6

16
16

13.3
13

5.7

6

Specimens examined. — Total number 105, from the following localities ;

Alberta: Henry House 2 (skins).

British Columbia: Sumas, 1 (skin, Miller coll.).

California: Nevada City, 1; Nicasio, 2.

Colorado: Rifle, 1.

Massachusetts: Nantucket, 1; North Truro. 6 (skins. Miller coll.).

Montana: Flathead Lake, 2.

PIPISTRELLUS. 87

Nebraska: Platte River, 1.

Nevada : Badger, 2.

New York: Lake George, 6 (2 skins); Leyden, 14; Locust Grove, 4; Lyon>

Falls, 4; Sing Sing, 47 (20 young).
North Carolina; Magnetic City, 1 (skin).
Oregon: Beaverton, 1 (skin, Miller coll.); Blue Mountains, 1; Crooked River,

1; East base Cascade Mountains, near Mount Thielsou 1 (skin); P^lgin, 3;

Harney, 1; Salem, 1.
Pennsylvania: Carlisle, 1.

General remarlts. — Lasionycteris noctivagans is one o.f the most easily
recognized of Nortli American bats. Its peculiar color alone is suffi-
cient to distinguish it from all others found in the region where it occurs.

Genus PIPISTRELLUS Kaup.

1829. Pipistrellus Kaup, Skizzirte Entwick.-Gesch.,u. Natlirl. Syst. d. Europ. Thierw.,

Th. I, p. 98. Type VespertUio pipiatreUus Schreber.
1839. Vesperugo Keyaerling & Blasius, Wieginann's Archiv f. Naturgesch., oter

Jabrg., Bd. 1, p. 312 (part).
1856. Nannugo Kolenati, Allgem. Deutscb. Naturbist. Zeitg., Dresden, Neue Folge

II, 131, 169-172. Based on natltusii, pipistrellua, and kuhlii.
1856. Hypsugo Kolenati, Allgem. Deutscb. Naturbist. Zeitg., Dresden, Neue Folge,

II, pp. 131, 167-169. Included the species maurus and krasclieninikowii.
1864. Scotopliilus H. Allen, Monogr. N. Am. Bats, p. 27 (part, not Scotophilus Leach).
1878. Veaperugo Dobson, Catal. Chiroptera Brit. Mus., p. 183 (part).
1893. Vesperugo H. Allen, Monogr. Bats N. Am., p. 121.

Type species. — Pipistrellus pipistrellus (Schreber).

Geographic distribution of genus. — The greater ])a,vt of the Eastern
Hemisphere, and throughout the southern half of North America. Exact
limits of tlistribution not known.

Generic characters. — Dental formula:

. 2-2 1-1 2-2 3-3 o,

*' 3=3 '^' 1-1 '^'"'212'"*' 3=3= ^'^^
Skull (figs. 21 and 22) small and lightly built, varying somewhat in form
among the different species. Braincase usually more inflated than in
Vesperiilio and Lasionycteris, but rostrum proportionally as broad as in
these genera. Ears (fig. 20) distinctly longer than broad and tapering
to a narrowly rounded tip. Tragus straight or slightly curved for-
ward. Back of interfemoral membrane sprinkled with hair on basal
third. Mammai, 2.

General remarks. — The members of the genus Pipistrellus may be
recognized by their dental formula and small size. The bats of the
European g,eui\s, Pterygistes ' {Pterygistes noctula and P. leisleri), which
have the same dental formula, are large, heavily built, and altogether
different in appearance.'^

'Pterygistes Kaup, Skizzirte Entwick.-Gesch. u. Naturl. Syst. d. Europ. Tbiervv.,Th.
I, p. 100, based on f'espertilio prater us Knh] (^ J'. Hoc<«/a Schreber) and V. leisleri Knhl

^For remarks on the generic characters of ' Noctulinia ' (^Pterygistes), .see H. Allen,
Proc. U. S. Nat. Museum, 1893, p. 30.

88 NORTH AMERICAN FAUNA.

In America the genns is ropresentetl by three species, all of which
are strictly coiijjeueric with PiplstreJlus pipistreUu.s. Of the American
species P. subjlarvs resembles P. pipisfrcJJns most closely, but is dis-
tinguishable at a glance by its much longer thumlt.

KEY TO AMERICAN FORMS OF I'lriSTllELLUS.
Tragus blunt with tip bent forward:

Forearm about 31 mm. ; colors very pale hcspcriis (p. 88)

Forearm about 28 mm. ; colors darker aiistralis (]i. 90)

Tragus tapering and straight:

Forearm 30 to 32 rrracntcls (p. 93)

Forearm 33 to 36—

Color yellowish brown sxhjiavus (p. 90)

Color drab brown ohscurus (]>. 93\

PIPISTKELLUS KIvSPEKUS (H.Allen).
1864. Scoiopliilu's hespenis H. Allen, Monogr. N. Am. P>ats, p. 43.
1878. Vesprrngo liesperus Dobson, Catal. Chiroptera Brit. Museum, p. 228.
1886. VesperiKjo merriami Dobsou, Ann. & Mag. Nat. Hist., .5th ser., XVIII, p. 124.
1893. resperitgo luspenifi H. Allen. Monogr. Hats N. Am., ]). 128.

Type locality. — Fort Yuma, Cal, Type No. 5406, CJ. S. National Museum.
Oeofjriipliic (listrihiition. — Lower Austral zone in the Western United
States from western Texas to the Pacific Coast. Limits not known.
General characters. — Size very small (forearm about 26); thumb short

(about one-eiglith of forearm); ear shorter

#and more bluntly rounded than in other
^-r;^ American members of the genus, reaching

/^)'' I'A barelj'- to nostril when laid forward ; tragus
jim^Mi Wunt and distinctly bent forward at tip;
Vwf''^^m^4 feet very small, about half as long as tibia:
W''W^0' barely 1 mm. of tip of tail free from mem-
■.i^^^^'^ / brane; color very pale.
^ Pars. — The ears (fig. 20 h) are short,

FiG.2o.-Ear of (rr) ripistreiius ml- reaching barely to nostril when laid for-

flavvs a^A (fi) r.hesverus (X2). , m, ,•■,-, n • i •

ward. The anterior border of auricle is
strongly convex from well develoi)ed basal notch to region about
middle, where it becomes straight and remains so almost to narrowly
rounded tip. Posterior border concave immediately below tip, then
strongly convex to basal notch. Basal lobe well developed, separated
from auricle by a deep notch and joining fiice at point below line of lips,
and slightly behind posterior corner of eye. The fur of the head extends
over dorsal surface of ear to slightly beyond the basal third. Otherwise
the ear is naked except for a sprinkling of fine hairs on inner surface.

Tragus less than half length of ear, broadest just below tip; ante-
rior border straight throughout greater part of its length, but strongly
concave immediately below tip; posterior binder strongly convex from
tip almost to notch above well developed basal lobe.

jMemhranes.—T\\Q membranes are thin and delicate. Uropatagium
very sparsely furred at extreme base, otherwise naked except for a few
scattered hairs which are most numerous on the basal half. Wing

PIPISTRELLUS HESPP:RUS.

89

membranes attached at base of toes. Uropatagiuui exteiuliiig almost
to extreme tip of tail.

Feet. — Foot small, distinctly less tlian half as long- as tibia, naked or
with a few almost invisible whitish hairs on dorsal surface. Calcar
about as long as tibia, scarely keeled on posterior edge, terminal lobe
absent or very indistinct.

Fur and color. — The fur extends on basal third of ears, but barely
reaches extreme base of interfemoral membraue, and ou wing mem-
branes invades merely a very narrow strip close to body.

Color light yellowish gray or whitish gray, the fur everywhere deep
plumbeous at base. In some specimens the hairs on the back have faint
dark subterminal areas which, however, are visible on close inspection
only. Ears, muzzle, face, and membranes black. A narrow whitish
border on wing membrane between foot and fifth finger.

This species is apparently much more constant in color than P. suh-
Jiavus, but the absence of a good series of skins leaves the range of
individual variation in color a matter of uncer-
tainty.

Skull. — The skull of PijristreUus lieaperus
(figs. 21 r(, and 22 h) is very small, thin, and
papery. That of an adult male from Fort
Bowie, Arizona, measures 11.4 mm. inoccipito-
nasal length, 0 mm. in zygomatic breadth, and
4 mm. in occipital depth. The dorsal outline
is nearly straight from external .nares to occi-
put, though there is a slight concavity between
the orbits and a slight convexity over the brain case. Muzzle broad
and nearly flat, slightly concave ou each side of median line. In gen-
eral the skull of PlpistreUus hesperus suggests a miniature of that of
Lasionycteris.

Teeth. — The teeth of Pipistrelhis liesperus (fig. 23 «) do not differ
materially from those of P. suhjianis. The anterior upper premolar is
minute (much smaller than the smaller upper incisor) and usually
thrown out of the tooth row by the second premolar, the anterior edge
of which is generally in contact with the canine.

Measurement H. — See table, page 95.

Specimens examined. — Total number 127, from the following localities:

Arizona: Beaverdam, 1; Fort Bowie, 1 (skin); Grand Cauon, 2; Guadalupe
Canon, Cochise County, 4 (skins); Little Colorado, 2 ; Dos Cabezas, l(skin);
Keam Canon, Navajo County, 1 (skin); New River, Maricopa County, 1;
Yuma, 2.

California: Borax Flat, Mohave Desert, 3; Colorado Desert, 1 (skin); Death
Valley, 4; Funeral Mountains, Inyo County, 1 ; Furnace Creek, Death Valley,
1 ; Grapevine Spring, Death Valley, 1 ; Independence, 1 ; Hot Springs Valley,
Inyo County, 2; Jacumba, San Diego County, 2 (skins); Keeler, 1; Kern
River, 3 ; Kernville, 1 ; Lone Pine, 3 ; Owens Lake, 1 ; Palm Springs, 3 ; Paua-
mint Valley, Inyo County, 6; Pananiint Mountains, 4; Poso Creek, Kern
County,! (skin); Saline Valley. Inyo Connty, 1; San Emigdio, 1; Santa

Fig. 21.— Top view of sknil of

(a) Piplstiellus hespervs and
(6) P.suhflavus (.-2.)

90 NORTH AMERICAN FAUNA.

Ysabel, Sau Diego County, 18; Three Rivers, 7; Twiu Oaks, 2; Vallecitas, San

Diego County, 3 ; Whitewater, 1 (siiiu) ; Yosemite Valley, 1 ; Old Fort Yuma,

1 (type).
Colorado: Grand Junction, 4.
Lower California: Sau Fernando, 7 (Miller coll.).
Nevada: Gold Mountain, Esmeralda County, 1; Pahranagat Valley, 1; Vegas

Mountains, 1; Vegas Valley, Lincoln County, 1.
New Mexico: Dog Spring, Grant County, 7 (skins) ; Fort Wingate, 1; Grant

County, 1 (skin).
Texas: Chinate Mountains, 1; El Paso, 4 (1 skin); Paisano, 5; Pecos River, 1.
Utah : St. George, 2.
Washinton: Alniota, 1.

General rcmarlcs. — Pipistrellus hesperus is readily distinguishable
among North American bats by its dental formula, small size, blunt
tragus, and pallid color. It needs no comparison with any other species.
At the southernmost extremity of its known range a subspecies slightly
different from that occurring farther north has been differentiated.
Otherwise the species is remarkably constant in all its characters.

PIPLSTRELLUS HESPERUS AUSTRALIS subsp. nov.

Ti/jje from Barranca Hiarra, .Jalisco, Mexico; altitude about 3,000 feet. Adult 9 (in
alcohol), No. 52112, U. S. National Museum (Biological Survey collection). Col-
lected May 14, 1892, by E. W. Nelson. Original number, 2614.

General characters. — Slightly smaller than true Pipistrelltis hesperus;
fur shorter; color apparently darker and browner.

Ears, membranes, and feet. — As in the typical subspecies.

Fur and color. — The fur is shorter than in specimens of true hesperus
taken in April and May, but in distribution it shows no peculiarities.

Color darker and browner than in specimens of true hesperus that
have been immersed in alcohol for a similar length of time. Until skins
of the southern animal are examined the actual color differences between
the two forms can not be determined.

Measurements. — The measurements of the type and the averages of
four specimens from the type locality are given in the table of measure-
ments on page 95.

Specimens examined. — Four, all from the type locality.

General remarls. — Pipistrelliis hesperus austraUs is a fairly well
marked subsi)ecies characterized by slightly smaller size, shorter fur,
and darker color than in the typical form. The material by which it is
represented is so poor, however, that all the characters can not be
determined with certainty.

PIPISTRELLUS SUBFLAVUS (F. Cuvier). Georgian Bat.

1832. iVespertiUo georgiaiius F. Cuvier, Nouv. Ann. Mus. d'Hist. Nat., Paris, p. 16.
(Not determinable.)

1832. Fesjjer<iKo s»&/?arMS F. Cuvier, Nouv. Ann. Mus. d'Hist. Nat., Paris, p. 17. (De-
scription good.)

1835-41. VespertUio eri/flirodactyhis Tommiuck, Monogr. de Mamm., II, 13rae Monogr.,
p. 238.

1861. Svotoj>hihi)i j/eorgiauiis 11. Allen, Monogr. N. Am. Bats, p. 35.

PIPISTRELLUS SUBFLAVUS. 91

1878. Vespenigo i/eoriiianiis ])obson, Catal. Chiroi»tera Brit. Mns., p. 235.
1893. Vespenigo caroJineiisis H. Allen, Mouogr. Bats N, Am., p. 121 (not Vespertilio
carolinensis Geoff.).

Type locality. — Eastern United States; probably Georgia.

Geographic distribution. — Austral zones and casually parts of Transi-
tion zone in tbe Eastern United States, from tbe Atlantic Coast west to
Iowa and eastern and soutbern Texas.

General characters. — Size small (forearm, about 34); tbumb long
(febout i forearm) ; ear wben laid forward reacbiiig- sligiitly beyond nos-
tril; tragus straight, tapering to a broadly rounded tip; feet small,
slightly more than half as long as tibia; terminal 2 mm. of tail free
from membrane; liairs on back mostly distinctly tricolored; general
color light yellowish brown, undulated with darker brown.

Ears. — The ears (fig. 20 a) are considerably longer than in P. hesperus,
reaching, when laid forward, just beyond nostril. In general form the
ear is much as in P. hesperns, but the auricle is slightly narrower, and
the basal lobe is smaller and separated from auricle by a slight notch
0Dl3^ On dorsal surface of ear the fur of head extends scarcely to basal
third. Otherwise the ear is naked except for a sprinkling of fine hairs
on inner surface.

Tragus about half length of ear, broadest opposite anterior base and
thence tapering gradually upward to bluntly rounded tip which is
turned slightly backward. Anterior border slightly concave at base,
then gently convex to tip. Posterior border slightly concave imme-
diately below tip, then strongly convex almost to notch above well
developed basal lobe.

Membranes. — The membranes are thin and delicate. Uropatagium
thinly furred on basal fourth, otherwise naked except for a few scat-
tered hairs along veins on lower side. Wing membranes attached at
base of toes. Uropatagium attached at base of terminal caudal
vertebra.

Feet. — Foot large, distinctly more than half as long as tibia, covered
with consj)icuous light-brown hairs on dorsal surface. Calcar dis-
tinctly longer than tibia, scarcely keeled on posterior edge, terminal
lobe absent or very indistinct.

Fur and color. — The fur extends on base of ears and interfemoral
membrane and on wing membranes to line joining knee and middle of
forearm.

Color light yellowish brown, uniform on the ventral surface, but on
the back clouded to a varying degree with darker brown. The hairs
on the back appear to be of two kinds. The main body of the fur is
made up of short hairs (about 0 mm. in length), which are deep plum-
bens from base to a little below middle, then yellowish brown almost
to extreme tip, which is dark brown. Intermixed with these shorter
hairs are others which are much longer (about 10 mm. in length) and
clear yellowish brown to extreme tip.

92

NORTH AMERICAN FAUNA.

Typical PipistrellHs suhflarKs presents a wide range of individual
variation in color. This is due to the extent of the terminal dark bands
on the hairs of the ])ack, and also to the exact shade of the yeUowish
subterniinal bands. The yellowest specimens that I have seen were
taken at Washington, D. (J., during May and June.

^kull. — The skull ot Pipisfrelliis suhfarits (figs. 21a and 22//) is
larger than that of P. hesperns. That of an adult
male from Washington, I). C, measures 13 mm. in
oceii)ito nasal length, S mm. in zygomatic breadth,
and r> mm. in occipital depth, Tlie dorsal outline
is nearly straight from the anterior nares to a point
immediately behind the orbits, then strongly con-
vex to occiput. Muzzle narrow and arched, the
concavities on each side nearly obsolete. In gen-
eral the skull of Pipistrelltis suhflavus suggests a
Fig. 22.— Side view of skull miniature of that of the smaller forms of Vespertilio.
ona)PipMushesi,eru. Teeth.— 'Hh^ tccth of PipistreUiis suhflavus (fig.

and (h) l'..sul)flavt(s { 2). -L j \ e>

23 h) are larger than those of P. hesperus but essen-
tially similar in form. The anterior upi)er premolar is large (about the
size of tlie larger upper incisor) and generally fully in the tooth row.

Measurements. — See table, page 057.

Specimens examined. — Total number, 2i;>, from the following local-
ities :

Alabama: C4roensboro, 2.

District of Columbia: Washington. 17 (11 skins).

Indian Territory : Stihvell, l;^. ,

Louisiana: Mer Rouiie, 10; Houma, 2 (skins).

Maryland : Marshall Hall,

5 (skins); St. Georges

Island, 2 (skins).
Mississii)pi : Washing-
ton, 8.
Missouri : Marble Cave,

Stone County, 70.
New York : Sing Sing, 33.
North Carolina: Raleigh,

7 (skins); B or tie

County, 2 (skins).
Pennsylvania : Carlisle,

7 (1 skin).

Tennessee : Hickman County, 1 (skin) ; Arlington, 3 ; Big Sandy, 10; Danville, 4.
Texas : Clear Creek, Galveston County, 1 ; Brownsville, 1.
Virginia: Cedarville, 6 (skins. Miller coll.) ; Fredericksburg, 6 (skins); Hanip-

sliire County, 1 (skin); Wytheville. 2.

General remarls. — The Georgian bat, Pipistrellus suhflavus, is so
readily distinguished among the species of the region it inhabits that
detailed com])arisons are scarcely necessary. Its dental formula, small
size, relatively large thumb, distinctly tricolored fur and general yel-
lowish color are unmistakable characters.

Fig. 23.— Teeth of (a) Pipistrelhis hespems aad (6) P.siibflavus
(X5).

PIPISTRELLUS VER.ECRUCIS. 93

PIPISTRELLUS SUBFLAVUS OBSCURU>S subsp. uov.

Type from Lake Giorge, Warren County, N. Y. Adult 9 (skin) No. 67723, U.S.
National Museum (Biological Survey collection). Collected September 6, 1894, by
Walter K. Fisher. Original number, 198.

General characters. — Size and proportions as in typical snhfavus, but
color duller and less yellow, and dark tips of shorter liairs on back
more conspicuous.

Ears, membranes, feet, and fur. — As in typical svhflavus.

Color. — Fur everywhere blackish slate at base. Middle baud ou
shorter hairs of back dull, pale, wood browu or Isabella color. Tii)s of
these hairs dusky brown, and much more consx)icuous than iu true sub-
fiavus. Long hairs of back pale wood brown. Belly uuiforni Isabella
color, iu some specimens inclining toward wood brown, but seldom show-
ing any ai)proach to the bright yellowish browu of true subjiavus.

A melanistic si^ecimeu is dark chocolate brown throughout. Two
others are rich reddish brown. Iu all three of these abnormal individ-
uals the characteristic variegation of the fur of the back still persists.

Shidl and teeth. — I cau find no cranial or dental characters to distin-
guish Pipistrelhis subjiavus obscuriis from the typical subspecies.

Measurements. — See table, page 95.

Specimens examined. — Thirty four (seven skins), all from the type
locality.

General remarks. — Pipistrellus subjiavus obscurus is readily distin-
guishable from true subjiavus by its darker, duller, less yellow color.
The difference is especially noticeable on the ventral surface, which is
generally a rich yellowish wood browu in tjincal s ubfavus, dull Isabella
color in obscurus. The darker hue of the back in obscurus is due partly
to diflterences in the color of the long hairs, and of the middle bands
of the short hairs, and partly to the more extended dark tips of the
short hairs. Like the typical form, Pi^ristrcllus subjlarus obscurus
varies considerably in color, so that individual specimens of either sub-
species, especially those that are not fully adult, are sometimes difficult
to identify. When series are compared, however, the difl'erential char-
acters at once become apparent.

PIPISTRELLUS VER^CRUCIS (Ward).
1891. reaperuyo vcrwcriicis Ward, Am. Naturalist, XXV, p. 745, August, 1891.

Ty2)e locality. — Las Vegas, Jalapa, Vera Cruz.

Geographic distribution. — This species is known from the type locality
Jiily.

Characters. — As I have seen no specimens of Pipistrellus verwcrucis,
I copy the original description.

I All six specimens were indistinguishable one from another in point of color. The
ollowing color-description is taken from a dried skin, whereas all the rest oi" the
lescriptiou is taken from a specimen preserved in alcohol.

94 NORTH AMERICAN FAUNA.

Hairs of back clove-browu for basal half, followed by two equal zones respectively
broccoli-brown and clove-l>rown; some of hairs furthermore tipped with light Van-
dyke-brown, giving a decidedly "rusty" tone to the back. Ventral surface ; bases
of hair slightly lighter than those of back, followed by light-hair brown, producing
a grayish or smoky effect.

Wing membranes naked, except a very limited area on upper surface along sides
of body, not exceeding 3 or i nun. in width ; and on lower surface, the area included
between a line passing from the middle of humerus to the knee and the side of the
bodj' is scantily haired.

Interfemoral membrane wilh a small, triangular patch of hair on its upper sur-
face, covering base of tail, and extending to one-fourth of its length.

Legs and arms naked. Wing extending from base of outer toe. Antebrachial
membrane losing itself at middle of radius. Two caudal vertebr;e free from
membrane.

Black glandular prominences between eyes and nostrils well developed, fringed
with longish hairs on both upper and lower edges, and with three or four long,
black, bristly hairs growing from its upper surface.

Inner edge of ear conch evenly convex. Outer edge coming up in an even, sweep-
ing curve from angle of mouth to level of tip of tragus, where it meets a slightly
concave line leading up to the obtusely rounded tip. A nearly semi-circular anti-
tragus is developed from that part of the conch passing below the tragus. Bone of
inner margin of tragus concave, thus throwing this organ forward, followed by a
straight margin. Bone of outer margin with a subtriangular lobe, followed by a
deep notch, above which the greatest width is (quickly reached. From here a nearly
straight line leads to the tip, which is obtusely rounded.

Measurements in millimeters: Length of head and body, from tip of nose to base
of tail, 37.5; length of tail, 36; length of tail beyond membrane, 3; length of head,
15; height of ear, from notch between antitragus and conch to tip, 10; height of
tragus, inner margin, 4.5; height of tragus, outer margin, 6; greatest width of
tragus, 2; length of antitragus, 2; height of antitragus (approximately), .75; length
of forearm, 31; length of thumb, including claw and excluding metacarpus, 7,5.
Second digit — metacarpal, 29. Third digit — metacarpal, 30.5; first phalanx, 11.5;
second phalanx, 11; cartilaginous tip, 5. Fourth digit — metacarpal, 29; first pha-
lanx, 10; second phalanx, 7; cartilaginous tiji, 2.5. Fifth digit — metacarpal, 28;
first phalanx, 8.5; second phalanx, 5; cartilaginous tip, 1. Interspace between
tips of third and fourth digits, 1() ; interspace between tips of fourtii and fifth digits.
37; interspace between tip of fifth digit and juncture of membrane with foot, 42;
extent of outstretched wings, 212; length of tibia, 13.5; length of foot, 9; length
of calcaneum, about 8.

0^0 i_i 9_g S-^

^^^^^^'o'l-l'i-jU^^^^tsi].

Middle upper incisors separated by 1.5 mm., inclined forwards and inwards; a
large internal cusp on posterior-external edge halfway up from base to tip. Outei
incisors simple, conical, inclined parallel to their respective inner mates, separated
from canines by about .75 mm. Lower incisors tri-lobate, evenly spaced. UppW
canines long, simple, slightly recurved. Lower canines straight, with basal cu8p>
on forward edge only. First upper premolar interior to tooth line, visible from tht
exterior. Second upper premolar longer than any of its corresponding molars.

A prominent conical oxcresc"ence is on the lower gum, opposite the space between
the premolars, in front of which the point of the upper canine passes. Two muc!)
less prominent excrescences are on the upper gum immediately above this lowei
one. Type No. 527 i , Las Vegas, V[era Cru]z, February 19, 1891. Collectors, H. L.
Ward and C. M. Teran.

General remarks. — Pipistrellns rera'eriicis differs from P. snhflaviisw
its smaller size, relatively longer tliumb, and browner, less yellow coloi

VESPERTILIO.

95

No specimens of tbis species have been obtained by the held agents of
the Biological Survey, nor are any known to be in American museums.

Average measurements of Xorth American forms of Pipistrellns.

Locality.

ht'iperus Washington : Almota

California : Fort Yuma . . .

Santa Tsabel .

Colorado: Grand Junction

Texas : Paisano

auitialis Jalisco: Barranca Ibarra.

Barranca Ibarra.

vera'cnteis I Vera Cruz : Las Vegas,

Jalapa.

suhflavus Louisiana : Mer Rouge . . .

Missouri : Marble Cave . . .
District of Columbia :
Washington.
obsctirus New York : Lake Georsje.

1?
'IcT
10
4
3

'1?
4
'1

10??
10

5??

10

77 32 jl2
70 '28 |11
72.8 32 111. 5
74.6|31 12
79 34. 5 1 12. 8
64 '28 jll.4
63.2 26.8110.7
73. 5^36 13. 5

B

j

0

bO

S

a
■■a

a

^

H

s

q
5

a

0

0

cS

l=H

H

h-!

w

85.1
84.6
84

40.7 15.3
39. 8i]6. 1;

37.8

84. 8 38. 9

15.2

5

5.5i

5.5;

5.4;

5

5.1;

9

7.9
8.1
7.8

30

4

28

3.8

31.6

4

3L3

4

32.5

4

29

4

28.6

3.9

31

7.5

34.6

6.8

33.7

6.8

34

6.8

36

6.8

52 12.4 9

48

51.4

52

55.5

47

45

58

10 j 8.6

n.ej 8.7

12.1! 9.3

12. 4l 9.6

10.4! 8.4

10.7 8

60.814.2

9.8

00

13.9

9.5

62.8

14

9.6

60. 6

14

10

5.4

4.6

5.2

5.4

5.1

5

4.5

6

6.6
6.4

' Type ; measurements by original describer.

Genus VESPERTILIO Linnseus.

1758. Vespertilio Liniupus, Systema Natiine, 10th ed., I, pp. 31-32. Type by elimina-

tiou VespertiUo miiriniis LiiinaMis (not F. muriniis Scbreher, 1775).
1820. Eptesiciis Ratinesque, Annals of Nature, p. 2. Tyjie Eptcs'wiis meJanops Rati-

uesqiie {^VesperiiJio fuscus Beauvois).
1829. Cnephn'us Kaup, .Skizzirte. Entw.-Gescb. n. Natiirl. Syst. d. Europ. Tbierw., I, p.

103. Typo Vespertilio serotinus 8cbrebei'.
1839. Vesperngo Keyserliug & Blasiua, Wiegmann's Arcbiv f. Naturgescb., 5ter Jabrg.,

Bd. l,p.312(part).
1839. T'esperiis Keyserling & Blasins, Wiegmann's Arcbiv f. Naturgescb., 5ter Jabrg.,

Bd. 1, p. 313. Based on tbe 32-tootbed species of ' Vesperugo.'
1841. Xoctala Bonaparte, Iconografia Fauna Italica, I, fasc. XXI, under Vespertilio

alcytlioe. Type ' Vesperugo' serotinus.
1856. Cateorns Kolenati, Allgem. Deutscb. Naturbist. Zeitg., Dresden, Neue Folge,

II, p]). 131, 162-163. Type ' Vesperugo' serotinus.
1856. Meieorus Kolenati, Allgem. Deutscii. Naturbist. Zeitg., Dresden, Neue Folge,

II, pp. 131, 167-169 (included nilssoni, discolor, stirii, leucippe, aristippe).
1861. Scotophilus H. Allen, ^Monogr. N. Am. Bats, p. 27 (part).
1878. T'cspcrugo Dobson, Catal. Cbiroptera Brit. Mus., p. 183 (part).

1892. Adelongcteris H. Allen, Proc. Acad. Nat. Sci. Phila. (1891), p. 466, Jan. 19, 1892.

(Proposed as a substitute for f >8j>erHs, preoccupied in Entomology).

1893. Adelonycteris H. Allen, Monogr. Bats, N. Am., p. 111.

Type species. — Vespertilio mtirinus liinni^ns {=V. discolor batterer)
I — not V. murinus Schreber.

j Geographic (Ustribviion. — Boreal, Austral and parts of Tropical re-
gions in both hemispheres.

9G NORTH AMERICAN FAUNA.

2-2 1-1 1-1 3-3

Generic characters. — Dental formula : *, .ya ; c, , . ; pm, .^^ ; t^h .> q=32.

Skull (figs. 24 aud 25) large and heavily built; rostrum broad in propor-
tion to l)rain case (less so than in Lasioiiyctcris), scarcely concave at sides
back of nasal aperture; dorsal iirofile nearly straight, rising gradu-
ally from external nares to occiput, which in the adult is strongly
angular and provided with a conspicuous sagittal crest. Ears short,
considerably narrower than long, basal lobe well developed, but not
excessively large. Tragus straight, short, directed slightly forward,
broadest near the middle and tapering to a moderately sharp point.
Back of interfemoral membrane wholly naked except for a sprinkling
of hairs on basal fourth. Mammae, 2.

General remarks. — The genus Vespertilio contains the largest Ameri-
can sjiecies'of the Yespertilionine group. Aside from the dental for-
mula, the large size of Vespertilio fiiseus, the only known North Ameri-
can species, is sufficient to distinguish the genus among those occurring
in the region now under consideration.

The North American species is separable into at least five tolerably
well-marked subspecies as follows:

KEY TO THE SCTBSPECIES OF VESPERTILIO FUSCUS.

Size small (total length, 96 to 107; forearm, 40 to 45; longest finger, 68 to 77).

Breadth of mnzzle greater than half length of head propinqiius (p. 100)

Breadth of mnzzle less than half length of head bahamensis (p. 101)

Size large (total length, 105 to 122; forearm, 43 to 52; longest finger, 77 to 96).

Membranes and ears thick and leathery, the ears distinctly thickened along

anterior border fnscus (p. 96)

Membranes and ears thin, the ears scarcely thickened along anterior border.

Forearm, 47 to 50; longest finger, 85 to 89 (average 86) cubeniiis (p. 102)

Forearm, 50 to 52; longest finger, 85 to 96 (average 90) ..miradorensis (p. 99)

VESPERTILIO FUSCUS Beanvois. Brown Bat.

1796. Vespei-liUo fnsGun Beanvois, Catal. Feale's Mnseuni, p. 14. (Philadelphia, Pa.).
1806. Vespertilio carolinensis GeofiVoy, Ann. Mns. d'Hist. Nat., Paris, VllI, p. 193.

(Carolina.)
1818. Fesprrtilio phaiops liafiuesixne, Am. Monthly Mag., Ill, p. 445. (Kentueky.)
1820. Eplesiciis mclanops Rafinesqne, Annals of Natnrc, p. 2. (Kentucky.)
1823. J^'spcrtilio arquatus Say, Long's Expedition to Rocky Mountains, I, p. 167,

footnote.
1835. Vespertilio MJ'si»MS Teinmincl\, Monogr. de Manuualogie, II, \}. 235.
1843. Scotophihis greenii Gray, List Spec. Mamm. Brit. Mns., p. 30 (nomen nudum).
1864. ScoiopIiih(sfit.scii>i H. Allen, Monogr. N. A. Bats, p. 208.
1878. Vesper It I/O serotinus var. Vesper us fiiscus Dobson, Catal. Chiroptera Brit. Mus.,

p. 193.
1893. AdeJonijcteris fmcus H. Allen, Monogr. Bats N. A., ]}. 112.

Type locality. — Philadelphia, Pa.

Geographic distribution. — Austral, Transition, and (lower edge of)
Boreal zones throughout the United States and adjoining British
provinces.

General characters.— ^izii large; total length, 110 to 112; tail verte-
bra, 41 to 52; forearm, 43 to 40; longest finger, 77 to 84; ear, 11.0 to 14;

VESPERTILIO FUSCUS.

97

ears and membraues thick and leatliery; crowns of upper molars nar-
row; color variable, but seldom very dark.

Ears. — Ears short, reaching barely to nostril when laid forward,
furred on basal third above and sprinkled with hairs on most of inner
surface, but especially near an terior bor-
der. The membrane of the ear is heavier
and more leathery than in the southern
subspecies, and the anterior edge is dis-
tinctly thickened.

Membranes. — Membraues naked, broad
and ample, that of wings attached to
foot a little beyond base of toes. Free
edge of interfemoral membrane a little
shorter than calcar and terminating at
baseof penultimate caudal A'ertebra. The
flight membranes, like the ears, are
thicker and less membranaceous tlian in
the subpecies occurring in or near the
tropics.

Feet. — Foot about half length of tibia;
calcar slightly longer than foot, keeled

on outer
edge, and
terminat-
ing indis-
tinctly or
in a faintly

defined lobe. Dorsum of toes with a
few short bristle-like hairs.

Fur and color. — On middle of back the
fur is about 1-5 mm. long. The fur ex-
tends along the sides in a line about 10
mm. wide on wing membranes both above
and below. The proximal third or fourth
of uropatagium is furred. Otherwise
the membranes are naked except for a
few scattered hairs on the under side of
the interfemoral membrane and on the
under side of the wings close to the lui-
merus and forearm.

Color brown throughout, but always
paler on belly than on back. The exact
shade varies considerably, but is usually
a clear bister or sepia. Sometimes, how-
Ears and membranes blackish in dry

Fig. 24. — Top view of skull of («) Yes23er-
tilio bakamensig, (&) Y. fuaeus, auil (c)
V. serotinus (X2).

Fig. 25. — Side view of skull of (a) Tesper-
tilio bahainensis, {b) V. fuscvs. and (c)
y. serotinus (^ 2).

ever, it approaches cinnamon,
specimens.

Sl-uU.—The skull of typical VesperliUo f uncus (figs. 24 Z>, l'5 h) averages
2772— is^o. 13 -7

98 NORTH AMERICAN FAUNA,

about 1S.5 intii. in occipito-iiasai length and 12.5 mm. in zygomatic
breadth; maudible, 14 mm. It has no tangible characters to distin-
guish it from the skulls of the other large subspecies.

Teeth. — The teeth of tyi)ical Vespertiliofu.scus (tig. 20 a) do not differ
appreciably in form or size from those of the other large continental
subspecies. They average slightly smaller, however, than in V.fu.seiis
miradorcnsis, and the crown of the mi(hlle upper molar is usually
narrower.

Measurements. — See table, page 103.
> Specimens excDiiined. — Total number, 330, from the following localities :

Alabama: Gieensboio, 1.

Arkansas: Fort Towson, 1.

Arizona: Ajiaclie, 4; Santa Cataliua Mountains, 3 (skins); Chiricabua Monu-
taius, .1 (skin); Gnadalupe Canon, Cocbise County, 2 (skins); Fort Yerde,
2 (1 skin); Fort Huacbuca, 0; Huacbuca Mountains, 4 (skins); New River,
2; San Fraucisco Mountain, 6; Yuma, 1.

Britisb Columbia: Asbcroft, 2 (skins).

California: Bear A^alley, San Bernardino County, 2; Cassel, 2; Cloverdale, 1;
Dulzura, 3 (Iskin); Horse Corral Meadows, Fresno County (altitude, 8,000
feet), 1; Kern Lakes, Nortb Fork Kern River (altitude, 7,000 feet), 1; Keru
River, 6; Kernville, 1; Soutb Fork Kings River. 2 ; Lone Pine, 7; Little Kern
River, 3; Mount Sbasta, 2 (skins ); Mount Wbitney, 2; Nevada City, 6;
Nicasio, 63; Owens Lake, 1; Old Fort Tejon, 1; Pine Yalley, 4 (skins); Poso
Creek, Kern County, 1 (skin) ; Round Valley, 1 ; Raymond, 2; Santa Barbara,
2; Sequoia National Park, 9; Sberwood, 1; Twin Oaks. San Diego County,
3; Tbree Rivers, 1; Tebacbapi, 1; Visalia, 2; Walker Basin, Keru County,
4; Yosemite Yalley, 2.

Colorado: Loveland, 6 (skins. Miller coll.).

Connecticut : Norfolk, 2.

District of Columbia: Wasbington, 53 (33 skins).

Georgia: Riceboro, 1.

Idabo: Fort Sberman. 1.

Illinois: Ricbland County, 1; Warsaw, 4.

Kansas: Fort Riley, 2; Neosbo Falls. 1 (skin).

Maine: Eastport, 4.

Massacbusetts: Cambridge, 4; Wilmington,!) (skins).

Mississippi : Bay St. Louis, 2.

Missouri: Marble Cave, Stone County, 5; St. Louis, 1.

Montana: Big Snowy Mountains. 1 ; Prospect Creek, 2; Kalispell. 2; Milk
River, 1.

Nevada: Pyramid Lake, 4; Carson Yalley. 1.

New Hampsbire: Cbarlestown, 1.

New York: Hammondville, 6; Sing Sing. 13.

Ontario: Toronto, 1 (skin).

Oregon: Anna Creek, 3; Des Cbutes River, 4 (skins); Fort Klamatb, 2.

Pennsylvani;i : Carlisle, 1; Center County, 2 (skins).

Soutb Dakota: Smitbville, 5; Custer, 1; Cheyenne River. 1; Fort Pierre,!;
Fort Meade, 1.

Texas: Brazos River, 1.

Utali: Cacbe County, 1; Laketown, 1; Ogden, 5; St. George, 4.

Wasbington : Spokane Bridge, 2; Geyser Basin, 1.

General remarl^s. — In size and general appearance typical Vesper-
tilio fuscKs occupies a somewhat intermediate position among the Xorth
American subspecies. It is considerably smaller than miradorensis

VESPERTILIO FUSCUS MIRADORENSIS. 99

and luucli larger tbauj;roj>^;i(/«^^s' and hahainensis. Very i)allid speci-
mens are occasionally taken in tlie Southwestern United States, but
the luiiiiber of skins available for comparison is so small that it is
impossible to determine the status of the form which these aberrant
individuals represent.

YespcrtUlo fuscus and Y. serotinus have been considered by many
writers as races of a circumpolar species. Six specimens of the sero-
tiue — four from Budapest, Hungary, and two from Berne, Switzerland
— kindly sent me by Mr. Oldfleld Thomas, of the British Museum, prove
conclusively that this view of the relationship of the two animals is
untenable. The differences between the American and Europeau forms
are so great that, taken in connection with the complete geographic
isolation which undoubtedly exists, they leave no doubt of the neces-
sity of recognizing each as a distinct species. Yespertilio serotinus is
a large and heavily built animal, approached in size by T". fuscus mira-
dorensis alone among- the races of Y. fuscus. The adult females from
Budapest measure, respectively: Total length, 131 and 131; tail ver-
tebra-, 52 and 53; tibia, 22.8 and 22.6; foot, 10 and 11; forearm, 52
in each; thumb, 8.1 and 8; longest finger, 93 and 96; ear from
meatus, 20 in each; width of ear, 13 and 11; tragus, 9 and 8.6. In
addition to its large general size Y. ser'otinus has relatively much larger
skull and teeth than any of the races of T^ fuscus {see figs. 24, 25, and
2G). The skull of an adult female from Budapest (Xo. 4489, Miller coll.)
measures: Occipito nasal length, 21.4; zygomatic breadth, 15; man-
dible, 17; upper tooth row (exclusive of incisors), 8; lower tooth row,
10. The skull is considerably broader in proportion to its length than
in Y. fuseusj and the audita! bulla^ are relatively smaller. The teeth
are much larger than those of Yespertilio fuscus, and the inner lobes of
the upper molars are broader, in this respect approaching T". fuscus
ciihensis. The upper incisors arc separated from the canines by a wider
space than in fuscus, and tin's space subtends a distinct groove on-the
surface of the premaxilla between the roots of the canine and incisors.
The paroccii)ital processes are much more strongly developed in T'.
serotinus than in any of the races of T". fuscus.

VESPERTILIO FUSCUS MIRADORENSIS (H. Allen).
1866, Scotophihts miradorensis 11. Allen, Proc. Acad. Nat. Sci., Phila., p. 287.

Ty2)e locality. — Mirador, Vera Cruz, Mexico. Tyi^e in the United
States i^ational Museum, but now mislaid or lost.

Geographic distrihution. — Costa Eica, Guatemala, and southern Mex-
ico. Limits of range not known.

General characters. — Size larger and color darker than in the more
northern form. Feet and distribution of far as in true fuscus ; ears
and membranes thinner and more membranaceous.

Color. — In color Yespertilio fuscus miradorensis averages darker than
tviie fuscus, thus agreeing with the other southern forms, propinquus
and cuhensis.

100 NORTH AMERICAN FAUNA.

Skidl. — The skull of Vcs2)eftiIio/i(scHs mirtidoycnsis is slightly larger
aiul souiewliat less flattened tliau that of true /hschs. The skull of au
adult male from Tehuacau, Puebla, measures: Occipito-uasal length,
19.5 mm.; zygomatic breadth, l."5; mandible, 14.."). The occiput, although
developing even more strongly marked ridges than in the tyi^ical sub-
species, appears less sharply 'peaked' behind when viewed from the side.

Teeth. — The teeth are heavier than in true fuscm, and the crown of
the middle upper molar is broader on its inner side, but no tangible
dental characters can be established to separate the large subspecies.

MecLsiwements. — See table, page 103.

Specimens examined. — Total number, 17, from the following localities:

Costa Rica: San Josp, 1.

Guateoiala: Zuuil, Qnezalteuango, 1.

Mexico: Valley of Toliica, 2 (skius); Ixtapalapa, 2.

Oaxaca: Cerro San Felipe, 1; Oaxaca, 1.

Puebla: Tehuacau, 3 (1 skin).

Tlaxcala: Mt. Maliucbe, 1.

Vera Cruz: Jico, 1; Las Vigas, 2; Tuxpaugo, 2 (skins).

VESPERTILIO FUSCUS PROPINQUUS (Peters).

1872. Vespents projniiquiis Peters, Monatsber. K. Preuss. Akad. Wiss., Berlin, p. 2(32.
1878. I'esperiKjo propinqami Dobsou, Catal. Chiroptera Brit. Mus., p. 203.

Type loi-aUty. — Santa Ysabel, Cruatemala.

Geographic distribution. — In addition to the type the only known
specimen of Vespertilio fuscus propinquus is from Greytown, Nicaragua.

Goteral characters. — Size very small (total length, 90 to 103; tail
vertebrae, 37 to 45; longest finger, 08; ear, 14 to 15) ; breadth of muzzle
distinctly more than half length of head; colors dark.

Ears. — The ears in Vespertilio fuscus propinquus are proportionally
shorter and broader than in typical /h«c».s;, and the tips are distinctly
more broadly rounded. They are haired in exactly the same manner
as in tv\x& fuscus. The ear membranes are thin and membranaceous,
like those of the other southern races.

Memhraues and feet. — Except for their smaller size, the membranes
and feet are exactly as in trne fuscus, though the membranes, like the
ears, are thinner and less leathery.

Ftcr and color. — The fur is shorter than in true fuscus, averaging only
about 7 mm. on back. There is nothing peculiar in its distribution.
In the single alcoholic specimen that I have seen the color is about as
in V. fuscus miradorensis. Peters, however, describes the color of the
type specimen as rust red.' This is much brighter than the Greytown
specimen, but the color may be due to staining while in alcohol.

Measurements. — See table, page 103.

Spccitucns examined. — I have seen onl}' one specimen of Vespertilio
fuscus 2)r(q)inq)( us. This Avas collected by Mr. Charles W. Eichmond at
Greytown, Nicaragua ( 9 ad., Xo. 52790, U. S. National Museum,
Dept. of Agriculture collection).

' "Oben rostroth, dieHaare an der Basis sch\varz';)raun, Bauchseite blasser, indeui
die au der Basis srhwarzbrauuen Haarc bier iiiebr rostnelbe Spitzen baben." '

VESPEKTILIO FUSCUS BAHAMENSIS,

101

General remarls, — Among the races of Vcspertilio /uncus, V. fuscus
lirophiquus differs most widelj- in size and in form of bead from its
nearest geograpliical ally, Y. fuscus miradoreusis. It combines the
small size of the West Indian bahamensisy the broad muzzle of true
fuscus, aud the delicate ears and membranes of tlie southern races in
general. Additional material may show that it is speciflcally distinct.

VESPEETILIO FUSCUS BAHAMENSIS siibsp. nov.

Type from Nassau, New Providence, Bahamas. Adnlt cj (iu alcohol) No. 76537, U. S.
National Musetnn (Biological Survey collection). Collected iu tlie spring of 1894
by C. J. Maynard.

Geocirapliic (listrihuiion. — This form is known from the type locality
only.

General characters. — Size about as in V. fuscus propinquus ; breadth

Fio. 26. — Teeth of (a) Ycspcrfilio fxiseiis, (h) Y. bahamensis, (c) T. cube7uis, (d) T. itiimdorensis, and

(e) V. serotinus (x5).

of muzzle less than half length of head; ears narrower than in propin-
(pius, about as in typical /'/r.s■c/^s•.

Ears. — Ears smaller than in tyincal fuscus, but of essentially the
same shape, thus narrower than in propinquus. The ear membrane is
thinner and more membranaceous than in true fuscus. in this respect
resembling that of the other southern races.

Membranes and feet. — The membranes and feet are as in typical
fuscus, allowance being made for the smaller size of bahamensis, and
the difference in texture of the membranes common to all the southern
races.

Fur and color. — The fur is shorter than in true fuscus, averaging only
about 8 mm. in length on the back. It is distributed exactly as in the
typical subspecies. So far as can l)e determined from specimens pre-
served in alcohol, the color is considerably darker aud duller than in
true fuscus.

102 NORTH AMERICAN FAUNA.

Skull. — The skull of Vespertilio fuscns baliamensis is much smaller
and more lightly built than that of typical fuscus (figs, 'l^a and 2oa).
The skull of a fully adult male measures: Occipito nasal length, 10.5;
zygomatic breadth, 11 ; mandible, 12.G. In form it differs from that of
true fuscns in its narrower, deeper, more cylindric brain case and less
sharply 'peaked' occiput.

Teeth. — The teeth, like the skull, are smaller than those of tTiie fnscits
(flg. 26h). In a fully adult male the upper tooth vow (exclusive of
incisors) measures G.4 mm.; the mandibular tooth row, 7.8 mm. Id
form the teeth differ slightly from those of true fuscus in the greater
breadth of the inner (lingual) side of the first and second upper molars.

2Ieasurements. — See table, page 103.

Specimens examined. — Total number 90, all from the type locality.

General remarks. — Vespertilio fuscus haliamensis needs no comparison
with typical fuscus or with Y. fuscus cuhensis, from both of which it
differs widely in size. Its superficial resemblance to V. fuscus pro-
pinquns is closer. Unlike the latter, it has a very narrow muzzle.

VESPERTILIO FUSCUS CUBENSIS (Gray).

1839. Scotojiliihis cuhensis Gray, Auu. Nat. Hist., IV, p. 7.

1840. Vespertilio diiterireits Gervais, iu Ramon de la Sagra's Hist, de I'lle de (.'aba,

Maminiferes, p. 6.
1892. Fesperiujo fuscus cubensis Chapinau, Bull. Am. Mas. Nat. Hist., IV, p. 3ir>.

Type locality. — Cuba.

Geographic distribution. — Cuba.

General characters. — Externally similar to Vespertilio fuscus mirado-
rensis, but slightly smaller in general size, and with much smaller ears.
Skull about as large as in true fuscus, thus much smaller than iu
miradorensis.

Ears. — The ears are delicate and papery, like those of the other
southern races. They are smaller than in either fuscus proper or mira-
dorensis. In form they differ nuxrkedly from those of true fuscus in
their general narrowness, and especially in their more pointed tips.
The characters of the ears have already been described by Mr. Chapman.

Memhranes. — In form the membranes do not differ from those of the
other subspecies. In texture they agree with the southern forms.

Fur and color. — The fur is distributed exactly as in the other sub-
species. In color the specimens, after five years' immersion in alcohol,
are darker and redder, especially on the whole ventral surface, than any
others that I have seen. They even surpass V. fuscus miradorensis in
darkness and richness of color.

Skull. — The skull of Vespertilio fuscus cuhensis is about the size of that
of \vn& fuscus or a little smaller, thus distinctly smaller than that of T'.
fuscus miradorensis, the form to which cuhensis bears the closest super-
ficial resemblance, and much larger than that of hahamensis, its nearest
geographical ally. In form the skull is similar to that of true fuscus

VESPERTILIO FUSCUS CUBENSIS.

103

but the braiu case is slightly less flattened. The sagittal crest is well
developed as in the other large subspecies.

Teeth. — The teeth of Vesperiilio fuscus cubcnsls (fig. 26 c) diiier from
those of true fuscus in the greater breadth of the inner (lingual) sides
of the maxillary molars. These teeth are also distinctly shortened in
their transverse diameter. These peculiarities are exaggerations of
the conditions found in miradorensis and haliamensis.

Measurements. — See table below.

Specimens examined. — Total number, 11, from the following locality:
Cuba: Trinidad. 10'; , 1.

General remarls. — Vespertilio fuscus cuhensis is a fairly well marked
insular form apparently most closely related to V. fuscus miradorensis
of southern Mexico. It differs much less from this large continental sub-
species than from V. fuscus haliamensis, its nearest geographical ally.

Averarje iiieasnrcnienfs of subspecies of resperiilio fuscus.

Subspecies

fURCHS

prnpiiiijuus
baliarinntiis
cuhensis

Massachusetts : Cambridge

Xew York : Sing Sing 10

District of Columbia : 5

Washington.
Mississippi: Bay St. Louis., j 2

California : Xicasio 10

Lone Pine I 5

Vera Cruz: Mirador ^\

Jico 1

Las Vigas 1

Tiaxcala: Mt. Malinche i 1

Piiebla: Tehuacau 2

Guatemala: ZnDil i 1

Costa Rica: San Jose [ 1

Guatemala: Sta. Tsabel j^l

Nicaragua: Greytown 1

In ew Providence : Xassaa . . . ' 1

Nassau... 10

Cuba : Trinidad 10

116 '47. .5
113. 9|44. 3
110. 8!46. 5

108. 5 42
113 '47

108. 6 46
57

118 48
120 50
120 50

119 50.
110 ...
118 49
105 45

96 37
103 44

101. 7 42.
110.7148.

.

/2

g

H

1^

tH

la 2

h^ iw

19.5,10.4 45 I 6.8 81
19. i; 9.7 45.3

19. l! 9. 7144. 8

19. 5 10
19.7 9.
19.6 9.
.... 10.

■22 10
22 10
21.611.

5*20. 7| 9.

.'20 110.

22 !lO

:18 10

17 j 7.

18 I 8
6 17. 9' 8.
519.9 9

:47

6 46. 4

244. 8

6 50

50

I

51

4 52

8 50. 5
4.50

52

40
840

42
6 42. 7

'48.4

18
7 J80 19.
6. 5 80. 5I18.

6.7 79.517.

6 82 !l7.
6. 179.4'l7
8.3'. ...I...

8 91 19.

7 85 |l8

6. 4 96 jl9
7 |90 19
6. 4.... 18.
0. 8 93 '20

9.5 .... 15

5 |08 '14

6 '77 16.
6 [74 15,
6. 6'86 ;i6.

13.3 8.3

12.7! 8.3
12. 81 8

12.5^ 7.8
12.7, 8.1
12.4 8.1
3

13

13.6

13

13.3
6112

13.6

11.4|)

11 7.8
811 Is
6 11. 3' 7.6
712.6, 8.4

For the opiiortnnityofexaminingthe.se .specimens I am indebted to Dr. J. A. Allen

of the American iluseniu of Natural Hi.story.
-Type.

104 NORTH AMERICAN FAUNA.

[Note. — The following specifs is not rt-presfuttMl in any of the extensive collections
of l)at.s recently in.ide in Mexico. As I have never seen the animal aiul hence can
form no opinion as to the weigiit of its cliaracters, I have not attempted to include
it in the synopsis of the North American forms of fesperlilio. Dobsou's description,
based on an examination of the type, may be iutrodnced here, however, as an aid
to the recognition of the species.]

VESPERTILIU AL15IGULAKIS (Peters).
1872. J'(S2>ertts (Alarsipohrmus) albujithirin Peters, Monatsber. K. Akad. Wiss., Uerlin,

p. 260.
187i^. VciiperiKjo albitjiilaris Dobson, Catal. C'hiroptera Brit. Mns., ^i- 207.

"Ears very broad and broadly rounded oft' above; the lower Lalf of
the outer margin of the ear couch broadlj' folded backwards, as in T.
novtuUii separated in front from the angle of the mouth by a Avart, hut
terminating heloic and intefnal to it under the lower jan- by a small inter-
nal prolongation; tragus broad above, attaining its greatest width above
the middle of the inner margin, which is slightly concave, narrowe.st
opposite the base of the inner margin, a prominent triangular lobe at
the base of the outer margin. oSTostrils rather wide a])art, opening sub-
laterally ; muzzle broad and obtuse ; crown of the head scarcely elevated
above the face-line.

•'Wings from the base of the toes; postcalcancal lobe long and nar-
row; last caudal vertebra free.

"Fur dark brown above, the extreme tips hoary, as in V. noctivagans^
paler beneath, the chin and throat, as far back as a line connecting the
posterior margins of the ears, pure irhite.

"Upper inner incisors long and broad and slightly bifid at their
extremities; outer incisors very short, shortly exceeding the cingulum of
the inner ones in vertical extent; the single upper premolar close to the
canine; lower incisors in the direction of the Jaws; first lower premolar
half the size of the second, which exceeds the molars in vertical extent.

"Length (of the type specimen, an adult S ), head and body 2".35
[59.7 mm], tail V'.o [38 mmj, head 0".l [17.8 mm], ear 0".65 x 0".13
[10.5 mm X 5;8 mm], forearm 1'''.05 [11.9 mm], thumb 0".35 [8.9 mm],
third finger 2".75 [09.8 mm], fifth finger 2" [50.8 mm], tibia 0".6 [15.2 mm],
footO".35 [8.9 mm].

''Hah. — Mexico. Type in the collection of the Berlin Museum.

"This species may be at once distinguished from all other species of
Y<spertilionidw by the very peculiar manner in which the outer margin
of the ear-conch terminates under the jaw, which has caused tlie
describer, Dr. Peters, to make it the type of a new subgenus, Marsipo-
la'inus. In the prolongation of the ear conch, in the form of the tragus,
and in dentition it resembles the African species of Chalinolohus.'"

General remarks. — This species is very difterent from any of those
recently collected in Mexico, and is probably well worth}' of subgeueric
or even generic separation from Vespertilio. Its characters are so
remarkable and Mexico has recently been so thoroughly explored that
doubt is thrown on the accuracy of Peters' information concerning the
type locality.

LASIURUS. 105

Genus LASIURUS Gray.

1831. Ldsiurus Gray, Zoological Miscellany, No. 1, p. 38 ( based ou the American hairy-
tailed bats).

1861. Lasiunis H. Allen. Monogr. X. Am. Bats, p. 14.

1,^70. Aialapha Peters, Monatslier. K. Akad. Wiss., Berlin, p. 907. (Xot Atalapliu Rati-
nesqne, 1814.)

1878. Aialapha Dobson. Catal. Chiroptera Brit. Mns.. p. 2(37. CSotAtalaplia Ratinosqne,
1814.)

18i;'3. AialaiihaU. Allen. Monogr. BatsN. Am., p. 141. (Not J/«/((j)/<a Ratinesijne, 1814.)

Type species. — Lasinrus horcalis (Miiller .

Geof/raphic distribution of genus. — The wliole of Nortli America and
South America, the West Indies, Sandwich Islands, and Galapagos
Islands.

1-1 1-1 2-2 3-3

Generic characters. — Dental lormula: r, c,—,] c, ^. ; prtn., ry^; m, o-o=32;

upper incisor in contact with canine; a minute upper premolar at base of
canine on inner (lingual) side ; dental formula otherwise as in Dasypterus^
Nycticeius, and Ixhofjevssa ; skull (ligs. 28, 29,31) broad, short and deep,
very different in form from that of any other North American genus
of VespertiUonidcc except Dasypterus; ear (fig. 27) broad, blunt, and
rounded at tip, hairy on most of dorsal surface; dorsal surface of inter-
femoral membrane furred nearly to extreme edge; mamma', 1.

The members of the genus Lasiurris are recognizable among Xorth
American bats by their thickly furred interfemoral membranes. Two
distinct species are known to occur north of Paimma; one of these is
divisible into at least five well-marked geographic races.

KEY TO XOKTH AMERICAX FOEilS OF LASIUEFS.

Size large (forearm more than .50 mm.) ciiieniis ip. 112)

Size small (forearm 36-44).

Underside of wing membrane very sparsely haired along forearm .?»if.r/crt?i us (p. Ill)
Underside of wing membrane thickly furred immediately back of forearm.

Ear small and Avith slightly develoiied external basal lobe telioiis (p. 110)

Ear large and with well-developed external basal lobe.

Color mahogany brown seminolus (p. 109)

Color varying from deep rich cherrj' red through orange and
yellow to light yellowish gray.

Color deep rich cherry red, forearm 42-44 pfelfferi (p. 110)

Color varying from yellowish gray to light red, forearm
36-43 borealis (}). 10-5)

LASIURUS BOREALIS (Miiller). Red Bat.

1776. J'esjycri'dio horealis Miiller, Xatursyst. Snppl., p. 21.

1777. Vespertilio novehoracensis Erxleben, Syst. Regui Anim., I, p. l')~^.

1781. VespertUio lasinrus Schreber, Siingthiere, Abth. I. Taf. LXII B (published with

Abth. IV Heft 4. See Sherborn, Proc. Zool. Soc. London. 1891, p. 589).
1785. Vespertilio noreboraciis Boddaert, Elenchus Auimalinm, I, p. 71.
1785. VesperfiUo lasnrus Boddaert, Elenchus Animaliura, I, p. 71.
1796. Vespa-tUio rubellus Palisot de Beauvois, Catal. Peale's Museum, p. 204.

106 NORTH AMERICAN FAUNA.

1814. Jtalapha americana Ratiuesque, Pr(lcis des ddcouv. soniiol., p. V2 (nomeu

nudum).'

1815. Vespertilio 7'iihra Ord, Gutlirie's Geography, 2d Am. ed., II, p. 291.
1818. Vespertilio tesselatits Ratiuesque, American Monthly Mag., Ill, p. 44.5.
1818. Vespertilio monaclms Raiinesque, Am. Monthly Mag., Ill, p. 445.

1820. Vesperiilio rufus Warden, Description des I5tat8-Unis de I'Amdrique Septentrio-

uale, V, p. 606.
1863. Lasiurus norehoracensis H. Allen, Monogr. N. Am. Bats, p. 15.
1870. Lasiurus funehris Fitzinger, Sitzungsber. k. Akad. Wissensch. Wien, LXII, p. 46.
1878. Atalaplia noveboraceusis Dobson, Catal. Chiroptera Brit. Mus., p. 269.

1893. Atalapha uovehoraccusis H. Allen, Monogr. Bats N. Am., p. 142.

1894. Atalapha horeaVis Rhoads, American Naturalist, XXVIII, p. 523.

Tifpe locality. — Xew York.

Geographic distribution. — The typical form of Lasiurus borealis ranges
tbrongh the' Boreal, Transitioii, and Anstral zones in eastern Xortli
America from Canada to Florida and Texas, west at least to Indian
Territory and Colorado. Southern and western limits of range not
known. Probably breeds throughout its known range.

General characters. — Size small (forearm, 38 to 43; longest finger, 78
to 88); forearm with no distinct tuft of far
near proximal end; color very variable, rang-
ing from bright yellowi.sh red or fawn color to
yellowish gray; a whitish area in front of
shoulder.
Ears. — The ears of ty])ical Lasiurus borealis
'~? .#^ Cb ^ tJ?'^/ (^^* ^^^') when laid forward reach a little
V ^^^ more than halfway from angle of mouth to

Fig. 27. — Ear of (a) Zn«iMn<s 6ow- j_ -i mi i. • i j • i. i i i.

au^anao,)L.M!otu^x2). I'ostril. The autcrior border is strongly but
irregularly convex from free point of anterior
basal lobe to tip, a distance through which it forms almost a semicircle.
The posterior border is slightly concave immediately below tip, then
evenly convex to basal lobe. The convex portion of the outline of the
posterior border forms the arc of a circle with considerably longer radius
than that of the anterior border. Posterior basal lobe strongly devel-
oped and deeply notched on anterior border. Inner side of ear naked
except for a few scattered hairs, which are especially numerous along
anterior and ijosterior borders. Outer side densely furred throughout
basal two-thirds, naked at tip.

Tragus triangular in general outline. Anterior border straight from
base to slight concavity just below tip; posterior border straight from
tip to widest point (opposite anterior base), where there is a strong
angle, below which the margin is straight to slightly developed basal
lobe.

'Raiinesque says: "J'ai observe cette espt'ce [A. sicula Eaf.] en .Sicile, eiledift't>re
de V Atalapha americana ( Vespertilio noveboracensis Lin.), autre espf'ce du memo geure,
par ses deux premier's et son dernier caractere."

LASIURUS BOREALIS. 107

Membranes. — The flight membranes are attached at base of toes, the
iu'Oi)ataginm at extreme tip of tail.

Feet. — The foot is small, less than half as long- as tibia. Dorsal snr-
face of toes thickly fnrred. Calcar about twice as long as foot and con-
siderably shorter than free border of iuterfemoral membrane. It is
slightly developed, indistinctly keeled, and seldom lobed at tip.

Fur and color. — The fur is everywhere full and soft. On middle of
back it is about 7 mm. in length and on neck about 10 mm. It covers
the basal two-thirds of dorsal side of ear, the whole dorsal side of the
iuterfemoral membrane, and the dorsal side of the flight membrane to a
line running from ankle to middle of humerus. There is a narrow strip
of fur running along basal third of lifth metacarijal and a squarish
clump at base of thumb. Xear base of forearm (in position occupied
by strip of. fur in L. cinereus) there are numerous fine scattered hairs,
which are so inconspicuous as readily to escape notice. On the ventral
surface the fur reaches about to middle of uropatagium and on flight
membranes to line joining knee and elbow. Beyond elbow a sparse
growth of hairs covering an area 10 mm. or more in width extends
along forearm to bases of fingers, where it
becomes much more dense. The ante-
brachial membrane is covered with a sparse
coating of hairs on the ventral surface.

In color typical Lasii<rus boreaJis varies
verj' extensively, but never shows the ma-
hogany brown of seminolusov the intense red
of the tropical races. Red specimens are ru-
fous red throughout (the exact shade some- ^'»- 28.-TorTiew of sknii of (a) Last-

, , , . ufux fjorcaUs and (b) L.teliijtis{ 2).

Where between rutous and burnt sienna),

paler and more fawn-colored on the belly, the hairs of the back usually
with distinct grayish tips, those on the throat and chest tipped with
whitish. A yellowish white patch in front of each shoulder. Frequently
the white on chest tends to connect the shoulder patches by a Avhitish
collar. The individual hairs on the back are blackish at base, then
light rufous to the narrow subapical baud which gives the characteris-
tic color to the back, and, finally, grayish white at extreme tips. Gray
specimens are yellowish gray on the back and bufi'y on the belly. The
red usually ijersists as a faint salmon suffusion.

Sk-iiJI. — The skull of typical Lasiurus borealis (figs. 28 a, 29 b) has
the broad rostrum and fiaring zygomata of L. cinereus. The dorsal
profile of the skull is nearly straight from external uares to highest
point of occiput. The skull of an adult female from AVashington. D. C,
measures: Greatest length, 13.8; zygomatic breadth, 10.2; breadth of
rostrum at jjosterior edge of large premolar, 0; mandible, 10; upper
tooth row, 5.4: ; lower tooth row, 0.4.

Teeth. — The teeth (fig. 30 b) are large, the upper molars broad on the

108 NORTH AMERICAN FAUNA.

inner (lingual) side, and the lower molars wide in tlieir transverse
diameter.

Measurements. — See table, page 115.

Sx^ecimens examined. — Total number, 387, from the following localities:

Alabama: Mobile Bay, 3; Greensboro, 9 (2 skins, Merriam coll.).

Arkansas: Fort Smitb, 1.

District of Columbia: Washington, -20 (6 skins).

Florida: Old Town, 1 (skiu, Miller coll.); St. Marys, 1.

Georgia: Ricel)oro, S.

Illinois: Mount Carmel, 1 (skin); Gluey, 3; Warsaw, 2; West NortUfield, 1
(skin).

Indian Territory : Hartshorue, 1 (skiu); Redland, 3.

Kansas: Cedar Yale, 1.

Kentucky : Hickman, 2.

Louisiana: Lafayette, 1; Nevr Orleans, 2; Pineville, 1; Shreveport, 1.

Massachusetts: Nantucket,^ 1.

Mississippi: Hancock County, 1 (skin, Miller coll.); Washington, 10.

Missouri: Golden City, 1; Marble Cave, Stone County, 1.

New York: Greene, 1; Hartford, 1 (skin) ; Oyster Bay, 1 (skin) ; SingSing. 86.

North Carolina: Fort Macon, 1 (skin); Magnetic City, 1 (skin); Roan Moun-
tain, 1 (skin).

Oklahoma: Ponca, 2.

Ontario: North Bay, 2 (Miller coll.); Toronto, 1 (skin).

Pennsylvania: Bainbridge, 1; Carlisle, 1 (skin); Kenneth Square, Chester
County, 1.

South Carolina: Mount Pleasant (near Charleston), 7 (skins, Miller coll.)

Tamaulipas: Matamoras, 2.

Tennessee: Alexandria, 1; Arlington, 2; Big Sandy, 11; Clarksville, 1 ; Dan-
ville, 1.

Texas: Arthur, 3; Brownsville, 1.58 (3 skins) ; Clarksville, 1; Corpus Christi, 2;
Fort Clark, 1 ; Nueces Bay, 1 ; Paris, 3 ; Waco, 1 (skin. Miller coll. ) ; Wichita
Falls, 1.

Virginia: Amelia Court-House, 1; Berryville, 1 (skiu); Dismal Swamp, 2;
Gainesville, 1.

General remarks. — Typical Lasiitrus horeaUs presents a wide range
of individual Yariation, but may always be distinguished from the
other subspecies by fairly constant charaeters. It never shows the
mahogany brown coloring of L.horealis seminolus and seldom approaches
the brilliant cherry red of L. borealis pfeif'eri. From L. boreal is teliotis
it differs in its larger ear with well-developed, strongly-notched external
basal lobe.

Specimens from Brownsville, Tex., where the animal breeds, are
slightly smaller than those from New York, and the ear is propor-
tionally shorter (see table of measurements, p. 115). In this respect
they are intermediate between true borealis and teliotis, though the ear
is formed exactly as in the tyjncal subspecies. Specimens from Okla-
homa and Indian Territory, on the other hand, are indistinguishable
from northeastern specimens.

LASIURUS BOREALIS SEMINOLUS. 109

LA8IUKUS BOKEALIS SEMINOLUS (Rhoads).
1895. Atalapha horealis scminola Khosuls, Proc. Acud. Nat. Sci. Pliila., ji. 32.

Tij2)t' locality. — Tarpon Springs, Fla,

(rciif/raphic (listrihiiUoii. — Lower Austral and Tropical zones from
South Carolina to southern Texas,

(ieneral cJun-acter.s. — In size and jiroportions similar to typical Lasi-
Hfus boreaiis; general color mahogany brown, slightly frosted with
grayish.

UarSj memhranes, feet, and distribution of fur. — In all external char-
acters except color Lasinrus horealls seminolus agrees with typical
horealis.

Cidor. — General color rich mahogany brown throughout, the back
(especially between the shoulders) slightly frosted Avith gray and the
throat and chest varied with whitish. A distinct whitish area in
front of shoulder as in true horealls. Muzzle, backs of ears, and fur
bordering forearm, yellowish brown. CIumiD of fur at base of thumb
whitish or yellowish. On middle of back the fur is about 12 mm. in
length. In this region the colors on the individual hairs are arranged
iu four bauds as follows: Basal band deep blackii^h plumbeous (this
band usually broader than in true borealis), middle band light gray,
siibapical band rich mahogany, extreme tip grayish white.

Color variation in Laslurus borealis seminolus is much less than in
typical borealis, and is chietly noticeable in the amount of red in the
mahogany brown, in the amount of white on the throat and chest, and
in the shade (»f gray in the broad middle band on the hairs of the
back. This is often strongly suffused with yellowish.

Skull and teeth. — As in typical borealis.

Measurements. — See table, page 115.

Hpeeimens examined. — Total number, 19, from the following localities:

Florida: Old To%Tn, 3 (skius. Miller coll.); Lake Haraey, 2.

Georgia : Nashville, 1.

Louisiana: New Orleans, 5.

Mississippi : Bay St. Louis, 3.

Sontb Carolina: Mount Pleasant (near Charleston), 4.

Texas: Bi'owusville, 1 .

General remarks. — Lasiurns borealis seminolus appears to be a well-
marked subspecies confined to the Austroriparian fauna. The single
specimen taken at Brownsville, Texas (No. 50970, U. S. National
Museum) was killed on September 8, 1891, and may have been a migrant.
No intermediates between seminolus and true borealis has yet come to
light, but tlie perfect agreenient of the two forms in all charaters
except color nmkes me unwilling to recognize them as species. The pos-
sibility that seminolus and true borealis are dichromatic phases of one
species lacks weight on account of the total absence of intermediate
specimens, and also from the fact that both forms have not yet been
found breeding at any one locality.

110 NORTH AMERICAN FAUNA.

LASIURUS BOREALIS PFEIFFERI (Guiidlacli).

1861. Atalapha pfcifferi Gundlacb, Monatsber. K. Preiiss Akad. Wiss., Berlin, p. 1.52.
1878. Aialapha iiovehoracensls xav. (i (Jtalaplui p/eiffcri) Dobsou, Catal. Cbiroptera
Brit. Mus., p. 271.

1892. Jtala2)lin vorchoracensis pfeifferi ('bapiuau, liull. Am. Mas. Nat. Hist., IV, i). 316.

Type locality. — Cuba.

Geograpluc (listrihution. — Cuba. Jamaica? Bahamas'?

General characters. — Slightly larger than typical Lasiurus horealls^
but similar in iiroportions; color brighter and more intense.

Ears., membrajies, feet, and (listrihution of fur. — As iu the typical
subspecies.

Color. — I have seen no skins of the Cuban red bat, and am therefore
unable to give a detailed description of the animal's color. The two
specimens collected by Mr. Chapman in 1892 have now been in alcohol
for five years. Hence their color furnishes no trustworthy basis for
comparison with that of continental material. When compared with
alcoholic specimens from the eastern United States they are appreci-
ably brighter.

Measurements. — See table, page 115.

Specimens examined, — Two from Trinidad, Cuba (Am. Mus. Nat.
Hist.).

A skull from Nassau, Bahamas (Miller coll.), and an imperfect skin
from Spanish town, Jamaica, may be referable to this race, but it is not
possible to identify them with certaiut3^

General remarlcs. — Lasiurus horealis ifeifferi is a tolerably well-marked
insular form, distinguished from typical horeaUs by its slightly larger
size and brighter color.

LASIURUS BOREALIS TELIOTIS (H. Allen).

1891. Atalapha teliotis H. Allen, Proc. Am. Pbilos. Soc, XXIX, p. 1.

1893. Atalapha teliotis H. Allen, Monogr. Bats N. Am., p. 153.

Type locality. — Unknown, probably some part of California (type in
U. S. National Museum).

Geographic distribution. — This form is known from a few localities iu
California and Lower California from the head of the Sacramento Valley
south to Comondu.

General characters. — Slightly smaller than typical Lasiurus horealis;
ear proportion allj^ much shorter than in the typical subspecies, and
with external basal lobe greatly reduced in size; color averaging
brighter than in the typical form.

Ears. — The ear (fig. 27&) is similar iu form to that of typical horealis,
except that the tip is slightly narrower and the external basal lobe is
reduced in size, indistinctly marked off from the rest of the ear, and
scarcely, if at all, notched on its anterior border.

Mcmhranes, feet, and distribution of fur. — The external form, with the
exception of the size and shape of ears, is as in true horealis.

Color. — I have seen only four skins of the Californian red bat. In

LASIURUS BOREALIS MEXICANUS.

Ill

Fig. 29.— Side vievv^ of skull of
(a) Lasiurus teliotis and {h)
L. borealis (X2).

these tlie color is uniformly slightly darker and redder than in ordinary
red specimens of true honalis. The difference is especially noticeable
on the interfemoral membrane, rump, and lumbar region. One skin
( S ) from Dulzura. Cal., almost lacks the grayish tips to the hairs on
the back. Another (also S ) taken at the same jjlace on the same day
(November 5, 1891) shows the gray tips very
distinctly on the neck and fore part of the back.

Skull. — The skull of Lasiurus borealis teliotis
(figs. 28 &, 29 a) is distinguishable from that of typ-
ical borealis by its smaller size, narrower rostrum,
and less flaring zygomata. That of an adult
male from Dulzura, Cal., measures: Greatest
length, 12.1; zj^gomatic breadth, 9; breadth of
rt)Strum at posterior edce of large premolar, 5.2;
ui)i)er tooth row, 4.(5. TLe mandible of this spec-
imen is lost. That of another adult male from
the same locality measures: Length, 9; lower
tooth row, 5.1.

Teeth. — The teeth are smaller than in the typ-
ical subspecies, the upper molars are narrower on the inner (lingital)
side, and the mandibular teeth are narrower in their transverse diam-
eter (fig. 30 «.)

Measurements. — See table, page 115.

Specimens examined. — Total number, 10, from the following localities:

California: Exact locality unkuoAvn, 1 (type); Bakersfield, 1; Berryessa,
Santa Clara Conuty, 1 (skin); Dnlzura, 2 (skins, Miller coll.): Fresno, 1;
Santa Ysabel, San Diego County, 1; Tehama, 1; Three Rivers, 1.
Lower California: Comondu, 1 (skin).

General remarks. — Lasiurus &ore-
(/Zisfe/io/isisreadily distinguishable
from typical borealis by its smaller
ear, with less developed and entire
external basal lobe, smaller skull,
with narrower rostrum, and weaker
dentition. From X. borealis mexi-
canus it differs in completely furred
dorsum of interfemoral membrane and more hairy under side of wing.

Fir,. 30.— Teeth of (a) Laiiunis teliotis and (6) L
borealis (X5).

LASIURUS BOREALIS MEXICANUS (Saussure).

1861. Aialapha mexicana Saussure, Revue et Mag. de Zool., 2e st^r., XIII, p. 97, Mars.,

1861 (southern Mexico).
1871. Atalaplia frantzii Peters, Monatsber. K. Preuss. Akad. Wiss., Berlin (1870), p.

908, 1871 (Costa Rica).
187X. Aialapha novehoracensis var. a {AtaJapha frantzii) Dobsou, Catal. Chiroptera

Brit. Mus., p. 271.

Type locality. — Xot stated, Init without doubt in some one of the
States of southern Mexico, probably Vera Cruz, Puebla, or (Jaxaca.

112 NORTH AMERICAN FAUNA.

Geof/rapliic. distyibntioii. — Ceutral America and southern Mexico.
Limits of range uuknown.

General characters. — Apparently most like Lasiurus horeaUs tcUotis,
but feet, iuterfemoral membrane, and under side of wings much less
hairy.

Ears. — In dried specimens the ears api^ear to be essentially as in
L. horeaUs teliot is, though the external basal lobe may be slightly more
developed.

Membranes and feet. — These shoAv no distinctive characters.

Fur and color. — On the body the I'ur shows no peculiarities as com-
j)ared with the other subspecies. On the iuterfemoral membrane it
extends thickly to about the middle, then becomes more sparse, and
finally disappears, leaving the edge of the membrane bare. The backs
of tbe feet are scarcely furred. On the under side of the wings, the
area behind the forearm which is densely furred in the other sub-
species, is merely sprinkled with inconspicuous hairs; these are, how-
ever, more dense at the bases of the fingers. The antebrachial mem-
brane is also very sparsely furred.

Color as in L. horeaUs teliotis.

Measurements. — See table, page 115.

Specimens examined. — Total number, 8, from the following localities:

Jalisco : , 6.

Tehuautepec : Guichicovi, 1 (skiuj.

Vera Cruz: Peuuela (uear Cordova), 1 (skin).

General remarlis. — From the unsatisfiictory material at my disposal it
appears that ic/«/«r»s horeaUs mcxicanus is a well-marked race, most
like teliotis, but differing from this, as well as from all the other known
subspecies, m the restricted peripheral distribution of the fur.

LASIURUS CIXEREUS (Beauvois). Hoary bat.

1796. Vesjurtilio linereiis Palisot de Beauvois, Catal. Peale's Museum, Philadelphia,

p. 14. (Obvious misprint for chierens.)
1823. VespertUio pruinosiis Say, Long's Exped. to Rocky Mts., I, p. 167 (footnote).
1864. Lasiurus cinereus H. Allen, Mouogr. N. Am. Bats, p. 21.
1878. AtaJapha cincrca Dobson, Catal. Cbiroptera Brit. Mus., p. 272.
1893. Atalupha ciuerea H. Alleu, Monogr. Bats N. Am., p. 155.

Type locality. — Philadelphia, Pennsylvania.

Geographic distribution. — Boreal Xorth America from Atlantic to
Pacific. The hoary bat breeds within the Boreal zone, but in autumn
and winter it migrates south to the southern border of the United States
and probably, much farther.

General characters. — Size, large (forearm, over 50 mm.); prevailing
color, gray; ears with black rims; forearm with distinct patch of fur
near base.

Ears. — The ears of Lasiurus cinereus are in general similar to those
of L. horeaUs, but are broader in proportion to their length (see table of
measurements, p. 115). The external basal lobe is less developed than

LASIURUS CINEREUS.

113

in borealis and without trace of notch ou anterior border. Margin of
ear membrane dark brown or blackish. Outer side of ear densely furred
to a little beyond middle. Inner side with conspicuous patch of yellow-
ish hairs above and in front of middle and a border of similar hairs along
lower part of anterior edge.

Tragus shaped as in L. borealis, covered with sparse coating of hairs
on outer side.

Membranes. — In form and attachment the membranes are as iu L.
borealis.

Feet. — Foot about half as long as tibia; dorsal side thickly furred.
Calcar twice as long as foot and slightly shorter than free border of
interfemoral membrane. It is distinctly though narrowly keeled on
posterior edge, aud usually lobed at tip. The terminal lobe is very
variable, aud may be well developed ou
one side and absent ou the other.

Fur and color. — The fur is distributed
much as iu L. borealis. As in that species,
it is distinctly longer on neck than ou back,
thus forming a ruff. On the neck it aver-
ages about 15 mm. iu length, ou the back
11 mm. General color, a mixture of light
yellowish brown, deep umber brown, aud
white, the yellowish brown clear and un-
mixed on throat, head, and under side of
membranes, the umber brown predominat-
ing on back and dorsal surface of interfem-
oral membrane, where, however, the hairs
are mostly tipped with silvery white, some-
times to so great an exteut as nearly to
conceal the dark tints beneath. Lips,
chin, aud cheeks sprinkled with short blackish hairs. Ventral surface
with white predominating ou belly, between which and yellow of throat
is a baud iu which the umber brown is more conspicuous than else-
where ou the under parts. Tufts of fur at bases of thumb, fifth finger,
aud forearm, light yellowish brown, like fur on under side of wing
membraues. On middle of back the individual hairs are colored as
follows: Deep plumbeous at base; light yellowish brown (shading
toward umber distally) through middle half; umber brown subapically ;
silvery white at tip.

Color variation is considerable, but never enough to obscure the char-
acters of the species. It appears to be wholly independent of locality,
as skins from such widely separated localities as Minnesota aud south-
ern California are practically indistinguishable.

One skin from the Santee Eiver, South Carolina, has the dusky tints
throughout the pelage so intensified and extended as to suggest melan-
277L*— Ho. 13 8

Fig. 31.

-Skull of Lasiurus cinereus

(X2).

114

NORTH AMERICAN FAUNA.

Fio. 32. — Teeth of Lamnrus cine mis ( 5).

ism. Another from Eureka, (Jalifornia, is iu a similar phase, though
not so extreme.

Slcull. — The skull (fi!ii. 31) resembles that of Dasyptcrns intermedius
and Lasiurus horeaUs, but is intermediate between the two in size. The
rostrum is broad and short and the zygomatic arches broadly flaring.
The skull of an adult female from Santa Ysabel, California, measures:
GreatestJength, 1G.4 ; zygomatic breadth, 12 ; breadth of rostrum at pos-
terior border of large premolar, 8; mandible, 12.(5 ; upper to )th row, G.4;
lower-tooth row, 8. That of an adult female from Fort ynelliiig,

Minnesota, measures:
Greatest length, 17;
zygomatic breadth, 12;
breadth of rostrum at
posterior edge of large
preuiolar, 8; mandible,
13.0: upper-tooth row,
7 ; lower-tooth row, 8.

Teeth.— ThQ teeth of
Lasiurus cinereus (fig.
32) are large and strong, but the minute upper premolar is proportion-
ally smaller than in L. borealis.
Measurements. — See table, page 115.
Specimens examined. — Total number, 50, from the following localities:

Alabama: Mobile Bay, 1.

Alberta: Eijibt miles N\V. of Red Deer, 1 (skiu. Miller coll. ).

Arizona: Temi)e, Maricopa County, 1.

California: Berryessa, Santa Clara County, 1 ; Cloverdale, I; Eureka, 1 (skiu);

Kern River, 1; Monterey, 1; Nicasio, 1; Pauamint Mountains, 2; Santa

Ysabel, 1 (skiu).
Cbibuabua; San Luis Mountains, 1.
Colorado: Larimer County, 3 (skins. Miller coll.).
District of Columbia: Washington, 1.
Georgia: Savannah River, 1.
Illinois: Warsaw, 1.

Kansas: Little Blue River, 1; North Falls, 1.
Louisiana: Pineville. 1.
Maryland: Laurel, 1.

Massachusetts: North Truro, tJ (skins, Miller coll.).
Minnesota: Fort Snelling, 1.

Nebraska: Fort Pierre, 1; Fort Union. 2; Loui^ Fork, 1.
Nevada : Vegas Valley, 1.

New Mexico: Dog Spring, Grant County, 2; Dona Ana, 1.
New York: Westville, Long Island, 1; Locust Grove, 4.
Nova Scotia : Halitax. 1.
Tamaulipas: Matamoras. 1.
Texas: Brownsville. 9 (1 skiu. Miller coll.).
Washington; Almota, 1; Fort Walla Walla. 1.

DAS YPTE KITS.

115

Average measurements of North American forma of Lasiurus.

^'ame.

Locality.

borealis .

seminulus .

ji/eifferi ..
teliolis

mi'xicanns
cinereitg.. .

Ontario : North Bay

New York : Sing Sing

District of Columbia:

Washington.
Mississippi: Washington.

Texas : Brownsville

South Carolina: Charles-
ton.

Florida : Lake Harney

Mississippi : Baj' St. Louis

Texas : Brownsville

Cuba : Trinidad

California : .* (type)

Three Rivers..

Tehama

Bakersfield . . .

Jalisco

New York: Locust Grove.
California: PauamintMts.

Cloverdale ....

Monterey

Kern lliver...

10

10
10

2dd

2dd
3??

1?

2dd

1

Id

Id

1?

6

4

29?

Id

Id

Id

^ I

H

P^

106 47.5J19.5: 7.2

110.4 50.9,19.6' 7.8

103 ;47.5il9.2 7.4

105.649. 119.2 7.4

108. 9 52. 7
100 46

97.5,45
104.3 50

115 ^54
106. 5 50.

18.6 7.9
19 ; 6.9

fM

39 6.7
39.7 7
38. 5' 6.3

40.1 7.3

40.6 6.4

40 i 7

96 47
107 57

100 '45

113.3,57
134. 5 57
136 58
130
138

140 158

19. 3j 7

19.3 7.6?

I i
i20 I 9

5^2L8J 7

.20 6.6 39

:.... 8 37
20 8 39
19.6 8.6 40
,20. 2 8. 1 41.

5 23. 2 10 50.

5 23.5 9 54
,23 10 '|46
24 9 55
23 1 9 51

9 (A

1^

79.511.8
81.7n.9
79. 3 11. 2

7.4

7
6.8

7

6.6

6.4

7

7.4,
10.6
10

10.4
10
10

1L4
11

10.1

6.5

7

6.5

80.411.810.1 6.7
82.810.51 9.8| 6.1
81 il2.710.7 6.9

79 11.

79

74 i 9.

82 , 9.

76 [10
86.5jl3

107 18

109 17.
101 |l7

110 '17
103 !l8

310

|ll
61L4
5 10. 7
7.6
8j 9.8
4 9.6
9

10.2

17.2
517.7

15

16

17.6

6.5

6.5

7

6.8

5.4

6.4

6

6

6.5

9.5

9

9

9

9

Genus DASYPTERUS Peters.

1864. Lasiurus H. Allen, Mouogr. N. Am. Bats, p. 25 (part).

1871. Dasypterus Peters, Mouatsber. K. Akad. Wi.ss., Berlin (1870), p. 912 (subgenus).

1878. AtaUtpha Dobson, Catal. Chiroptera Brit. Mns., p. 267. (Part — not Atalapha

Kafinesque, 1814.)
1893. Dasypterus H. Allen, Monogr. Bats N. Am., j). 137 (genus).

Tyjye species. — Dasypterus intermedius H. Allen.

Geographic distribution of type species. — Gulf States and northeastern
Mexico.

Geographic distribution of genus. — The range of the genus is the same
as that of the only known species.

Generic characters. — Dental formula :

. 1-1 1-1 1-1 3-3_.,..

'' 313; ^'? r_v ^^*'^' 2-2' "*' 3=3 ~'
upper incisor in contact with canine; skull (fig. 33) easily distinguish-
able from that of any other American genus of Vespertilionida; except
Lasiurus, hy its extreme shortness, depth, and breadth; ear consider-
ably higher than broad, somewhat tapering at ti]), naked on half of
dorsal surface; dorsal surface of interfemoral membrane furred on
basal half only; mamma', 1.

IKJ

NORTH AMERICAN FAUNA.

General remarlcs. — Without seeiug the South American species orig-
inally associated with D. intermedms by Peters, it is impossible to
determine whether these belong in the genus as now understood.

DASYPTEKiry INTEKMEDIU.S H. Alleu.

1863. Laaiuriis intermedins H. Alleu. Pioc. Acad. Nat. Sci. Phila. (18(j2), p. 14(i.

1864. Lusiuriis intermedius H. Alleu, Mouogr. N. A. Bats, p. 25.

1878. AtaJapha intermedia Dolisou, Catal. Chiroptera lirit. Mus., p. 274.
1893. DdsiiplevHs intermedins H. Alleu, Mouogr. P>ats N. Am., p. 137.

Type locality. — Matamoras, Tamaulipas, Mexico.
Geographic distrihidion. — Gulf States and northeastern Mexico.
General characters. — Size, large (forearm, 45-50); color, light brown.
Uars. — The ears are short, reaching barely to nostril when laid for-
ward. The dorsal surface is densely furred on basal half, but other-
wise the ear is naked except for a sprink-
ling of hairs on inner side, especially along
anterior edge. Beginning at lower edge
of basal lobe the anterior margin is first
strongly convex, then nearly straight for
a distance of abont 0 mm., then abruptly
convex (or even angular), after which it
continues nearly straight to narrowly
rounded oif tip. Posterior border slightly
concave immediately below tip, then
gently and evenly convex to notch above
posterior basal lobe. Posterior basal
lobe well developed, slightly notched on
lower side, and joining face about 5 mm.
behind angle of mouth.

Tragus blunt and bent forward, anterior
border Jiearly straight to slight concavity
just below tip. Posterior border convex
immediately below tii^, then straight to point opposite anterior base.
Here a sharp angle is formed, below which the margin is irregularly
crenulated to base.

Feet. — The foot is moderate in size, a little less than half as long as
tibia. Calcar slightly shorter than tibia, very indistinctly keeled and
terminating obscurely or in an ill-defined lobe.

Membranes. — Membranes thick and leathery. Wing membrane at-
tached at base of toes, uropatagium near tip of last caudal vertebra.
Free border of uropatagium slightly longer than calcar.

Fur and color. — The fur is full and soft. On the middle of the back
it is about 12 mm. in length. The fur of the back extends on basal half
of outer side of ear, basal half of dorsal surface of interfemoral mem-
brane, and base of wing membranes. On the latter it occupies a strip
about 10 mm. in width. There is a sliiiht tuft of hair at the base of the

Fig. 33.

-Skull of Dasi/ptenis intennc-
dius (X2.)

DASYPTERlTft TNTERMEDIUS.

117

tliiiiub, and in many specimens a faintly indicated tuft near proximal
end of forearm. On the ventral surface the fur barely reaches the
uropatagium except along- the basal fourth of tail. A thin coating of
fur occupies the under side of the wing membrane to a line joining-
elbow and knee. Beyond this it extends in a strip about 10 mm. wide
along posterior edge of forearm to bases of fingers. The greater part
of the propatagium is thinly furred.

Color light yellowish brown of variable shade, the hairs throughout
tlie body with narrow dark plumbeous bases and those of the back
with faintly dusky tips. The general effect is suggestive of the color
of PipistreUiis snhfanis.

Sh(U and teeth. — The skull (fig. 33) and teeth (fig-. 34) have been suf-
ficiently described under generic characters. The skull of an adult

Fig. 34.— Teeth o{ Dafttrpferus intermedius (X5).

female from Brownsville, Tex. (Xo. 52r>40, U. S. National Museum),
measures: Greatest length, 19.6; zygomatic breadth, 15; breadth of
rostrum at posterior edge of premolar, 9; mandible, 15; upper tooth
row, 8; lower tooth row, 9. The skull of an adult male from Houma,
La., measures: Greatest length, 18; zygomatic breadth, 13; breadth
of rostrum at posterior edge of premolar, 8; mandible, 14; upper tooth
row, 7; lower tooth row, 8.0.

Measurements. — Average measurements of 18 specimens of Dasypte-
rus intermediKS from five localities are given in the following table:

ArercKje measurements of IS specimens of Dasupierus intermedins from 5 localities.

Locality.

o ^

a

D

O
H

1

'3

H

o
o

1

o

e

a
to

§

ID

S

ll

O

Texas: Brownsville

Louisiana : Lafavette

10??
2??
2cfcf
3
Id

145

126.5

130

127

120

65.9

52

61

03.5

54

24.9

20

18.9
20
18

10
8.7
9
9

8

55
48
46
45.5

47

8.9

7

6

111

96
95.5

18.8
18.5
18

14.4
15.5
15

8.8
9 3

9 4

Florida: Old Town

Mullet Lake

7

95

17

U

8

118 NORTH AMERICAN FAUNA.

Specimens examined. — Total iiiiiubei' 7li, Jroin the following localities:

Florida: Davenport, 1 (skin); Mullet Lake, 1; Old Town, 3.

Lonisiana: Lafayette, 2; Houma, 2 (1 skin).

Mississiiipi : Hancock County, 1 (skin).

Tanianlipas: Matanioras, 3 (2 skeletons).

Texas: Brownsville, 57 (2 skins); Padre Island, 1; Cameron Connty, 1.

General remarls. — Aside from its generic characters Dasypterus
intermedins i.s distinguishable among North American bats by its large
size, small ears, and yellowish brown color.

Specimens from Louisiana, Mississippi, and Florida average dis-
tinctly smaller than those from Brownsville, Tex. (which are essen-
tially topotypes). More extensive material than that now available
may show the necessity of recognizing two subspecies, a larger Tarn au-
lipan (typical) form, and a smaller Austroriparian form.

Genus NYCTICEIUS Rafinesque.

1819. Xycticeiiis Rafinesqne, Jonrn.de Physique, LXXXVIII, June, 1819, p. 417.

1827. Nyciicens Lesson, Man. de ISIanim., p. 98.

1827. Nycticejiis Temminck, Monograpbies de Mamm., I, p. xviii.

1830. Nycticeyx Wagler, Natiirl. System der Ampbibien, p. 13.

1831. Nycticea Le Conte, McMnrtrie's Cuvier, Animal Kingdom, p. 432.
1864. Nycticejus H. Allen, Monogr. N. Am. Bats, p. 11.

1878. Nycticejus Dobson, Catal. Chiroptera Brit. Mus., p. 266.
1893. Nycticejus H. Allen, Monogr. Bats N. Am., p. 131.

Type species. — Nycticeins humeraUs Eafinesque.

Geographic distriimtion of type species. — Austral zones in the Eastern
United States.

Geographic distribution of genus. — Austral zones in the Eastern United
States. Cuba.

Generic characters. — Dental formula:

*' 3-3' ^' 1-1' •^*'*' 2-2' *'*'3-3 — '
upper incisor distinctly separated from canine; lower incisors scarcely
crowded; outer lower incisor tricuspidnte and not smaller than others;
skull low and narrow ; uropatagium furred at extreme base only ; tragus
blunt and bent forward; tip of tail free from membrane; mammae, 2.
The genus Nycticeins as thus defined is peculiar to America, where it
is represented by one species. It differs in dental formula from all
other genera of American Vespertilionidw except Basypterus and
Rhogel'ssa. From the former it is distinguishable by its differently
shaped skull, wide space between upper incisor and canine, and essen-
tially naked uropatagium. From Rhogei'ssa it is separated by details
in the structure of teeth and skull, as well as by external characters,

NYCTICEIUS HUMERALIS Rafinesqne. Rafiuesqne's Bat.

1818. VespertUio humeralis Rafinesque, American Monthly Mag., Ill, p. 445.

1819. Nyctieeins humeralis Rafinesque, Journ. de Pbysi(|ue, LXXXVIII, p. 417.
1831. Nycticea crepuscuJaris Le Conte, McMnrtrie's Cnvier. Animal Kingdom. I. p. 432.

NYCTICEIUS Hr.MKRALIS

119

18t)4. Xiicticciiia cnpiiscularin i[. AlU'ii, ilonogr. X. Am. Hats, p. 11.
1878. Nycticejiis crepnscularis Dobsou, Catal. Chiroptera Brit. Mus., p. 266.
1891. XijcticeJHs humeralis Thomas, Ann. & Ma<^. Nat. Hist.. 6th ser., VII, p. 528.
1893. XtjcticeJHS limncralis H. Allen, Monogr. Bats, N. Am., p. 132.

Ti/2)e localittj. — Kentucky.

Geof/rajjhic distribution. — Austral zones in the eastern United States
west to Arkansas and southern Texas.

General characters. — Size, medium (total length, 88 to 95; forearm,
34 to 38); color, dull brownish, slightly paler beneath.

Ears. — The ears are small and for their size remarkably thick and
leathery. They are naked throughout except at extreme base above.
Lower anterior half of inner surface with a few short scattered hairs.
Anterior border strongly convex immediately above small but dis-
tinct anterior basal lobe, then very slightly
convex to narrowly rounded off tip. Poste-
rior border gently concave from immediately
below tip to a little below middle, then con-
vex to slightlj' developed external basal lobe.

Tragus short, broad, and blunt, bent slightly
forward; posterior base with distinct lobule.

Membranes. — The membranes, like the ears,
are thick and leathery. Wing membranes at-
tached at base of toes, uropatagium at middle
of terminal caudal vertebra.

Fur and color. — The fur is sparse and short,
that on middle of back averaging about G
linn, in length. It is closely confined to the
body, barely reaching extreme base of uropatagium and flight mem-
branes.

Color duU umber brown above, paler below, the fur everywhere
plumbeous at extreme base, but the dark basal color less well defined
than in other species with which Nycticeius is found associated. The
exact shade varies slightly, but is usually burnt umber or mummy
brown on the back and raw umber or hair brown on the belly. One
skin from Hickman County, Tenn. (No. 30037, U. S. National Museum),
is dark sepia above, broccoli brown below.

Slcull. — The skull (fig. 35) is short, broad, and low. That of an adult
female from Sans Souci, N. C. (No. 43037, U. S. National Museum), meas-
ures 14 ram. in greatest length and 10 mm, in zygomatic breadth; great-
est length of mandible, 10.6. Dorsal profile nearly straight from external
nares to occiput, but slightly convex over front part of brain case.
Occiput never developing strongly marked ridges. Length of bony
palate behind molars (exclusive of central spine), about half width of
interpterygoid fossa.

Teeth. — The teeth (fig. 30) are not so large as might be expected from
the massiveness of the skull. Upper tooth row of adult female from
Sans Souci, N. C, G; lower, 0.8. Upper incisor close to canine, but

Fig. 35.— Skull of Nycticeius
humeralis (X2).

120

NORTH AMERICAN FAUNA.

separated from it by a space less tlian half as great as the diameter of
the incisor. Upper molars Diuch narrower on the inner side than on

the onter side. Outer
lower incisor with trans-
verse diameter of crown
slightly greater than
that of second or third.
MeasuremenU. — In the
following table average
measurements are given
of 35 specimens of Nyc-
ticekis humeralis from the United States, and for comparison, Gund-
lach's measurements of a dry specimen of N. humeralis cuhmius.

Fig. 36. — Teeth of XiicHceiv n humeralis ("-'5).

Average vwosurements of 36 specimens of XiicUceius.

^

P

o •

,a

u

HI

^

^1 0

tn

-S

«

H

<D

Name.

Locality.

1

>

C8

i

B

+2 ^

<c fell

O

.a

til

!='S'

o

cS

X:

g

o

eS

10

El

^

H

H

H

1=^

bH

H

h)

H

H

Pennsylvania : Carlisle

10

93.5

36.9

13.8

6.7

36.2

5.4

64. 5'l3. 9

C.l

Virginia: Dismal Swamp

5

91.4

35.8

13.4

7.7

35.8 6.6 63.413.510.5

6.2

Tennessee : Big Sandy

10

92.2

36.3

13.7

7.3

34.3 5.5 63.413.1 9.9

5.9

10

1

92.3

37.2
.9

13.6 7.2
11 fi-7

36.4 5.3 65.212.7! 8.8

5 1

cnbanus

Cuba

SO

5. 5 55

1

Specimens examinrd. — Total number 154, from the following localities:

Arkausas: Fort Smitlt, .'i.

District of Columbia: Wasliington, 2.

Florida: Titnsville, 1; Chattahoochee, 1.

Georgia: Ricoboro, 3.

Indian Territory: Redland, 4.

Kentucky: Hickman, 7.

Louisiana: Mer Rouge, 19; Pineville, 1.

Mississippi: Bay St. Louis, 17; Washington, 2.

North Carolina: Bertie County, 9; Sans Souci, 4 (skins).

Pennsylvania: Carlisle, 12.

Tamaulipas: Matamoras, 1.

Tennessee: Big Sandy, 13; Danville, 2; Warner, 1; Arlington, 4; Hickman

County, 1 (skin).
Texas: Brownsville, 32; Paris, 3; Arthur, 1 ; Lomita Ranch, 2; Hidalgo, 1.
Virginia: Dismal Swamp, 5; near Riverton, 1 (skin, Miller coll.).

NYCTICEIUS HUMERALIS CUBANUS (Gundlach).

1861. Vesperns eiihauus Gundlach, Monatsber. K. Preuss. Akad. Wiss., Berlin, p. 150.
1877. Nycticeins ciihauiis, Gundlacl), Coutribncion ;i la Mamalogia Cubana, p. 33.

Type locality. — Cuba.
Geographic (Hstributio n. — C uba .

NYCTICEIUS HUMERALIS CUP.ANUS. 121

Characters. — I have not seen specimens of XycHcchis from Cuba, but
Gundlach's careful description of the animal leaves no doubt that it is
distinct from the form occurring on the mainland. It is distinguished
from the latter by smaller size and apparently also by paler color. A
translation of Gundlach's second and more perfect account of the
animal is as follows: "Pelage above light tawny (the fur blackish at
base), beneath pale reddish tawny (the base of the fur likewise black-
ish). Face and flight membranes blackish brown. The nose appears
somewhat divided by the projecting nostrils; between the nose and the
eyes there is on each side a i)rotuberance with bristly hairs. Ears oval,
lengthened (8 mm. high in front). The anterior base rounded and
spreading outward; the posterior at the angle of the mouth forms a
semicircle. This semicircle, which bends inward to the tragus, forms
another rounded enlargement. Tragus oblong, scarcely narrowed
throughout, somewhat bent in the form of a sickle forward and pro-
vided with a tooth-shaped lobule at the base of the exterior border.
Nails tawny.

"The measurements of a dry specimen are as follows : Width between
extremities of wing, 0.180 m.; total length of body, 0.045; length of
tail, 0.029; length of head, 0.016; length of ear, 0.012; length of fore-
arm, 0.030; length of thumb, 0.005^; length of second or index finger,
0.029i; length of third finger, 0.055; of fourth finger, 0.046; of fifth fin-
ger, 0.040; length of tibia, 0.011; length of foot to the end of nails,
0.0003; length of calcar, 0.013.

" This is a rare species. I have only observed it at Habana (Cerro) in
a house where it lived in a crack above the window, and in the field
near Cardenas, where I killed it while flying about at dusk. A female
contained two embryos in May." ^

' Pelaje, por encinia p;tlido-pardo (los polos con la base uogruzca), por debajo palido
bermejizo-pardo (labase de los pelos tambiou negruzca). Cara y membranas vola-
doras, morenas. La nariz aparece algo dividida por las ventanas saltonas ; entre la
nariz y los ojos hay en cada lado un lollo con pelos tiesos. Orejas ovales, alargadas
(por delante con 8 mil. de alto). La base anterior, redonda y extendida bacia fuera;
la posterior, pelada en el angulo de la boca y extendida en Tin semicircnlo, ([ue incli-
nado bacia dentro basta la orejnela, forma una segnnda ampliacion redonda. Ore-
juela oblonga, adelgazandose ap(^na.s, algo encorvada en forma de boz bacia delante,
y provista en la base del borde exterior con \\\\ lobnlo deutiforme. Unas pardas.

Las mcdidas, touiadas de nn ejemplar disecado, son:

Anchnra entre las puutas del ala, 0.180 mil. ; longitnd basta el fin del cuerpo, 0.045;
longitiid del rabo, 0.029; longitnd de la cabeza, 0.016; longitnd de la oreja, 0.012;
lougitud del antebrazo, 0.030 ; longitnd del pulgar, 0.005i ; longitnd del seguudo dedo
6 iiidice, 0.0291; longitnd del tercero dedo, 0.055; longitnd del quarto dedo, 0.046;
longitnd del qninto dedo, 0.040; longitnd de la tibia, 0.011 ; longitnd del pi<S basta el
fin de les nuas, 0.006J; longitnd del espob'm, 0.013.

Es eapecie rara. La be observado solamente en la Habaua (Cerro) en una casa
donde vivia, en las rendijas sobre una ventana, y en el campo cerca de Ciirdenas,
donde la mate al oscurecer volando. Una 9 tenia en mayo dos embriones.

122

NORTH AMERICAN FAUNA.

Genus RHOGEESSA H. Allen.

1866. Rhogeessa H. Allen, Proc. Acad. Nat. Sci., Phila., p. 285 (genus).

1873. Iiho{joessa Marscliall, Nomeuclator Zoologicns, Mauim., p. 11.

1878. Iiho(/c(--isa Dobsou, Catal. Cliiroptera Brit. Mus., i>. 24;") (subgenus of ' Vesperitf/o').

1893. Bhogeessa H. Allen, Monogr. Bats N. Am., p. 132 (genus).

Type species. — Rhogecssa tumida H. Allen.

Geof/raphic disiribuiion. — Tropical Mexico,
Central America, and probably northern South
America (known from Margarita Island, Vene-
zuela).

Generic characters. — Dental formula :
. 1-1 1-1 1-1 3-3 ^,.

*'3=3^ ^M-1' ^''^2^ '"'3-3=^'^'
lower incisors crowded, the outer cusp of first
and secomi obsolete; third lower incisor greatly
reduced in size, unicuspidate (figs. 37 a and 38«);
upper incisor very close to canine or in contact
with it; skull small, light, and papery, narrow
and deep; external form variable, but tragus
always straight or bent backward, and tail in-
cluded to tip in interfemoral membrane.

Remarks. — The genus Rhof/ei'ssa has received
varying treatment. It was originally described
as a full genus whose relationships were sup-
posed to be with Nyciiceius and Nyctinomns.
In 1878 Dobson referred it to ^Vesperugo^ as a
subgenus. This view has been adopted by
most subsequent writers except Mr. Oldfield
Thomas and Dr. Harrison Allen, both of whom
have recognized Rhogecssa as a
full genus related more closely
to NycUceitis than to any of the genera usually included
under the name ' Vesperugo.^ Mr. Thomas has pointed
out characters in which Rhogecssa resembles Antro-
zons. These characters, the reduced size of the outer
lower incisor and slightly crenulate i)osterior border
of tragus, seem to be instances of parallel develop-
ment rather than indications of genetic relationship. The genus
Rhogecssa is closely related to Nycficeius, but the peculiarities of the
lower incisors and the general form of the skull are enough to warrant
its recognition.

Fig. 37.— Left manflibnlar in-
cisors of (a) JOiogcetsa and
(b) Nycticcius ( ■ 20).

Fig. 38.— Cro-mi.s of in-
cisors of right manili-
ble of (a) Rhogeessa
and (6) Kyctieeim

(20).

RH0GEKS8A TUMIDA. 123

KEY TO SPECIES OF IJHOGEESSA.

Lateral mandibular incisor scarcely one-twentieth as large as central

incisors alleni (p. 128)

Lateral mandibnlar incisor one-half to two-thirds as large as central
incisors.

Ear laid forward, reaching ahont 6 mm. beyond tip of nose <jraeilis (p. 126)

Ear laid forward, rcacliing abont to tip of nose.

Fur grayish brown at base par nil a (p. 125)

Fur yellowish throughout.

Forearm about 30 mm tumida (p. 123)

Forearm about 25 mm minutilht (p. 125)

RHOGEESSA TUMIDA H. Allen.

1866. L'liof/eessa tumida H. Allen, Proc. Acad. Nat. Sci. Phila., p. 286.
1877. Vesperugo parrnlus Dobson, Catal. C'hiroptera Brit. Mus., p. 245.

Type locality. — Miiador, Vera Cruz, Mexico.

GeogyapMc (li.stribntion. — Central America and soutliern Mexico.

General characters. — Size small; lengtli, 70 to 75; tail, 30 to 33; fore-
arm, 27.4 to 30. Calcar strong, distinct, slightly longer than free border
of uropatagium, terminating in a small but evident lobule conspicuously
keeled on tlie posterior border. Free border of uropatagium naked.
Ears moderate, laid forward tliey reach about to tip of nose. Wings
from base of toes. Legs and feet short and strong, the feet when out-
stretched reaching to within 5 mm. of tip of tail. Fur yellowish, the
hairs on the back with dusky tips.

Ears. — The ears (PI. I, fig. 8) are moderately long, reaching, when
laid forward, about to tip of nose; the substance of the conch thick
and leathery. Anterior border strongly concave from base to a little
past middle, then straight to narrowly rounded-off tip. Posterior border
concave just below tip, then gently and evenly convex to base. No
indication of basal notch.

Tragus directed slightly forward; the anterior edge nearly straight,
but slightly concave at base, and curved a little backward at tip.
Posterior edge faintly crenulate, concave below tip, then concave to
slightly developed basal lobe. Greatest width of tragus at about
middle of posterior border.

MemhranoH. — The membranes, especially the uropatagium, are remark-
ably thick and leathery for so small a bat. Throughout they are wholly
naked except close to the body and along the veins on the interfemoral
membrane. Wings from base of toes. Uropatagium (PI. I, fig. 13)
attached at tip of terminal caudal vertebra.

Feet. — The feet and legs (PI. I, fig. 13) are short and strongly built,
in this respect resembling N. humeralis. The foot is scarcely oue-half
as long as the tibia, and the toes are slightly longer than the sole.
The toes are not united by membrane at base. Calcar distinct and
strong, slightly longer than free border of uropatagium. Lobule at

124 NORTH AMERICAN FAUNA.

tip of calcar «niall but dist inct. Keel well developed and supported by
one or two cartilaginous outgrowths.

Fur and color. — There is nothing peculiar in the distribution of the
fur. It extends in a very narrow line on the wings along the side of
the body both dorsally and veutrally, and on the uropntagium covers
the basal fourth dorsally but scarcely reaches the membrane on the
ventral side.

In color the fur is dull yellowish brown throughout, scarcely paler
\rentrally, the hairs dusky at tip. Ears and membranes dark brown.

Sliull, — In general appearance the skull of Rhogeessa tumida (fig. 39)
stands between that of Kycticeius humeralis and Plpisirellus suhjiavuH.
The skull of an adult female from Santo Domingo, Oaxaca (iSTo. 73-JG7,
United States National Museum, Biological Survey collection), meas-
ures: Greatest length, 13 j zygomatic breadth, 8.4; breadth of rostrum
at anterior edge of first molar, 5; mandible, 9;' upper tooth row, 5.6;
lower tooth row, 6. That of an adult female from Patuca, Honduras
(No. 21017, United States National Museum), meas-
ures: Greatest length, 12.4; zygomatic breadth, 8;
breadth of rostrum at anterior edge of first molar,
4; mandible, 9.4; upper tooth row, 5; lower tooth
row, 6. The rostrum is relatively narrower than
in Kycticeius and the occiput is more elevated.
The muzzle is distinctly concave in front of orbits
instead of flat or almost convex as in Kycticeius.
Teeth. — Upper incisor usually in contact with
riG.?.9.-skuiiotAv«)<7<>;>«sa cauiuc, though occasionally separated by a narrow
tuwida{x2). space (fig. 40«). Maxillary teeth essentially as in

Nyeticeius, but premolar relatively larger and i)osterior molar narrower
in proportion to its length.

Lower incisors greatly crowded, the outer cusp pf i 1 and i 2 much

smaller than middle and inner cusp. Outer lower incisor uni uspi-

date, about one-half the size of i 1 or i 2. Other mandibular teeth

essentially as in Kycticeius, but premolars more crowded.

Measurements. — See table, page 129.

Specimens examined. — Total number, 10, from the following localities:

Colima: Colima, 3.

Costa Rica: — , 1.

Guatemala: Huehiietau, 1.
Guenevo : Amula, 1 (Merriam coll.).
Honduras : Patuca, 2.
Oaxaca: Santo Domingo, 1.
Vera Cruz: Mirador, 1.

General remarks. — Rhogee'ssa tumida needs comparison with R. par-
rula and R. minutilla only. From the former it is distinguished by its
clear yellowish fur without darker base, and from the latter by its con-
siderably larger size.

As the mandible is imperfect, this measurement is only approximately correct.

RHOGEESSA FARVULA. 125

Dobson aud most subsequeut authors have wiougly applied the spe-
cific uanie parvula to this species. So far as known R. parvula is
restricted to the Tres Marias Islands.

[Tlio followiug species is not North Americau, but is iutroduced here to complete
the accouut of the genus I\hofjei!ssa.~\

RHOGEESSA MIXUTILLA Miller.

1896. Vesperiigo parcidiis Robinson, Proc. U. S. National Museum, XVIII, p. 651 (not

Ehof/ei'ssa parvnla H. Allen).

1897. Rhogves=ia minuiiUa Miller, Proc. Biol. Soe. Washington, XI, p. 139, May 13, 1897.

Type locality. — Margarita Island, Venezuela. (Type in U. S. JS^ational
Museum, Xo. 6321G.)

Geographic distributiofi.— This, species is probably confined to Mar-
garita Island.

General characters. — Similar to Ehof/ei'ssa tumida, but considerably
smaller (forearm, only 25 mm.).

JEars. — The ears of the type (when relaxed by soaking in water)
appear to be smaller and narrower than in B. tumida, but otherwise not
peculiar.

Membranes, feet, and distribution of fur. — As in K. tumida.

Color. — Fur everywhere light yellowish brown to base, the hairs on
the back tipped with chestnut. The color appears to differ slightly
from that of i^. tumida, but I have too few skins to make an adequate
comparison.

S'lcull. — The skull of the type and only known specimen is so much
injured that its characters can not be determined with certainty, but it
appears to be smaller and relatively narrower than that of R. tumida.
Greatest length, 11.8; length of mandible, 9; upper tooth row, 5 ; lower
tooth row, 5.6.

Teeth. — The teeth are essentially as in R. tumida.

Measurements. — The measurements of the type specimen are given in
the table on page 129.

Specimens examined. — One, the type.

General remarJcs. — Rhogeessa minutilla is a small insular form most
closely related to 7^. tumida, but apparently perfectly distinct. So far
as I know the genus Rhogel'ssa has not yet been recorded from the
mainland of South America, where, however, it doubtless occurs.

RHOGEESSA PARVULA H. Allen.

1866. EliogeVssa parvula H. Allen, Proc. Acad. N'at. Sci. Phila., p. 285

Tpye locality. — Tres Marias Islands, Mexico.

Geographic distribution. — Tres Marias Islands.

Characters.— As I have seen no specimens of this species, I cpiote
the original description entire. It is as follows:

"Ear sub-acute at tip; lips whiskered; eyes very small, each fur-
nished with a wart above; similar growth seen beneath chin. Fur
above silky, not thick, of a light greyish-brown at basal third, fawn-

126 NORTH AMERICAN FAUNA.

chestuut-brown at apical two-tliirds; that of head same color, iimijing
ou to the ears one-half their height. Beneath, basal third inclined to
greyish; apical two-thirds grayish fawn. Membranes almost black,
naked, excepting basal fourth of interfemoral membrane behind, which
is furnished with a small, short patch of glistening fur.

" Measurem en ts — 7841 .

"Height of auricle 6" [12.7 mm.]; height of tragus 3" [6.4:]; length
of head 7" [14.8J; length of body 10" [21.1]; length of tail 1' 2" [30.o];
length of forearm 1' 1" [27.4]; length of longest finger V 11" [48.5];
length of thumb 2" [4.2]; length of tibia o" [10.6]; length of foot 2h"
[5.3]; expanse 6' 7" [16.7].

"Two individuals, $ and 9 ; Nos. 7841, 7842, Museum of Smithsonian
Institution. Alcohol.

"Tres Marias, Mexico, Col. Grayson."

RHOGEESSA GRACILIS sp. nov.

Type from Piaxtla, Puebla. Adult ^ (in alcohol). No. 706'J4, U. S. Nat. Museum,
I5iological Survey collection. Collected Nov. 24, 1894, by E. W. Nelson and E. A.
Goldman. Collector's number, 7099.

Geograpliic distribution. — Southern Mexico (Puebla and ' Isthmus of
Tehuantepec').

General characters. — Size, medium; length, 79 to 82; tail, 38.6 to 41;
forearm, 32 to 33. Calcar slender but distinct, a little shorter than free
border of uropatagium, terminating in a small lobule, distinctly keeled
on the posterior border. Free border of uropatagium naked. Ears long;
when laid forward extending about 6 mm. beyond tip of nose. Wings
from base of toes. Feet and legs long and slender, the outstretched
feet reaching to within about 10 mm. of tip of tail.

IJars. — The ears (PI. I, fig. 7) are long, and at the same time broadj
laid forward they reach about 6 mm. beyond tip of nose; the substance
of the conch thin and translucent. Anterior border strongly convex
from base to a little below middle, then straight or very slightly con-
vex to the rather broadly rounded oft tip. Posterior border concave
below tip to about middle, where it bends abruptly outward, then grad-
ually convex to base. A very faintly indicated basal notch and basal
lobe. About 5 mm. above the crown and an equal distance from the
tip of the ear conch in the male is developed a conspicuous, flattened-
pyriform, glandular thickening with the large end toward the anterior
margin of the ear and the main axis nearly perpendicular to that of the
auricle. The thickened mass is 5 mm. in length, 4 mm. wide at the
broad end, 2 mm. at the narrow end, and 1 mm. thick. It is most con-
spicuous on the dorsal side of the ear, where, although not different in
color from the rest of the ear, it is noticeably raised above the surface,
and the boundaries are sharply marked. On the inner side of the
ears the thickenings are less definite in outline, but are noticeably
paler than the surrounding integument. When these structures are
examined with a lens it is seen that they are thickly covered on the

RHOGEESSA GRACILIS. 127

outer side with pores lyiug mostly at tlie bases of tlie fine hairs with
which the surface is beset. The thickened masses are of exactly the
same size aud shape in the two ears and are placed symmetrically with
respect to the outlines of the couchs.

Tragus slender and taper pointed, slightly bent backward at the
tip, and broadest opposite anterior base. The anterior border is
slightly concave at base, then evenly convex to tip. The posterior
border is strongly concave from tip to a point slightly above the middle,
where the tragus attains a width nearly equal to that at level of ante-
rior base. From this point to the basal lobe the posterior border is
nearly straight and about parallel with the lower part of the anterior
border. Basal lobe small but prominent. Posterior border of tragus
crenulate, especially near the middle, where there are five or six minute
sharply projecting jioints, from the bases of which thickened processes
may be traced a sliort distance into the substance of the tragus when
the latter is held to the light.

Memhranes. — The membranes are thin and semitransparent, the uro-
patagium not different in texture from the wings. Throughout they
are entirely naked, except for a narrow line of hair on the wings extend-
ing along sides of body about to a line drawn halfway between knee
and elbow. On the uropatagium there is also a narrow hairy area
close to body and a sprinkling of fine haii'S along the veins. Wings
from base of toes. Uropatagium (PI. I, fig. 12) attached at tip of ter-
mimil caudal vertebra.

Feei. — The feet are small and weak, distinctly less than half as long
as the slender tibiae (PI. I, fig. 12). Toes longer than sole, cleft to
base. Calcar slender but very distinct, about as long as free border
of uropatagium aud terminating in a small and ill-defined lobule.
Keel remarkably well developed, extending from near tip of calcar
almost to base and supported by four cartilaginous processes.

Fur and color. — The fur is long, that on middle of back averaging
about 9 mm. It extends farther on the membranes than in B. parvula^
but otherwise shows no peculiarities. In color it is everywhere light
sepia at base, then dull yellowish brown, that on the back tipped with
chestnut. As this description is from a specimen that has been
immersed in alc(»hol for nearly two years and a half, it ca)\ not be more
than approximately accurate.

SkulL — The skull of Bhogeessa gracilis is longer and more slender
than that of R. tumida, and the forehead appears to be more abruptly
raised above the face line. The zygomata are less widely flaring in
front. Apparently the occiput is considerably narrower than in B.
tumida. From the material at hand it is, however, impossible to deter-
mine the cranial characters with accuracy, since of the skull of the
female topotype there remains only the mandible and rostral portion,
while the skull of the type is so nuich injured that it would not hold
together if removed from the skin and cleaned.

128

NORTH AMERICAN FAUNA.

Fig. 40.

-Teeth of (a) lihogeessa tumida and (6) R.fjracUis

(>5).

Teeth. — The teeth of Rhogel'sa gracilis (fig. 40) differ from those of
B. tionida in uiimerous details. The crowns of the upper molars are
much narrower on the lingaal side and the posterior upper molar is
considerably broader. The front lower preniolar is slightly larger than
in li. tumida, but otherwise the mandibular teeth show no distinct

differences.

Measurements. — See table,
page 129.

Specim ens ex a m ined. —
Total number, 3, from the
following localities:

Isthmus ofTeluiantepec, 1.
Puebla: Piaxtla, 2.

General remarks. — Bhoge-
i'ssa gracilis is so readily
distinguished from the other
species of the genus by its
slender form and very large ears that no detailed comparisons are
necessary.

This is the only species of North American Yesperiilionida' in which
I have found any sexual differences in cutaneous structures.

RHOGEESSA ALLENI Thomas.

1892. L'hogeessa uUeni Thomas, Ami. & Mag. Nat. Hist., 6th ser., X, j). 477, Decem-
ber, 1892.

Type loealiti/. — Santa Rosalia, near Autlan, Jalisco, Mexico. Type
in British Museum.

Geographic distrihiition. — Bhogeessa alleni is known from the type
locality only.

Characters. — As I have not seen the type and only known specimen
of this bat, I copy Mr. Thomas's original description:

"Decidedly larger than Bh. parrula; muzzle obliquely truncate as
in that species. Ears large, laid forward they reach about 1 or 2 millim.
beyond the nostrils; their inner margin very convex forwards below,
straight or even slightly concave above; tip narrowly rounded off;
outer margin concave below the tip, then straight, becoming slightly
convex below, outer basal lobe but little marked. Tragus long, its
broadest point opposite to base of its inner edge; inner edge straight
or slightly concave, tip rounded, outer margin slightly convex, the edge
indistinctly crenulate, somewhat as in Antrozoiis pallidns ; a marked
lobule at the base of the outer margin, above and below which there is
a concavity. Thumb very short and thick, no longer than in Bh.parvida.
Posterior edges of wing-membrane bordered with white, bifid tip to
fourth finger unusually distinct; wings from the base of the fifth toe;
post-calcareal lobe small and narrow: tip of calcar projecting slightly
from the back of the membrane; tail included in membrane to the
extreme tip.''

RHOGEESSA ALLENI.

129

^^ Teeth. — Upper incisors one on each side, long, slender, nnicuspidj
upper premolars large, quite close to tbe canines; no trace of a minute
anterior premolar. Lower incisors six, the lour median ones broad,
tricuspid; the outer ones unicuspid, exceedingly minute, i)ractically
invisible from in front, and scarcely one-twentieth of the size in cross
section of the median incisors; far smaller therefore both absolutely
and relatively than in Eh. parvula.

"Dimensions of the type (an adult female in spirit): —

"Head and body 47 millim; tail 41; ear above head 12.2, from notch
10; tragus, inner margin 7; forearm 35; thumb 5; metacarpal of third
finger 33.5; lower leg 15.5; hind foot 7.1; calcar 15.

"Skull of a second specimen: Occiimt to gnathion 14.7; greatest
breadth 9.5; distance from front of canine to back of m. 3 5.4."

Measurements of specimens of lihogeissa.

Xame.

Locality.

tumida Vera Cruz : Mii-ador

Colima : Colima

C'olima

Colima

Oaxaca: SaiitoDomingo.
Guatemala: Huehuetau.

Honduras: Patuca

Patuca

parvula i Tres Marias Islands

ininntilla Venezuela: Margarita

Islam).

f/racilig Puebla : Piastla

Piaxtla

Isthmus of Teliuantepec .
alkni Jalisco: Autlan

8195' J' ad. '65. 5 25. 4 10. 4 5. 129. 5 4. 8 57. 2 .

-;

,

<u

CS

ffl

p

(O

:p

*"

«M

?.^

^

t[j

'd

o

fA

^

)7.2

7.3

12. S 9.6 7
12.4 9.4 7.4

52102 9 ad. 75 34 12 5 29. 8 4 52

52005 cfad. 170 Xi 11.4 5.4^0 4 52

52060 cTad. 70 j30 ;11 | 5.428 4 51 jl2.6|9

732G9 9a<l I. ...12 7 \s3 4.463 13. 6' 9. 6

78600 cT ad. 75 133 11. 4 6 30 -1. 6 .56 14 9

21016 9 ad. 73 |30 12 6 30 4 55 12.6 9

210171 ? ad. ]75 31 12.4 5.6 30 i 57 13 19

7841 'rf ad. 65. 5 29. 5 10.4 5. 3 27.4 4.148.5^ {

63216 -'^ ad 25 11 5 25 3.6 51 111.8 8

7U691 9 ad. ,79 |38. 6 14 6
70604 Vad. 82 41 1 14. 6 5
11240' ? ad. 77 37 13 5
1 sjad. ....:41 i-..., 7.1

4 60
4.4 61

58

17

16.6

16

11

11.8

II

7

7.2

7

7

7

6.4

6.4

10
10

9

7

'Type, measurements from H. Allen.

2772— Is^o. 13 9

'Typo.

8 Type, measurements from Thomas.

INDEX TO GENERIC AND SPECIFIC NAMES.

[Names iu italicH :ue licre I't-cognizecl as valid.

abramus {Fipiatnllufi), lu.
Ailelonycteri.s, 12,95.

fiiscus, 39, 96.
Aeorestes, 12, 55.
atKiii.s ( Vespertilio), 20, 38.
agilis (Ve.spertilio), 73.
alancengif; (ilifotie), 39, 40, 56, 63-64.
albescens (2[i/oti,s), 12,59,84.
albescens ( Vespertilio), 20, 39, 64, 66.
alb!niilaris ( Vesjiertilio), 20, 39, 41, 104.
albigtilaris (Vesperugo), 39. 104.
albigularis (Vespenis), 15, 20, 104.
alboliinbatus (Vesperugo), 20.
alcytlioe (X'cspertilio), 15, 95.
allm! (Rhnrjecssn), 20, 40, 41, 123, 128-129.
aiuericaua (Atalapha), 13, 20, 106.
Antrozohtiv, 41.
AntrozoHS, 12, 42.

pacificus, 38, 40, 45-40.

pallidus, 38,40,43-45.
avi.stippe (Vesperugo). 20.
aristippe (Vesperus). 15.
anpiatus (Vespertilio), 20, 96.
Artibni^,\S.
Atalapha, 13, 105, 115.

araericaua, 13, 20, 106.

borcalis, 106.

laudata, 13.

ciiierea, 40, 112.

ega, 13.

egregia, 13.

frautzii, 25, 40, 111.

fiis.;ata. 26.

interniodia, 13, 40. 116.

mexicana,31. 111.

iioveboracensis, 39, 106.

pfeitteri, 33. 39, 110.

sewinola, 35, 109.

sicula, 13.

teliotis, 36, 39, 110.
auduboni (Vespertilio). 20, 86.
awitdg (Plecotus). 140.
aiiiitns. (Vespertilio), 18.
mil traliti {I'ipi.strclliis), 39, 40, 90.
aiistroriparius ( Vesperrilio), 20, 60.
Ixthaininsh ( Vespertilio). 39,41, 101-102.
Barbastella, 17.

barbastellus (Vespertilio). 17.
beelisteiiiii (Vespertilio), 16.
belli (Scotopbiliis), 21.
blossevillei (Scotophilus), 15.

borealis (Atalapha), 106.
borealis ( Laeiurus), 39, 41, 105-108.
borealis (Vespertilio), 21. 105.
braelmiaiius (Taphozous), IS.
Ijrachyotus, 13, 55.
brevirostris (Vespertilio), 21.
calcaratus (Vespertilio), 21.
calif oniicHS (J/i/ofis). 39. 40, 5(). 69-72.
ealifornicus (Vespertilio), 21, G9.
caiiacciiiii (Comastes). 13.
carolii (Vespertilio), 22, 38, 60.
caroliuensis (Vespertilio), 22, 96.
carolinensis (Vesperugo), 39, 01.
Catcorus. VJ, 95.
caiidata (Atalaplia). l:i.
caiiriniis {Jliiotit), 39, 40, 56. 72.
Clialinulobut!. 104.
I chrysonotus (Vespertilio), 23, 77.
ciliatus (Vespertilio) 14.
clliolabniia (Myotis), 39, 40, .56, 72-73.
ciliolabruni (Vespertilio), 23, 39, 72.
ciiierea (Atalapha), 40, 112.
cinereus ( Lasiiirui-), 40, 41, 105, 112-114.
cinereus (Vespertilio), 23, 112.
Cuephieus, 13, 95.
Cnepliaiophiliis, 13, 85.

ferriigineiis. 13.

niaeellus, 13.

noctivagaua, 13.

pellucidus, 13.
Coiuastes, 13, 55.

capaccinii, 13.

dasyeuenie, 13.

linmopbilus, 13.

megapodius, 13.
Cori/norhinus. 13. 49-50.

wncrei'". 13. 38, 40, 51-52.

pallescen.f. "8. 40, 52-53.

toimscndii, 38, 40, 52, 53-54.
erassus (Vespertilio), 23.
ereeks (Ve.spertilio), 24.
erejiuscularis (Nycticea), 24, 118.
crepiiscularis (Nycticejus), 40, 119.
cubanug (Xycticeitis), iO, 41, 120-121.
cubanus (Vesperus), 24, 120.
cubensis (Scotophilus), 24, 102.
cubensis ( Venpertilio), 39, 41, 102, 103.
cubensis (Vesperugo), 102.
cyanopterus (Vespertilio), 24.
cynocephalus (Nycticea), 25.
dasyeuenie (Comastes), 13.

131

132

INDEX.

(lasyciu'me (A'espertilio), 13.
JDaiypterus, 13, 54, 115-116.

intermedius, 40, 41, 116-118.
daubeu'onii (VeH]>ertilio), ID.
di.scolor (Vespertili"), 19.
discolor ( V^esperiigo), 14, 19, 20.
di.scolor (Yesperiis), 15.
dotnesticus ( Vespcrtilio), 25.
dutertreus (Vespertilio), 25, 102.
ega (Atalaplia), 13.
erjregia (Atalapba), 13.
Eptesicu-s, 13, 18, 95.
nielanops, 14, 30, 96.
nielas, 30.
iiiydas, 14, 32.
erytlirodactjlus (Ve.spertilio), 25, 90.
Enderma, 14, 46.

maculatum, 14, 38, 40, 46-49.
evoHs (MyotU), 23, -39, 40, 56, 77-80.
evotis (Vespertilio), 25, 39, 77, 80.
exili.s (A'espertilio;, 25, 69.
ferrugineiis (Cnephaiophilus), 13.
frantzii (Atalaplia), 25, 40, 111.
funebris (Lasiiirus), 26. 106.
fnseata (Atalapba), 26.
fuscus ( Adelouycteri.s), 39, 96.
fiiscus (Scotopbilus), 39, 96.
fnfcus ( Vesperlilio), 26, 39, 40, 96-99.
fuscus (Vcsperus), 39, 96.
georgianns (Scotophilus), 39, 90.
georgianus (Vespertilio), 26, 29, 90.
georgianus (Vcsperugo), 39, 91.
gracilis {Khogecssa), 40, 41, 123, 126-128.
greeuii (Scotopbihis), 27, 96.
gryphus (Vespertilio), 27, 39, 60, 75.
heiisbawii (Vespertilio), 27, 39, 69.
hesjients {rijiistrellns). 39, 40, 88-90.
hesperus (Scotopbihis), 27, 39, 88.
besperns (Vesperugo), 39, 88.
Histiotus, 14, 46.

niaeulatus. 14, 30, 46.
humcralis (Xrjcticeitis), 16, 40, 41, 118-120.
humeralis (Vespertilio), 16, 27, 118.
Hypexodoii, 14.
mystax, 14.
Hypsugo, 14, 87.

incantus (Vespertilio), 28. 56, 59.
ijiteriuedia (Atalapba), 13, 40, 116.
interiiicdius (Dasy2>terus), 40. 41, 116-118.
iutennedius (Lasiurus), 28, 40, 116.
Isotus, 14. 55.

keenii ( 2ryotis), 39, 40, 56, 77.
keenii (^'espertilio), 28, 77.
krascbeninikowii (Vcsperugo). 14.
ktihlii (Scotox'hilus), 17.
kublii (Vesperugo), 15, 20.
lanceolatus (Vespertilio), 28.
Lasionyctciis, 14. 54, 85.

noctivag-ins, 39, 40, 86-87.
Lasiurus, 14, .54, 105, 115.
horeaUs, 39, 41, 105-108.
cinereus, 40, 41, 105, 112-114.
funebris, 26. 106.
intermedius, 28, 40, 116.
mexicanvs. 40, 41, 105, 111-112.
noveboracensis, 39, 106.

Laxiurus, pfeifferi, 39, 41, 105, 1 10.
seminolus, 39, 41, 105, 109.
teliotii, 39, 41, 105, 110-111.
lasiurus (Vespertilio), 28, 105.
lasurus (Vespertilio), 29, 105.
lecontii (Plecotus), 29, 51.
leibii (Vespertilio), 29.
leisleri (Pterygistes), 87.
leisleri (Vesperugo), 20.
leporinus (Vespertilio), 18.
leptiirus (Vespertilio), 17.
leucippo (Vesperugo), 20.
leuci])pe (Vesperus), 15.
limnopbilus (Comastes), 13.
longicrus (Myotis), 39, 40, 56, 64-65.
longicrus (Vespertilio), 29, 39, 64.
lueifiigvs (Myotis), 38, 40, 56, 59-63.
lucifugus (Vespertilio), 29, 38, 59, 60.
luacellus (Cuepbaiopbilus), 13.
macleayii (Scotopbilus), 30.
macrojius (Vespertilio), 30, 39, 66. 67.
macrotis [Corynorhirms), 13, .38. 40, 51-52.
macrotis (Plecotu.s), 13, 30, 38, 51.
macrotis (Synotus), 38,51.
maculatum (Eudcrma), 14, 38, 40,46-49.
maculalus (Histiotus), 14,30,46.
Marsipobvmns, 15, 20, 104.
maurus (Vesperugol, 14.
Mcgaderma, 18.
niegalotis (Vespertilio), 30.
melanops (Eptesicus), 14,30,96.
niegapodius (Comastes>, 13.
melanorbmns (Vespertilio), 30,39,61).
melauotus (Vespertilio), 30.
nielas (Eptesicus). 30.
merriami (A'esperugo), 31,88.
Meteorus, 15, 95.
mexicana (Atalajiba), 31, 111.
mexicanus (Lasiurus), 40, 41, 105, 111-112.
mcxicanus (ilyotis), 39, 40, 56, 73.
7iiexicanus (\'espertilio), 31, 73.
minutilla, (Rhogecssa), 123,125.
miradoreniis (Vespertilio), 39,40,99-100.
miradoreusis (Scotopbilus), 31,99.
mouacbus (Vespertilio), 31,106.
inoiiticola (Vespertilio), 31.
nwrinus (Tespcrtilio), 14, 15, 18, 19.
iiiydas (Eptesicus), 14,32.
mydas (Vespertilio), 32.
Myotis, 15, 54, 55-56.

alascensis, 39, 40, 56, 63-64.

albescens, 12, 59, 84.

caUfornievs, 39, 40, 56, 69-72.

catirinus, 39, 40, 56, 72.

ciliolahrum, 23, 39, 40, 56, 72-73.

evotis. 23, 39, 40, 56, 77-80.

keenii, 39,40,56,77.

longicrus, 39, 40, 56, 64-65.

hicifugus, 38, 40, 56, 59-63.

mexicanus, 39, 40, 56, 73.

myotis, 13, 15.

nigricans, 12, 39, 40, 56, 74-75.

saturatus, 39, 40, 56, 68-69.

suhulatus, 36, 39, 40, 56. 75-76.

thysanodes, 39, 40, 56, 80-85.

velifer, 38, 40, 56-59.

INDEX.

133

MyotU, villosissimus, ]2.

yumancnsis 39, 40, 5t), 66-6S.
myotis (Myotin), 13, 15.
niyotis (Vespertilio). 15.
mystaciims (Vespertilio), 13, 17.
niystiix (Ilypexciloii), H.
iiiy.stax (X(ictilio). 32.
niystax (Vespertilio), 32.
Naimugo, 15, 87.
iiatliusii (Vcsperugo). 20.
iiatiereri (Vespertilio), 14, 15.
ninricanf! (Myotis). 12, 30, 40. 56, 74-75.
nigricans (Vespertilio). 32, 69, 74.
iiil.ssoiii (Vesperugo), 20.
iiilssuui ( Vesperii.s), 15.
iiitidus (Vespertilio), 32,39,60.69.
X.cliU', 18.

nij-stax, 32.

novcljoraceiisis. 26.
noctivagans (Ciiephaiopliilus), 13.
noctivagans (Laiiionycterix), ;!9, 40, 86-87.
nottivagan.s (St-otophilus), 39. 86.
noctivagans (Vespertilio). 14, ^2,86.
noctivagans ( Vesperugo), 3?, 86.
Xoctula, 15,95.
noctnla (Ptcryyistef,), 87.
noctiila (Vesperugo), 20, 104.
Noctulinia, 87.

novcboracensis (Atalapba), 39, 106.
uoveboraecusi.s (Lasiurus), 39,106.
novcboracensis (Xoctilio), 26.
novcboracensis (Vespertilio), 32, 105.
noveboracus (Vespertilio), 32. 105.
Xycticea. 16, 118.

crepiisc'iilaris, 24. 118.

cynocepbalus, 25.
Xyc'.iccius, 15, 54, 118.

cubanufi,40. 41, 120-121.

huvieralii, 10, 40, 41, 118-120.

tesselatus, 10.
Nycticejus, 10, 118.

crepuscularls, 40, 119.
Xycticeus. 10, 118.
Xycticeyx, 10, 118.
Xyctikstes, 16.

Si'rutiims, 10, 35.
Xyctitherivm, 16.

priicus, 33.

velox, 37.
Nyctopliiliis, 42.
Nystactes, 16, 55.
ohscunm (Pipistrellus), 39, 40, 93.
obscurus (\'espertilio). 33, 69.
oregonen^is (Vespertilio), 33, 69.
Pacliyotus, 15, 16.

paci/icvt {Antrozovs), 38, 40, 45, 46.
palkfcens {Corynoiiiintis), 38, 40. 52, 53.
pallifiKg (Antrozoui), 38, 40, 43-45.
pallidns (Vespertilio), 12, 33, 43.
parvvla (Phogecua). 17, 33, 40. 41, 123, 125-126.
Varviilus (Vesperugo), 40, 123. 125.
pelh.citlus (Cncpbaiophihis), 13.
per.'^picniatus (Vespertilio), 18.
ptcifif ri (Atalapba), 33, 39, ll(i.
rfeiffeii (I.athirva). ?9. 41, 10.3, 110.
pbaiops (Vespertilio). 30, 33, 90.

I'tpistielluf, 16, 54, 87-88.

abramug, 10.

australis, 39, 40, EO.

he>.perx(S, 39, 40, 88-90.

obscurus, 39, 40, 93.

pipistrellus, 88.

iubfiavus, 35, 39, 40, 88, 90-92.

vcrcecrucis, 39, 40, 88, r3-95.
pipistrelhis {Pipigtrcllna). ^9.
pipistrellus (Vesiiertilio), 16.
pipistrellus (Vesperugo), 15, 20.
riecofiita;,4h46.
Plecolut, 17, 18, 49.

auritus, 140.

lecontii, 29, 51.

iuacroti.s, 13, 30, 38, 51.

ratinesquii. 34.

townsendii, 13, 37, 53.

velatns, 14.
priscus {Nyctitherivm) , 33.
propinqxnis (Veapertilio), 39. 40, lOO-lol.
propinquus (Vesperugo), 30, IdO-lOl.
propinqiius (Vespcrus). 33, loO.
proterus (Vespertilio), 87.
pruinosus (.Scotopbilus), 15.
pruinosus (Vespertilio), 34, 112.
Pterojiui, 18.
PteryyUtes, 20. 87.

leulcri, 87.

noctida, 87.
pulverulentua (Vespertilio), 34, 80.
ratinesquii (Plecotus), :'4.
Rhogeesm 17.54,122-123.

alleni. 20, 40. 41, 123, 128-129.

gracilis, 40, 41, 123. 120-128.

miiivUlla, 123, 125.

parvula, 17, 33, 40, 41, 123. 125-120.

tumida, 17, 37,40, 41, 123-125.
Rhogoessa, 17, 122.
rubellus (Vespertilio), 34, 105.
rubra (Vespertilio), 34, 100.
rut'us (Tapbozous), 17.
riifus (Vespertilio). 34, 100.
salaril ( Vespertilio), 34.
saturatus (Myotis), 39, 40, 50, 08-6;).
savii (Vesperugo), 20.
savii ( Vesperus), 15.
Scotopihihts, 17.
Scotophilus, 85, 87, 95.

bellil, 21.

blossevillei, 15.

cubensi-s. 24, 102.

fuscus, ;;9, 90.

georgianus, 39, 90.

grcenii, 27, 96.

besperus, 27, 39, 88.

kuhlii, 17.

uiacleayii, 30.

iniradorensis, 31, 99.

noitivagans, 39,86.

liruinosus, 15.
Selysius, 17, 55.
seininola (Atalapba), 35, 109.
seminolusiLayiuriis), 39, 41, 105, 1;;9.
septentrionalis (Ves])ertilio), 35, 75.
serotimts (Xyctileste.s), 16,35.

134

INDEX.

sirotiniis {Vespertilio), 99.

serotimis (Vesperugo), 20,39,95,06.

serotinus (Vesperus), 13.

sicula (Atalapba), 13.

siiasnia (Vespertilio), 18.

siiectrum (Vespertilio). 18.

siibjiants {FipistreUus). 35, 39, 40. 88. 90-92.

subtiavus (Vespertilio), 35. 90.

itibulatia (Myotis), 36. 39. 40, 56, 75-76.

.snbulatns (Vespertilio), 29,36, 39, 75.

Synotus, 17, 49.

macrotis, 38, 51.

towiisendii, 38. 52.
Taphozous, 17.

braclimanus, 18.

rufus, 17.
teliotis (Atalapba). 36. 39, 110.
teliotis (Lasiurus), 39. 41. 105, 110-111.
tenuidorsalis (V.esi)ertilio). 36, 69.
tesselatus (Xycticeius), 16.
tesselatus (Vespertilio), 36, 106.
th!/ia7wdes (J/i/ofi's). 39, 40, 56, 80-85.
toirnsendii ( Corynorhiniig), 38, 40, 52. 53-54.
townseiulii (Plecotns), 13. 37.53.
to-wnsendii (Sjnotus), 38, 52.
tumida (BhogeVssa). 17. 37, 40, 41, 123-125.
ursimis (Vespertilio), 37, 96.
Tampynts, 18.
vampyrus (Vespertilio), 18.
velatus (Plecotns), 14.
relifer {2I;iotis). 38. 40, 56-59.
velifer ( Vesi>ertilio). 37, 38, 39, 56, 80.
relox {yyctitheriiim}, 37.
vercBcrucis (Piputrelhis), 39, 40. 88, 93-95
vera>crucis (Vesperugo), 37,93.
Tesperides, 18, 85.
Vespertilio. 18, 54, 55, 95.

affinis, 20, 3S.

agilis, 73.

albescens, 20, 39, 64, 66.

alhigularis. 20, 39, 41. 104.

alcytboe, 15, 95.

arquatus. 20. 96.

auduboni, 20. 86.

auritus, 18.

austroriparius, 21, 60.

baha)iH'nsis. 39,41, 101-102.

barbastellus, 17.

becbsteinii, 16.

borealis. 21, 105.

brevirostris, 21.

calcaratiis, 21.

californicus, 21, 69.

carolii, 22, 38. 60.

carolinensis, 22, 96.

cbrysoiiotus, 23, 77.

ciliatus, 14.

ciliolabrnm, 23. 39,72.

cinereus, 23, 112.

crassus, 23.

creeks, 24.

cubeiisis, 39, 41, 102-103.

cyanopterns, 24.

dasycneme, 13.

daubentonii, 13.

discolor, 19.

VeiperliUn, doniesticus. 25.
diitertreiis, 2."), 102.
erytlu oclactylns, 25, 90.
evotis. 25, 39,77,80.
exilis, 25. 69.
fuseuK, 26. 39. 40, 96-99.
georgianiis, 26, 29, 90.
grypbus, 27, 39, 60, 75.
bensbawii, 27, 39, 69.
liunieralis, 10.27.118.
incaiitus, 28, 56, 59.
keenii, 28, 77.
lanceolatus, 28.
lasiurns, 28, 105.
lasurus, 29, 105.
leibii. 29.
leporinus, IS.
lepturns, 17.
longicrus, 29, 39, 64.
lucifngns, 29,38,59,60.
niacropus, 30, 39, 66, 67.
megalotis, 30.
raelanorbinus, 30, 39, 69.
melanotus, 30.
niexicanns. 31, 73.
miradoremis. 39, 40, 99-100.
numacbus, 31, 106.
nionticola, 31.
imifinus. 14. 15, 18. 19.
mydas, 32.
myotis, 15.
niystacinus, 13. 17.
niystax, 32.
nattereri, 14.
nigricans. 32. 69, 74.
nitidus. 32.39,66,69.
uoctivagans, 14,32.86.
noveboracensis, 32, 105.
noveboracus, 32, 105.
obscurus, 33, 69.
oregonensis, 33, 69.
pallidus, 12, 33, 43.
perapicillatns, 18.
pbaiops, 30, 33, 95.
pipistrellus. 16.
propinqmis. 39,40, 100-101.
proterus, 87.
pruiuosiis, 34, 112.
piilrernlentns. 34, 86.
nibellus, 34, 105.
rubra, 34, 106.
rufus. 34. 106.
salarii, 34.

septentrionalis, 35, 75.
serotinus. 99.
spasma. 18.
spectrum. 18.
subflavus. 35, 90.
.subulatus, 29, 36, 39, 75.
tenuidorsalis, 36. 69.
tesselatus. 36, 106.
urBinus, 37, 96.
vampyrus. 18.
vehfer. 37,38,39,56,80.
villosissimus. 16.
virginiauus,37.

INDEX.

135

Veiipertilio. volans. 38, 69.

yumanensis. 3S. 39, 66. 69.
Vespertilionidce, 41.
Veirpertilionince. 41. .54.
Vespenigo, 19. 8.">. 87. 9.5.

albigulari.s 39. 104.

albolimbatus. 20.

aristiiipe. 20.

carolinen.sis. .39. 91.

cnbensis. 102.

discolor. 14. 19. 20.

georjianns. 39, 91.

lie.*perus. 39. 88.

kra.scheninikowii, 14.

kablii, 15, 20.

]ei8leri. 20.

leucippe. 20.

roaarus. 14.

merriami. 31, 88.

nathnsii. 20.

uattereri. 14, 15.

niUsoni. 20.

noctiragans, 39. 86.

noctala, 20, 104.

Vesperngo. parroluis. 40, 123. 125.

pil»i»tr<-lln.«, 15. 20.

propinquus. 39. 100-101.

.sarii,20.

serorintw. 20. 39. 95, 96.

vera-cmcLs. 37. 93.
Tespeni.-i, 20. 95.

albignlari-s. 15, 20. 104.

ari.stipi)e. 15.

cubana.s. 24. 120.

•Uscolor. 15.

fn.-«-n.-. 39. 96.

leacippe, 15.

nilssoni, 15.

propmqnn.s, 33. lOO.

savii. 15.

serotinns, 13.
viUogiigimusi {ilyotii), 12.
Tiilo.«is.«ijnn9 (Vespertilioi. 16.
^nrginianiis ( Vfesx)ertilio>..37,
volan.s ( Vespf-rtilio) 38. 69.
yumanengia {Mfiotiift. 39. 40, 56, CC-Go.
ynmanensis ( Vesi)ertlIio), 38, 33, C6, CD.

PLATE I.

[One iiml oiie-lialf times uatural size.]

Fig. 1. Myoiis relifer (J. A. Allen). Patzcnaro, Micboacau, Mexico.
(No. 52179, U. S. Nat. Mus.)

2. Mijotis caVifoinicus (•Audubon ct Bacbman). Nicasio, Cal.

(No. 1512, Meiriam collection.)

3. Myoiis niimaiiensis (H. Allen). Tulare, Cal.

(No. 30709, U. S. Nat. Mus.)

4. Nycticdiis humeralis Kafin('8(]ue. Brownsville, Tex.

(No. 52613, U. S. Nat. Mus.)

5. Myoiis ihysanodes Miller (topotype). Old Fort Tc.jon, Cal.

(29824, U. S. Nat. Mus.)

6. Myoiis eroiis (H. Allen). Bull Lake, Wyoming.

(No. 55846, U. S. Nat. Mus.)

7. Ithogeiissa gracilis Miller (type). Piaxtla, Puebla, Mexico.

(No. 70694, T;. S. Nat. Mus.)
X. L'hoyei'ssa tiimida H. Allen. Colima, Mexico.

(No. 52065, U. S. Nat. Mus.)
9. Corynorliinits macroiis ioansendii (Cooper). Gold Beacb, Oregon.

(No. 88542, IT. S. Nat. Mus.)

10. Antro^oiis 2)aUidiis (Le Coute). .Sycamore Creek, Texas.

(No. 24155, U. S. Nat. Mus.)

11. Eudervia macnlatuvi (.1. A. Allen) (type). Veutura County, Cal.

(No. :j;ii';. Am. Mus. Nat. Hist., N. Y.)

12. lihogei'ssa (jracilis Miller (type). Piaxtla, Puebla, Mexico.

(No. 70694, \:. S. Nat. Mus.)

13. lihogeiissa iumida H. Allen. Colima, Mexico.

(No. 52065, U. S. Nat. Mus.)

14. Nyciiceius humeralis Ratinesqne. I5rowusville, Tex.

(No. 52613, U. a. Nat. Mus.)
136

North American Fauna, No. 13.

Plate I.

1. Myotis velifer.

2. Myotis californicus.

3. Myotis !/iiinanensis.

4. Nyrficeiii.'i Jnimeralis.

5. Mi/otis tliyaanoden.
0. Myotis (Totis.

7. RhoyeHssa gracilis.

8. Rhogeessa tumida

9. Corynorhinus macrotis townsendi.

10. Antrozous pallidus.

11. Euderma maculatum.
\9. Rhogeessa gracilis.
1.3. Rhogeessa tumida.

14. Nycticeius humeralis.

PLATE II.

[One and oiic-balf times uatiu-al size.]

Fig. 1. Miioiis californiciis (Aiidubou & Bachmaii). Nicasio, Cal.
(No. 1.512, Merriani collection.)

2. Mi/oiis iinmatiensis (H. Allen). Tulare, Cal.

(U. S. Nat. Mas.)

3. Mi/ofls ei'otiff (H. Allen). Bull Lake, Wyoming.

(No. 55846, U. S. Nat. Mns.)

4. Mi/otis erotis (H. Allen). Perote, Vera (!ruz, Mexico.

(No. 88.541, U. S. Nat. Mns.)

5. MijoHh Ihijsanodes Miller (type). Old Fort Tejou, Californi£

(No. 29827, IT. S. Nat. Mus.)

6. Mjiotis relifer (J. A. Allen). Patzcnaro, Michoacan, Mexico.

(No. 52282, U. S. Nat. Mus.)
138

North American Fauna, No. 13.

Plate II.

1. Mijol is calif ornicus.

2. MyoTis yumanenxis.

3. Myotisevotis (.Bull Lake, Wyo.).

4. Myotis evotis fPerote, Mexico).

5. Myotis tliyscniodes.

6. Myotis velifer.

PLATE III.

[Two- thirds natural size.]

Fig. 1. Flecotiis auritus (Linn.). Hollsteig, Baden, Germany.
(No. 4495, Miller collection.)

2. Corijnorh'iuns macroiis pallescens Miller. Owens Lake, Cal.

(Xo. 28954, U. S. Nat. :\[us.)

3. Euderma maciilatum (J. A. Allen) (type). Ventura County, Cal.

(No. f ;Ji?, Am. Mus. Nat. Hist., N. Y.)

4. Lasiitriis einereus (Beauvois). Vegas Valley, Nevada.

(No. 27976, U. S. Nat. Mus.)
140

North American Fauna, No. 13.

Plate III.

1. Plecotus auritus.

2. Coryiiorhinus niacrotis pallfsceiis.

:i. Eudemia maculatuin.
■i Lasiuras cinereiis.

North American Fauna, No. 14.

U. S. DEPARTMENT OF AGRICULTURE
DIVISION OF BIOLOGICAL SURVEY

FORTH AMERICAN FAUIA

]sro. 14

[Actual data of publication, April 29, 1899]

t^Zi-

NATURAL HISTORY OF THE TRES MARIAS ISLANDS, MEXICO

General Account of the Islands, ^vith Reports on Mammals and

Birds. By E. W. Nelson
Reptiles of the Tres Marias, liy Lkoxhakd Stejneger
Notes on Crustacea of the Tres Marias. By Mary J. Rathbun
Plants of the Tres Marias. By J. N. Rose
Bibliography of the Ties Marias. By E. W. Nelson

Prepared under tlie direction of

Dr. C. HART MERRIAM

CHIEF OK DIVISION OF BIOLOGICAL SUIIVEY

WASHINGTON

GOVERNMENT PRINTING OFFICE
1899

^AaJ^/L' J>S 0<A^A.yiyvUAjjL^

570

LETTER OF TRANSMITTAL.

TJ. S. Department of Agriculture,

Division of Biological Survey,

Washington, D. C, January 25, 1899.

Sir : I have the honor to transmit herewith for publication as North
American Fauna No. 14 a report by E. W. Nelson on the natural his-
tory of the Tres Marias Islands, Mexico. These islands are the largest
off the west coast between Cape St. Lucas and the Isthmus of Panama,
but have seldom been visited, and very little is known of their fauna
or flora. For several years Mr. Nelson has had charge of the field
work of the Biological Survey in Mexico, and in May, 1897, visited
the Tres Marias. During the course of this visit he made a thorough
collection of birds and mammals and also secured specimens of reptiles,
fishes, mollusks, crustaceans, and plants, so that his report contains a
fairly complete account of the natural history of the islands. In work-
ing up the material collected, Mr. Nelson has had the assistance of
several well known naturalists in the United States National Museum
and United States Fish Commission, who have prepared reports on
special groups, as credited in detail on page 13.

Mention should be made also of the unfailing courtesy and interest
of the Mexican Government in the investigations conducted by the
Biological Survey in Mexico. Letters have been furnished by ofiQcials
in the City of Mexico, and by the late Mexican minister in Washing-
ton, Sefior Don Matias Romero, which greatly facilitated the work in
various ways, and on the occasion of the visit to the Tres Marias
euabled Mr. Nelson to borrow a large boat at San Bias and secure
comfortable quarters on the islands.

Several attempts at agriculture have been made on the Tres Marias
Islands, but the results have thus far been unsuccessful, owing to the
dry climate and the scarcity of permanent water. Corn and beans
have been grown on a small scale, but the crops suffer from the severe
storms which occur at certain seasons. Experiments have been made
with a view to utilizing the native species of agave for fiber and mes-
cal, and the cultivation of cotton has also been tried without success.
Recently it has been proposed to establish an American colony on one
of the islands for the purpose of growing coffee, bananas, Australian

3

4 LETTER OF TRANSMITTAL.

chestnuts, aud date palms, and to engage in the manufacture of
banana and chestnut flour. Such a scheme, Mr. Nelson tells me, could
only result in failure, as the islands are entirely unsuited to growing
these products. It therefore seems desirable to publish at once all the
information in the possession of the Department, for the purpose of
making it available to those who may be interested in the islands or
their products.

Eespectfully,

C. Hart Merriam,

Chief J Biological ISurvey.
Hon. James Wilson,

Secretary of Agriculture.

CONTENTS.

Page.

General d«^scription of the Tres Marias Islands, Mexico. By E. W. Nelson 7

Mammals of the Tres Marias Islands. By E. W. Nolson 15

Birds of the Tres Marias Islands. By E. W. Nelson 21

Reptiles of the Tres Marias Islands. By Leonhard Stejneger 63

Notes on Crustacea of the Tres Marias Islands. By Mary J. Rathbun 73

Plants of the Tres Marias Islands. By J. N. Rose 77

Partial Bibliography of the Tres Marias Islands. By E. W. Nelson 93

ILLUSTRATIONS.

PLATE.
Map of the Tres Marias Islands Frontispiece.

FIGURES.

Page.

1. Erythrina lanata Rose 81

2. Euphorbia nelsoni Millspangh 89

5

No. 14. NOETH AMERICAN FAUNA. April 29, 1899.

GENERAL DESCRIPTION OF THE TRES MARIAS ISLANDS, MEXICO.

By E. W. Nelson.
INTRODUCTION.

The Tres Marias islands are situated off the west coast of Mexico,
about Go miles west from the port of San Bias. These islands have
been known since early in the history of the i^ew World, and in 1532
were named Las Islas de la Magdalena by Diego de Mendo/a. Many
of the early explorers sailed about them, and Dampier states that
they were familiar to the buccaneers who visited these shores. They
are mentioned by several of the later voyagers, especially the English
exploring- expeditions which visited the west coast of Mexico in the
first half of the present century. During all this time, however, they
remained uninhabited and nothing definite was known or published
concerning their character or products. It is said at San Bias that the
first men who lived upon the islands were bandits, who took refuge
there, and had a secure retreat from which they harried the mainland
settlements for several years. Finally, the abundance of Spanish
cedar became known, a settlement of woodcutters was established on
Marie Madre, and this island has since been continuously inhabited.

Col. A. J. Grayson, a naturalist who lived for many years on the west
coast of Mexico, was the first to publish any detailed information about
the islands.' Most of this information is contained in the various paj)ers
embodying the results of his three trij)s to the Tres Marias in 1865, '06,
and '67, published by himself, George K. Lawrence, and W. E. Bryant.
In 1881 Alphonse Forrer, a natural history collector, spent some time on
Maria Madre collecting specimens for the British Museum, but no de-
tailed account of his work has been published. No other naturalist is
known to have visited the islands until the spring of 1807. In April of
that year Mr. E. A. Goldman and I visited the port of San Bias for the

' Mr. John Xantus, who spent several years subsequent to 1859 on the west coast of
Mexico, was supposed to have visited the islands, on account of several specimens
of birds which he sent to the Smithsonian Institution, labeled "Tres Marias Islands,
1861." But as no one else has collected any of these species, and as Xantus sent in
no birds which have been taken by others on the islands, it is safe to concludt; that
he did not visit the Tres Marias.

7

8 NORTH AMERICAN FAUNA.

purpose of outfitting an expedition to the Tres Marias. A letter to the
collector of customs at San Bias, kindly furnished me by the Mexican
Minister in Washington, the late Don Matias Romero, proved of the
greatest service. The collector of customs rendered every assistance
in his power, including the loan of a large open boat 25 feet long, and
a letter to his deputy which secured us very pleasant quarters in the
custom house on Maria Madre. While iireparations for the trip were
in progress a party from Socorro, N. Mex., consisting of Prof. 0. L.
Herrick, his son Harry, and Dr. T. S. Maltby arrived at San Bias, also
bound for the Tres Marias, and we made the trip together. On the
evening of April 28 the boat crept out of the lagoon, and by the aid of
a faint land breeze edged slowly oft" shore. The islands came in sight
the next morning, but it was impossible to reach them for several days,
owing to calms, head winds, and the lack of a keel to the boat. The
stock of water was on the point of exhaustion when Maria Madre was
finally reached, three days later, on the afternoon of May 2.

The landing was made at the settlement at the head of a shallow
bay on the east side of the island. Our letters secured a cordial
welcome from the customs inspector and the agent of the owner of the
islands. In a couple of hours the outfit was snugly installed on the
broad upper verandas of the custom house, where our headquarters
were located. Collections were made near this place, the island trav-
ersed both on foot and horseback, and on May 20 a boat trip was made
to the north end of the island and across to San Juanito. On May 23
the party returned to the settlement, and two days later proceeded to
Maria Magdalena, where camp was made near the beach for four days.
On May 29 we crossed to Maria Cleofa, where we remained two days,
and then started, May 31, on the return to the mainland. The wind was
fair, and a quick trip was made, San Bias being reached on the evening
of June 1.

When Colonel Grayson visited the islands, in 1865, he found a settle-
ment on Maria Madre, but the other islands uninhabited. In the
spring of 1897 there was a branch custom-house, with three inspectors,
at the main settlement on Maria Madre, which had supervision of the
shipment of salt and Spanish cedar. The settlement contained about
twenty-five families, all of whom, except the customs inspectors, were
in the service of the owner of the islands, Senora Gil de Azcona,
who lived in the city of Tei)ic, on the mainland. In May and June
the workmen are employed in salt-making at a lagoon near the south
point of the island, where there is a small group of houses. The rest
of the year they are occupied in cutting cedar and hauling it to the
beach for shipment. The available supply of this valuable timber is
now approaching exhaustion. Subseqnent to Grayson's visit a settle-
ment of woodcutters was made on the northeast side of Maria Magda-
lena, and a number of houses were built and a field cleared. We
found the place deserted, the houses in ruins, and the field overgrown
with thorny bushes.

GENERAL DESCRIPTION OF THE TRES MARIAS ISLANDS. \)

The amount of laud suitable for agriculture upon the islands is very
limited and forms but a small percentage of tbe total area. A few
cattle are raised on Maria Madre, but tbe scanty berbage and great
scarcity of water during the long dry season limit this industry to tbe
most insignificant proportions. Tbere is a small field near tbe settle-
ment, wbere coarse grass is grown for stock. Attempts bave been
made to grow corn and beans to supply tbe residents, but tbe fierce
summer storms of wind and rain, called 'cbubascos,' wbicb beat tbe
crops to tbe ground, bave rendered tbese efforts futile. At present
all food supplies are brougbt from tbe mainland. A number of years
ago a house was built and a field cleared and fenced near tbe north
end of tbe island for tbe purpose of growing cotton. A warehouse
was also built at tbe main settlement, but after a trial tbe owner was
forced to abandon the industry, tbe field and house were deserted, and
the place is now overgrown with bushes. Subsequently it was proposed
to utilize the agaves, which grow abundantly near tbe north end of the
island, for fiber and for distilling from their fleshy bases the alcoholic
product known as 'mescal.' Machinery was obtained, but the owner
died before the industry was exploited.

In winter tbe weather is dry and pleasant, and small coasting steam-
ers stop every now and then to take on wood for fuel, and sailing
vessels call for Spanish cedar or, in spring, for salt. In May the
inhabitants are obliged to lay in a stock of provisions sufficient for
several months, as they are practically cut off" from communication
with the mainland during summer, when the islands are avoided on
account of tbe storms that sweep over them. Many objects drift out
jfrom the Gulf of California after storms and are cast up on tbe shores.
In September, 1890, a great tornado of wind and rain swept over
northern Sinaloa and tbe Gulf of California; the coast lowlands were
devastated by tbe flooded rivers, and crops and forests were alike
overwhelmed and swept to sea. In May, 1897, tbe shore of Maria
Madre was still strewn with cornstalks, driftwood, and other wreckage
that had been stranded after this storm.

Our obligations to the collector of customs at San Bias and his agent
on Maria Madre have already been mentioned, and acknowledgments
are due also to the owner of the islands, Senora Gil de Azcona, whose
letter procured us the use of horses and other courtesies.

PHYSIOGRAPHY.

The Tres Marias are situated between latitude 21° and 22° and
longitude 106° and 107° (see frontispiece). Between the islands and
the mainland, 20 miles offshore, lies Isabel Island, only about a mile
long and 150 feet high. The soundings in the channel between the
mainland and tbe islands gradually deepen to less than 300 fathoms,
but just west of tbe grouj) the sea bottom drops rapidly to more than
1,500 fathoms. Tbe absence of a deep channel shows that they are
continental islands, as distinguished from the oceanic Eevillagigedo
group, farther west.

10 NORTH AMERICAN FAUNA.

The Tres Marias group comprises four islands, San Juanito, Maria
Madre, Maria Magdalena, and Maria Oleofa, arranged in a northwest
and southeast direction. Maria Madre, the largest, measures about
8 by 15 miles, and rises over 2,000 feet above the sea, North of this,
and separated from it by a channel 4 miles wide and 5 or 6 fathoms
deep, is San Juanito, an islet 3 or 4 miles in diameter and about 100
feet high. Kext southeast of Maria Madre is Maria Magdalena,
roughly triangular in outline and 7 or 8 miles across, with its central
summit rising to an altitude, of about 1,500 feet. A shallow channel
8 miles wide separates it from Maria Madre. Southeast of Maria Mag-
dalena lies Maria Cleofa, the last of the group. It is irregularly
rounded in outline, about 3 miles across, and its altitude is apparently
much less than 1,320 feet, as given on the charts. The channel between
the two last-named islands is about 12 miles wide and much deeper
than the others.

"With the exception of San Juanito, which is nearly flat with a narrow
border of low bluffs along the north shore, the islands are mountainous
and rise in successive slopes from the shore to the culminating point
near the center. The interior of Maria Madre is occupied by a moun-
tainous ridge extending almost the entire length of the island, but
descending to a gently sloping area near each end. The eastern side
of the island has the longer slope, while the westward or seaward face
is much more abrupt, thus corresponding with the formation of the
mountains parallel to the coast on the adjacent mainland. Both slopes
of the island are scored at intervals with canyons which usually descend
in a nearly direct line to the sea. Maria Magdalena and Maria Cleofa
are occupied by a central mountainous elevation, from which canyons
descend in all directions to the sea. The northeastern points of both
these islands are low, flat, sandy areas of limited extent, and the west-
ern faces are rocky and precipitous. Permanent fresh water is very
scarce on all the islands. There are three little streams on Maria
Madre, which sink several miles from the sea during the dry season,
and one each on Maria Magdalena and Maria Cleofa.

The relative situation of the islands, with the narrow, shallow chan-
nels between them, shows conclusively that at one time they formed a
single island at least 45 or 50 miles long, and at a still earlier stage they
must have been connected with the mainland. One of the strong-
est proofs of this former connection is shown by the correspondence
between the fauna and flora. The breaking down of the original island
into several smaller ones and the evident continuous encroachment of
the sea appear to indicate that the subsidence is still in progress. The
country back of the coast on the mainland was, within a comparatively
recent period, the scene of great volcanic activity, and the Tres Marias
bear evidence of having undergone various oscillations in level. On
Maria Madre there are great beds of marine deposits, hundreds of feet
above sea level, containing quantities of shells and corals of species
now living along the shore. Isabel Island, near the mainland, is of

GENERAL DESCRIPTION OF THE TRES MARIAS ISLANDS. 11

volcauic origin and exhibits similar evidence of having- once been a
much larger island which is now sinking. Apparently it consists mainly
of the remains of an old volcano, and a small crater still occupies the
center of the island. Although no craters were seen on the Tres
Marias, yet there are lavas and other volcanic rocks on all the islands,
but a large part of the formation is made up of other rocks elevated by
the volcauic uplift.

FAUNA.

The Tres Marias, like the adjacent coast, lie within the Arid Tropical
life zone. The evidence furnished by the fauna of the former connec-
tion of the Tres Marias with the mainland is as follows: Six species
of land shells were obtained, which, according to Dr. William H. Dall,
are widely distributed on the mainland. These sj)ecies are Folygyra
ventrosula Pfr., Orthalicus undatus Brug., Orthalicus undatus melano-

cheilus Yal., LameUaris ?, Opeas suhula Pfr., and Glandina turris,

Pfr. A fresh- water fish taken on Maria Magdalena and Maria Cleofa
has been identified by Prof. B. W. Evermann as Agonostomus nasiitus
Giinther, a common species on the mainland. In fresh-water pools on
Maria Magdalena two or three individuals of another small fish were
seeu, which were very similiar to common mainland species of Atvaous,
and undoubtedly belong to this or a closely allied genus. Six of the
seven species of lizards inhabit the mainland, and only one is peculiar
to the islands; the mud turtle and crocodile are also found on the
mainland, as are the eight species of snakes. Concerning the reptilian
fauna Dr. Stejneger remarks: "Thus most of the species are common
on the opposite mainland and generally distributed over tropical Mexico
and Central America. Then again it seems as if the species are i)rac-
tically identical on all the islands of the group. This would indicate
a comparatively recent severance of these islands from each other, as
well as from the opposite mainland of Mexico."

The birds and mammals seem to have been more susceptible to modi-
fying influences than other forms of life. Thirty-six species of resident
laud birds were found on the group, of which twelve are identical with
those on the mainland, and twenty-four can be distinguished specifically
or subspecifically. We found ten species of indigenous mammals, seven
of which, acccording to Dr. Merriam, are peculiar to the islands, but
closely related to sjjecies living on the mainland.

ANIMALS PECULIAR TO THE TRES MARIAS.

So far as known, the following species and subspecies (with the excep-
tion of Compsothlypis insularis) are peculiar to the islands :

MAMMALS.

Marmosa insularis Merriam. Procyon lotor insularis Merriam.

Oryzomys nelsoni Merriam. Ehogei'ssa imrvula H. Allen.

Peromyscus madrensis Merriam. Glossopliaga mutica Merriam.
Lepua graysoni Allen.

12

NORTH AMERICAN FAUNA.

Columha flavirosirls madrensls Nelson.
Leptotilit, capitalis Nelson.
Biiteo horcalis fumoSHS Nelson.
rolyboriis cheriwaij pallidiis Nelson.
Psittacula iyisularis Ridgway.
Trogon amhiguus goldmavi Nelson.
Drijohates scalaris graysoni Baird.
Nyctidromus albicolJis insularis Nelson.
Amazilia graysoni Lawrence.
lache Imvrencei Ridgway.
Platypsaris agJaioc, insularis (Ridgway).
My io^yagis placet! s minimus Nelson.
Icterus graysoni Cassin.

Cardinalis cardinalis mari(v. Nelson.
Piranga hidentata Jlammea (Ridgway).
Virco Jlaroriridis forreri (Von Madnrasz).
Virco hypochryseus sordidus Nelson.
Compsothlypis insularis (Lawrence). Oc-
curs also on the mainland near San IJlas.
Granatellus francesca; Baiid.
Thryotlioriis lawrencii (Ridgway).
Thryothorus lawrencii magdalena Nelson.
Melanotis cwrulescens longirostris Nelson.
Myadestes ohscurus insularis Stejneger.
Merula graysoni Ridgway.

Cneniidophorus mariarum Giinther.

KKPTILES.

FLORA.

The islands were visited near the end of the long dry season, when
most of the herbaceous plants were withered and lifeless, but repre-
sentatives of 13G species, largely shrubs and trees, were secured.

The general appearance of the vegetation was the same as that in
similar situations on the mainland. Among the most notable plants
were the Spanish cedar (Cedrela), three species of wild fig {Ficus), two
of Pithecolohium, five of Solnnum, two of Ipomoca, a Passijiora, cassias,
euphorbias, a large agave, a large cereus, and two oj)untias.

On San Juanito the vegetation is largely made up of bushes and
scrubby trees 8 to 15 feet high, with many agaves on the sandy southern
end. Agaves are very numerous also on the northern end of Maria
Madre. On the latter island the forest is rather low and scrubby near
the shore, but increases in luxuriance farther up the slopes, especially
along the bottoms and sides of the canyons, where Spanish cedars, wild
figs, and several other trees attain a large size. In its primeval con-
dition, before the advent of woodcutters, it must have presented a fine
example ot tropical forest growth. Now, only a few specimens remain
to show what the original condition must have been. Along the sum-
mit of the island the dense forest is made up of slender-trunked trees,
called 'palo prieto' by the natives, which I was unable to identify.
On Maria Magdalena the conditions were similar to those on Maria
Madre, but a larger percentage of the original forest still remains intact,
although the Spanish cedars are mainly gone. Maria Cleofa is more
rocky and sterile, and the trees are stunted and brushy. Several spe-
cies found on the other islands appeared to be wanting here. The
report on the plants shows that the flora of the islands is very similar
to that of the mainland, and the fact that several new species were
found may be due to our imperfect knowledge of the mainland flora.

GENERAL DESCRIPTION OF THE TRES MARIAS ISLANDS. 13

PLANTS DESCKIBED FROM THE TRES MARIAS.

MgipMla pacifica Greeuman.

Btlopirone nelsoni Greenman.

Bujcus pubescens Greenman,

Cordia insularis Greenman.

Erythrina lanata Rose sp. nov. (also on

mainland).
Euphorhia nelsoni Millspaugli.
Euphorbia subcwriilea tresmariw Millsp.

var. nov.

Gilibertia msularis Rose sp. nov.
Pilocarpus insularis Rose, sp. nov.
Ternostroemia maltbya Rose sp. nov. (also

on mainland).
Zanihoxylum insularis Rose sp. nov.
Zanthoxylum nelsoni Rost, sp. nov.

SUMMARY.

The following statement shows the number of species of animals and
plants now known from the Tres Marias :

Laud mammals 11

Birds 83

Reptiles 18

Fresh- water fish 2

Fresh-water shrimp 1

Land mollusks 6

Plants 136

ACKNOWLEDGMENTS.

Much of the value of this report is due to the cordial cooperation of
several eminent specialists. Through the courtesy of Mr. F. V. Coville,
curator of the National Herbarium, Dr. J. N. Rose, assistant curator,
was enabled to prepare the report on the plants. Dr. Leonhard Stej-
neger, curator of the division of reptiles of the National Museum,
Dr. William H. Dall, honorary curator of the division of conchology,
and Miss Mary J. Rathbun, assistant in the division of invertebrates,
reported on the Tres Marias material; and Prof. B. W. Evermanu,
ichthyologist of the United States Fish Commission, kindly identified
the collection of fishes from the islands and the adjacent mainland.
Finally, I wish to express my great indebtedness to Mr. Robert Ridg-
way, curator, and Dr. Charles W. Richmond, assistant curator, of the
division of birds in the National Museum, for having so freely i)laced
at my disposal, not only the material in their charge but also their
knowledge of tropical American birds.

I

MAMMALS OF THE TRES MARIAS ISLANDS.

By E. W. Nelson.

Mammals are not numerous either in species or individuals upon the
Tres Marias. So far as known, they number but eleven species, of
which seven are peculiar to the islands; one is introduced, and the
other three are widely ranging bats. A sea lion and two species of
porpoise were found near the shores, and whales were reported to
occur during certain seasons. As with the birds, one of the most
unaccountable features of the mammal fauna is the absence of a num-
ber of species that are common on the adjacent mainland. Considering
the i^rimitive condition of the islands, it is diificult to explain the
presence of field mice, the pigmy opossum, rabbit, and raccoon, while
the large gray opossum, nasua, skunk, fox, coyote, deer, xieccary,
squirrel, and various small rodents of the adjacent mainland remain
unrepresented. The Tres Marias mouse was rather common above 200
feet on all of the larger islands ; the rabbit was very numerous near
the north end of Maria Madre, on San Juanito, and in some places on
Maria Magdalena, and two species of bats were abundant in caves on
Maria Madre. Aside from these species, mammals were uncommon
and diflicult to find. One cause of their general scarcity may be the
very limited supply of ijermanent fresh water, and the absence of small
species from a broad belt near the shore was easily accounted for by
the abundance of carnivorous crabs.

The mammals obtained by our party have been identified by Dr. C.
Hart Merriam, who has described the new forms and given critical
notes on other si^ecies.^ Of the land mammals taken, five were new
and two, Lepus graysoni and Rhogeessa parvula, had been previously
described. We failed to secure two species of bats {Myotis nigricans
and Lasiurus horeaUs mexicanns) which were taken by Mr. Forrer. Not-
withstanding the fact that collections were made in several branches
of natural history, I feel confident that representatives of all the resi-
dent land mammals were secured, but it is quite possible that future
work may add other bats to the present list.

ANNOTATED LIST OF SPECIES.

Marmosa insularis Merriam. Tres Marias Pigmy Opossum.

Marmosa intiiilaris Merriam. Proc. Biol. Soc, Washington, XII, pp. 14-15, Jan.
27, 1898. Type from Maria Madre Island.

These pretty little opossums were not found except in the high inte-
rior of Maria Madre, between 1,200 and 1,800 feet above sea level, where

1 Proc. Biol. Soc. Washington, XII, pp. 13-19, 1898.

15

16 NORTH AMERICAN FAUNA.

they were apparently ratber commou about the wild fig trees in the
forest and were feedin^^ upon the figs. They may occur also on the
other islands, especially upon Maria Magdaleua. Two men living on
the island described the nests of these animals as globular masses of
dry leaves and small plant stems, lined with shreds of softer vegetable
matter. The nests are built in the forks of bushes, from 3 to 8 feet
from the ground, and have the entrance on the lower side. One of
the men found a nest situated as described and about 3 feet from
the ground. He saw the owner peering out of a hole near the lower
side, but as he approached the head vanished, and the entrance was
suddenly closed by the opossum drawing some of the nest material
across it. The nest was quickly thrust into a game bag, and when
examined was found to contain a female opossum and a number of
young clinging to her fur with their feet and tails twined closely
about hers. The weight of the young was so great that the parent
could only walk very slowly.

Oryzomys nelsoni Merriam. Nelson's Rico Rat.

Oryzomys nelnoni Merriam. Proc. Biol. Soc. Washington, XII, p. 15, Jan. 27,
1898. Type from Maria Madre Island.

This rice rat is probably a rare species, as only a few specimens were
secured after much trapping. They were found only in damp places
near springs about the summit of Maria Madre, about 1,800 feet above
sea level. This seemed the most suitable location for them on account
of the juicy herbaceous vegetation mingled with the undergrowth.

Peromyscus madrensis Merriam. Tres Marias Mouse.

Peromyscua madrensis Merriam. Proc. Biol. Soc. Washington, XII, p. 16, Jan. 27,
1898. Typo from Maria Madre Island.

This is the mostly widely distributed and probably the most numer-
ous rodent. Specimens were taken on the three large islands, but
its occurrence on San Juanito, where land crabs are very numerous,
is doubtful. They were generally distributed over the forest-grown
slopes bordering the shore, above the belt infested by crabs. On
Maria Madre they were most common about the wild fig trees near the
summit (1,500 to 1,800 feet), where the pigmy opossums were secured.
Here their burrows entered the ground under logs or projecting roots,
but elsewhere these mice were found living beneath rocks and small
ledges. They are apparently restricted to the forest, and while
nowhere so abundant as were the rabbits in one place near the north
end of Maria Madre, yet they were much more generally distributed.

Mus rattus Linn. Black Rat.

These rats were found in small numbers about the houses and dis-
tributed over the forested parts of Maria Madre and, as on the main-
land of western Mexico, we found only the gray form.

Lepus graysoni Allen. Tres Marias Cottontail.

Lej)U8 graysoni Allen, Mon. N. Am. Rodentia, pp. 347-348, 1877. Type from Tres
Marias Islands (undoubtedly from Maria Madre).
The cottontail is abundant in some places on San Juanito, Maria
Madre and Maria Magdalena, and was reported to occur on Maria

MAMMALS OF THE TRES MARIAS ISLANDS. 17

Cleofa. They were very numerous about a deserted ranch on the north
side of Maria Magdalena, but were rather scarce elsewhere on that
island. We found them extraordinarily abundant and surprisingly
tame about old fields on an abandoned ranch at the northern end of
Maria Madre. Some were killed with stones near camp, and it would
have been easy to kill over a hundred in a morning. They would sit
in their forms among the bushes while one peered at them from a dis-
tance of a few feet, and when driven out into au open space they often
sat quietly while the camera was brought up and focussed within a
short distance. The old fields at this ranch had been long abandoned
and were covered with a scattered growth of bushes, which seemed
more suitable for the rabbits than the forested areas, where they
occurred much more sparingly. The cottontails frequented the wood
roads leading from the shore up over the forested slopes, and after 3
o'clock in the afternoon could be found sitting quietly in little open
places in the undergrowth waiting for the nearer approach of sunset
before coming out into the roads.

The skin of these rabbits was surprisingly delicate, and it was
diflScult to skin them without tearing it in many places. It was found
almost impossible to carry a specimen by the hind legs even a short
distance without having the skin tear and slip where it had been
grasped by the hand.

It is strange that the rabbits are not more abundant on the islands,
considering the fact that the raccoon is the only predatory mammal,
and that the few red-tailed hawks and caracaras are the only birds
that prey upon them.

Procyon lotor iusularis Merriani. Tres Marias Raccoon.

Procyon lotor insularis Merriam. Proc. Biol. Soc. Washington, XII, j). 17, Jan-
uary 27, 1898.

The raccoon was rather common on Maria Madre and Maria Magda-
leua, but no sigus of them were seen on Maria Cleofa, where, however,
they may occur. In May they were feeding on wild figs and other
fruits and on the crabs, which were very abundant near the shore.
Every morning freshly made raccoon tracks were seen in trails leading
from the seashore to higher parts of the islands, but the animals
usually passed our traps without paying the slightest attention to the
bait. They were semi-diurnal in habits and several were seen in
the woods in broad daylight. One afternoon one was seen crossing
the bed of a dry wash near the northern end of Maria Madre, and
instead of trying to escape through the woods it climbed a wild-fig
tree on the bank and stood looking down from a horizontal branch
until shot.

Zalophus californianus (Lesson). Sea Lion.

A large seal or sea lion, called ' lobo mariuo ' or sea wolf by the Mexi-
cans, was reported to occur at several places on the rocky shores of
Maria Magdalena and Maria Cleofa. We first heard of them before
13950— i^o. 14 2

18 NORTH AMERICAN FAUNA.

leaving San Bias and again ui)ou reacliing the islands. It was evident
that the sea lions had been hunted for sport by previcfas visitors until
they had become comparatively scarce and are now in a fair way to
become extinct. After learning the location of the most frequented
places on both islands, we visited them under the guidance of a tortoise-
shell hunter who was very familiar with the shore, but we saw only a
single sea lion. It was on a rocky islet off the shore of Maria Cleofa,
and took to the water and disappeared before we could get a shot.
Our guide said that sometimes the sea lions leave the islands for a few
days, and this may account for the failure to find them about their
usual haunts. The consensus of opinion among the residents of Maria
Madre was that these animals are now very scarce. Formerly they
were found in many places; but at i)resent a rocky i^oint on the north-
west side aiid a jutting reef on the south side of Maria Magdalena
and some islets west of Maria Cleofa are the only landing places used.
It is possible that the Guadalupe Island fur seal {Arcfocephalus
toicnsendi Merriam) may also occur at times about the islands.

Rhogeessa parvula H. Allen. Tres Marias Rhogecssa,

Rhogeessa parvula H. Allen. Proc. Acad. Nat. Sci. Phila., 1866, p. 285. Type from
the Tres Marias.

These little bats were rather common on Maria Madre, where they
live in the forest and fly at dusk along the trails and about small open
places. At times they appear in such situations in broad day. Two
were killed while flying up and down a trail in the brilliant sunshine in
the middle of the forenoon, and I saw one hawking for insects among
the tree tops along a trail two hours before sunset. As a rule, however,
they only come out Avhen it is too dark for one to see more than an
indistinct form as they flit about among the trees. A few were also
seen on Maria Magdalena.

My Otis nigricans (Maximilian). Maximilian's Black Bat.

According to Mr. Oldfleld Thomas, a specimen of this bat was taken
on the Tres Marias by Mr. Forrer.' We took none, and they probably
occur on the island only as stragglers.
Otopterus mexicanus (Saussnre). Big-eared Bat.

A colony of over a hundred big-eared bats was living in an old ware-
house at the settlement on Maria Madre, and others were found in sev-
eral caves situated in various parts of the island. The warehouse where
these bats were found had a large open window and wide cracks, so
that it was quite light inside, yet they were found hanging from the
ceiling and roof, in j)lain view, and evidently had lived there a long
time. The specimens were mostly females heavy with young.

Glossophaga mutica Merriam. Tres Marias Glossophaga.

Glossophafja mutica Merriam. Proc. Biol. Soc. Washington, XII, pp. 18-19, Jan-
uary 27, 1898. Type from Maria Madre Island.

This was by far the most numerous bat on Maria Madre, where it was
found in every cave sufficiently deep to be dark. One cave was among

' Biologia Ccntrali- Americana, Mammalia, 206, 1881 (under y'es])crtilio nigncans).

MAMMALS OF THE TRES MARIAS ISLANDS. 19

some huge projecting rocks lying at the water's edge, near the settle-
ment. Manj^of the females collected contained large embryos. These
bats were feeding on the fruit of the wild fig.

As surmised by Dr. Merriam,' the record of Cheer onycter is mexicana
from these islands, given by Mr. Thomas in the Biologia, proves to be
referable to the present species. In reply to a letter of inquiry, Mr.
Thomas states that he discovered the mistake in identification too late
to correct it in the Biologia, and agrees with Dr. Merriam in referring
his specimen to G. mutica.

Lasiurus borealis mexicanus (Saussure). Mexican Red Bat.

Forrer added this species to the fauna of the Tres Marias as recorded
by Mr. Thomas.- We did not see any red bats, and I doubt their being
found on the islands except as stragglers from the mainland. Bats
are such wide ranging animals it is to be expected that several addi-
tional species will eventually be found to occur on the islands.

? Phocaena communis (Lesson). Common Porpoise.

Pori)oises supposed to belong to this species were common around
the shores of the Tres Marias and also in bays and mouths of
streams or lagoons along the coast of the mainland. They were always
seen in the belt of shallow discolored water within a short distance of
the shore. As soon as blue water, with a depth of over 40 fathoms,
was reached, the other porpoise {Prodelphinus lotigirostris) was encoun-
tered. The common porpoise was seen in schools of 10 to oO or 40 indi-
viduals swimming in loose order. At Maria Madre they came into the
shallow bay in front of the settlement in the early morning and followed
close along shore.

Prodelphinus longirostris (Gray). Long-nosed Porpoise.

In the blue water between the mainland and the islands these por-
poises were very abundant in schools of from 100 to 200 individuals.
They are much slenderer and more graceful animals than the preceding
species. While swimming about their feeding places at sea they were
accompanied by swarms of terns, gannets, and shearwaters. On one
occasion, while crossing to the islands, a school of about 200 porpoises
came directly toward us and passed under and on all sides of the boat.
While they were passing, the water was broken into foam on every
hand by their glistening black bodies, and overhead swarmed a shriek-
ing crowd of sea birds. Mr. Goldman made a fortunate rifle shot and
killed two of them, but one sank before it could be harpooned.

'Proc. Biol. Soc. AVashington, XII, pp. 13-19, footnote, January, 1898.
^Biologia Centrali-Americana, Mamm., p. 205, footnote.

BIRDS OF THE TRES MARIAS ISLANDS.

By E. W. Nelson.

The present paper is based mainly upon tlie birds found on the Tres
Marias, but for the sake of completeness the results of our work on
Isabel Island have also been introduced.' The situation of Isabel
Island between the mainland and the Tres Marias renders its bird life
of peculiar interest in the present connection, Mr. Xantus sent speci-
mens of birds to the National Museum labeled 'Tres Marias, 1861,'
but only one of these can be an authentic island species, and it seems
almost certain that Xantus did not visit the islands.

Colonel Grayson's notes on his three visits to the group and his
trip to Isabel Island were published by George N. Lawrence in the
'Proceedings' and 'Memoirs of the Boston Society of Natural History,'
while the descriptions of new birds in his collections appeared in various
publications and are mentioned in the bibliography (see pp. 93-94).
Grayson constantly refers to the 'Tres Marias Islands,' but the internal
evidence of his writings, in addition to the information given me by the
inhabitants, indicates that all of his work was done on Maria Madre.

Mr. A. Forrer visited Maria Madre in 1881, but the publication of
Vireo Jiavomridis forreri by Von Madarasz and a few notes in the
'Biologia Cen trail- Americana' and in some of the British Museum
Catalogues are all we know of his work there.

As already stated in the general introduction, our work was done on
Isabel Island on April 22 and 23, on Maria Madre from May 2 to 25,
and six days were spent working about Maria Magdalena and Maria
Cleofa. It is quite certain that the bird fauna of Maria Madre is
now fairly well known, and it will be advisable for anyone visiting this
group in the future to give attention chiefly to the two smaller islands.
It is certain that a large proportion of the birds found on Maria Madre
occur also on Maria Magdalena, but some of the species living in the
dense forest at higher altitudes on these islands probably do not occur
in the more scanty forest of Maria Cleofa.

At present 83 species and subspecies of birds are kn6wn from the
Tres Marias, and further observations will, no doubt, add to the list
various stragglers from the mainland. The bird fauna may be grouped
under the following headings: Resident land birds, 3C species or sub-

' The notes iu the following pages refer to Isabel Island only when so stated.

21

22 NORTH AMERICAN FAUNA.

species. Visitant land birds, 26 species or subspecies. Resident water
fowl, 13 species. Visitant water fowl, 8 species.

Of the 36 resident species or subspecies of land birds all but 5 were
observed by Colonel Grayson, These exceptions are: Melopelia leucop-
tera, Tyrannus melanchoUcns couchi, Ornithion imherbe, Vireo fiavoviridis
forreri, and Thryothorus lairrencii magdalence.

Twenty-four of the 36 resident land birds are specifically or subspe-
cifically distinct from their mainland representatives. Of this number
12 were described from Grayson's collections, 1 from Forrer's, and 11
from our own. A study of our collections from the islands, and near
San Bias on tbe mainland, brings out the interesting fact that several
species from the latter district show a decided approach to their island
representatives. This is very marked in Gompsotlilypis which is very
nearly the same at San Bias as on the islands. The Polybortis and
Platypsaris from that locality seem to be intermediate between the
island races and the birds of the mainland. Specimens of Thryothorus
felix from the same part of the coast are much nearer T, tawrencii than
they are to typical T. felix.

Among the 24 species or subspecies of land birds peculiar to the
islands 15 are larger than their relatives of the nearest mainland.
These are Golumhaf. madrensis, Leptotila capitaUs, Psittacula insularis,
Dryohates s. graysoni, Nyctidromus a. i7i,sularis, Amazilia graysoni,
Icterus graysoni, Cardinalis c. mar ice, Firanga h. Jfarmnea, Vireo f.
forreri, Vireo h. sordidus, Compsothlypis insularis, Qranatellus francescce,
Thryothorus lawrencii and MerUla graysoni.

Six of the island birds average smaller than their mainland repre-
sentatives. These are Polyhorus c. pallidus, lache lawrencei, Platyp-
saris a. insularis, Myiopagis p. minimus, Melanotis c. longirostris, and
Trogon a. goldmani. The two first named are generally smaller, but
Platypsaris a. insularis has a longer tarsus, Myiopagis p. mittimus a
longer bill and tarsus, Melanotis c. longirostris a longer bill, and Trogon
a. goldmani a longer bill and tarsus.

Although Compsothlypis insularis also occurs in a limited area along
the coast, I have considered it as a typical island species. The
difference in size between island birds and their mainland repre-
sentatives varies greatly, being slight in some and very well marked
in others. Nyctidromus a. insularis is a larger bird than alhicollis
proper, but has a shorter bill and tarsus. Among the birds peculiar
to the islands Thryothoriis laivrencii magdalenw and Myadestes o. insu-
laris are almost the only ones which do not show more or less well-
defined differences in size from their nearest mainland relative; a series
of the first named, however, may show that it also differs.

One of the most puzzling features of the fauna of these islands is
the absence of various land birds found on the adjacent mainland.
Although the physical conditions appear so much like those of the
mainland, yet some change must have occurred to upset nature's fine
balance and render these isolated areas unsuitable for many species.

BIRDS OF THE TEES MARIAS ISLANDS. 23

The death by starvation of the Louisiana Tanagers on Maria INIadre
Island (p. 52) is an example of the manner in which the island fanua
may be maintained in its present state. As the climatic conditions on
the islands and on the mainland are very similar and the vegetation
nearly alike, this paucity of species presents one of the curious prob-
lems of distribution.

It would be hard to find an equal area of similar country on the
mainland, near San Bias, where so few species of land birds could be
found. The only reasonable explanation seems to be the scarcity of
water and the long, dry season, which combine to reduce the food sup-
ply and perhaps render the country unsuited to some species. It was
very surprising to find a total absence on the islands of such com-
mon and widely spread mainland genera as Conurus, Moniotus, Piaya,
Campephilus, Melanerpes, Myiozetetes, Cissolopha, Cyanosjnza, Pipilo,
Pyrgisoma, Saltator, and others. The absence of Pipilo is especially
unexpected, for this genus is represented on Socorro Island, which
lies very much farther at sea off the same part of the coast.

ANNOTATED LIST OF SPECIES.

? Brachyrhamphus brevirostris (Vigors.) Short-billed Murrelet.

Bracliyrliamplius brevirostris was described from San Bias and B.
hypoleucus from Cape St. Lucas. Colonel Grayson mentions having seen
'guillemots' at Isabel Island (Mem. Boston Soc. I^at. Hist., II, p. 318,
1874) and ofi" theTres Marias group (Proc. Boston Soc. Nat. Hist., XIV,
p. 288, 1871). This led me to anticipate finding at least one of the
species there, and it is with some disappointment that I have to record
our failure to see either species about the islands, although I watched
for them constantly. From this experience I am inclined to think that
they breed only along the coast of Lower California, and visit these
islands sporadically.

Larus argentatus smithsonianus Coues. American Herring Gull.

A single immature specimen was taken on San Juanito Island May
22, and a few others were seen. These birds were fiying back and forth
along a strip of beach where a large colony of blue-footed gannets
were breeding, and the gulls probably had an eye on the nesting
ground for the i)urpose of capturing any unprotected eggs. They were
noted singly a few times along the shores of the Tres Marias and at
Isabel Island. No fully adult individuals were seen.

Larus heermauni Cassiu. Heermann's Gull.

On April 23 a fine adult bird of this species was shot on the shore of
Isabel Island. In company with its mate it had harried a blue-footed
gannet into disgorging a number of small fish upon a rock at the edge
of the water, and was i)icking up the spoils by a series of little down-
ward swoops and hoverings. The gannet had shuffled into the water
and was making off, with backward glances at its tormentor, when I
drew near. These giills are bold and noisy aggressors when they wish

24 NORTH AMERICAN FAUNA.

to take advantage of the gannets, and about the breeding places of the
latter they feed largely at the public expense. But few of them were
seen about the islands — two or three pairs at Isabel and half a dozen
pairs about the Tres Marias. A nest, which had been occupied earlier
in the season, was seen on the ledge of a rocky islet off the shore of
Maria Oleofa May 30, and full grown young of the year were also seen
on the rocks.

Sterna maxima Boddaert. Royal Tern.

None were seen at Isabel Island, although they were not uncommon
during April along the mainland coast. During May they were seen
in small parties about the shores of all the Tres Marias group, where
they probably breed in very limited numbers. The only specimen
saved was taken May 31 from a flock of six which was coasting along
the beach at Maria Cleofa.

Sterna elegans Gambel. Elegant Tern.

Sterna (jaUricidata Lawr., Mem. Boston See. Nat. Hist., II, p. 317 (1874).

Colonel Grayson found these terns on Isabel Island, but none were
seen by us.

sterna fuliginosa crissalLs (Lawr.). Pacific Sooty Tern.

Haliplana fuliginosa var. crissalis Lawr. (ex Baird MS.), Proc. Boston Soc. Nat.
Hist., XIV, pp. 285, 301, June, 1871; Mem. Boston Soc. Nat. Hist., II, p. 318,
1874.

These handsome terns are common about Isabel Island where Colo-
nel Grayson found them breeding. My observations from the last of
April to the first of June led me to believe that at this season Isabel
Island is their central roosting point. During the week w^e were cruis-
ing about Isabel and the Tres Marias islands many flocks were seen.
From about noon until the middle of the afternoon or later the flocks
were generally flying directly toward Isabel at an altitude of from
50 to 200 yards above the water. This was noted also near the islands,
while we were crossing the straits between the Tres Marias, and oft'
the mainland near San Bias. Many of the birds were perched
along the top of an inaccessible rock just oft" Isabel, and were also
seen alighting on the cliffs of the northern and northeastern side of
the island, but the boat was too unwieldy for us to venture near enough
to closely examine these haunts. The birds have a peculiar shrill
cry which they often utter while feeding and when flying about at
night. The night before we landed on Isabel Island it was necessary
to anchor about midway between the island and tbe shore. The wind
blew strongly in the afternoon but fell at sunset, a dead calm ensued,
and heavy clouds overspread the sky. During the day only a few
sooty terns had been seen, but from about 9 p. m. until near daybreak
they were evidently much more numerous, for their cries were heard at
short intervals. Several times the notes were uttered directly overhead
and the birds seemed to be scarcely higher than the top of the mast,
where they apparently paused and hovered while they examined the

BIRDS OF THE TRES MA.RIAS ISLANDS. 25

boat with great curiosity. As they were heard every night while we
were at sea, it is evident that they were both diurnal and nocturnal
in habits.

They feed well out at sea, and were not found anywhere along shore,
except when they came to their roosting place on Isabel Island. There
were no signs of their roosting about the Tres Marias, although they
may roost on some of the outlying rocky islets. Grayson found
them in small numbers farther west, about the Revillagigedo Islands.
During our trip to the Tres Marias many schools of large fish were
encountered swimming close to the surface and constantly breaking,
often with such force and rapidity that the water boiled and foamed
over considerable areas. These schools of fish were commonly accom.
panied by flocks of sooty terns and gannets, which appeared to be
animated by the wildest excitement. The terns hovered over the
foaming sea, uttering shrill cries and darting down into the water, evi-
dently after food; and in the midst of the turmoil the blue-footed gan-
nets swam about, beating the water with their wings and adding to the
noise made by the terns and leaping fish. While on Maria Madre
I saw a flock of terns some distance off shore, and taking a canoe,
managed to get out to them, and directly in the course of the
school of fish they were accompanying. Letting the boat drift, I
stood up and watched the swarm go by. Thousands of large fish and
hundreds of terns and gannets passed the boat on every side, amid loud
cries from the terns, a rushing sound from the fish and gannets, and a
bewildering complexity of motion in sea and air that was intensely
exciting. This novel sight was so interesting that I came near losing
the chance to secure some of the birds.

These terns were seen also following schools of porpoises off shore —
in the latter case accompanied by the wedge-tailed shearwater. In the
passage between Maria Magdalena and Maria Cleofa a flock of sooty
terns was seen soaring in wide circles high overhead and finally start-
ing off' for their roosting i^'ace on Isabel Island.

The 'variety crissalis,' named in manuscript by Professor Baird and
published by Mr. Lawrence, was characterized as "having the under
tail coverts tinged with ashy, instead of being pure white." A series
of 17 specimens from the west coast of Mexico, and from widely scat-
tered islands of the Pacific and Indian oceans, agree in having the
posterior part of flanks, under wing coverts, and entire crissum dis-
tinctly ashy, not a single individual being white on these parts, as is
commonly the case with birds from the Atlantic and Gulf coasts of
North America. Unfortunately the series of Atlantic birds at hand
is very small, but there is little doubt that crissalis is a valid subspe-
cies. Birds from the west coast of Mexico, the Galapagos Islands,
and Hawaii agree in having an average shorter bill and tail than
those from elsewhere. Specimens from the Indian Ocean have even
a longer bill and tail than thos^ from the Atlantic, but are ashy below,

2G

NORTH AMERICAN FAUNA.

like those from western Mexico. Specimens from Ascension Island, oflf
tlie west coast of Africa, also liave a light ashy shade on the lower
tail coverts.

The following average measurements show the sizes of these birds
from various parts of their range:

Table of measurements of Sterna fuliginosus and Sterna f. crissalis.*

Num-

Name.

Locality.

ber of
speci-
mens.

Wing.

Tail.

Cul-
men.

Tarsus.

sterna fuliginosus . .

East coast of North America and
west coast of Africa.

7

288.1

151

42.8

23.5

Sterna fuliginosus

West coast of Mexico, Hawaii, and

10

288. G

143. 5

41.8

23.6

crissalis.

Galapagos Islands.

Sterna fuliginosus

Kruseustern Islands { west of Hawaii) .

2

292.5

203.5

39.5

24.2

crissalis.

Sterna fuliginosus

Glorioso Island (Indian Ocean)

3

2n2.6

192

43.3

23.8

crissalis.

* All measurements are in millimeters.

In the foregoing measurements the length of the tail is unreliable,
owing to its variability, on account of wear and other causes.

Anous stolidus ridg-wayi Anthony. Pacific Noddy Tern.

AnoHS stolidus Lawr., Mem. Boston Soc. Nat. Hist., II, p. 318, 1874.
Anous stolidus ridgwaiji Anthony, Auk, XII, p. 36, 1898.

Common the last of April on Isabel Island, and a few seen oft" the
Tres Marias during May. Between San Bias and the islands a
number of these birds were seen. We usually saw one or two indi-
viduals at a time, and did not find them in flocks anywhere except
when congregated on the rocks at their roosting i)laces. At sea they
usually flew close along the surface of the waves with long, graceful
wing strokes. From their dark color and habit of keeping close to the
water they were several times mistaken for black petrels.

They were found in considerable numbers on the ragged faces of
clifls and rocks along the northeastern point of Isabel Island, and were
very unsuspicious, iiermitting ns to approach quite near in the boat.
While perched on the black lava clift's, their dark color blended so
closely with the background that it was very difticult to distinguish
them, even when within fair gunshot. The day we left the island we
visited their resting place and fired a dozen or more shots while they
were on the rocks or flying about, but the noise of the reports did not
seem to give them much alarm. They would circle out a short dis-
tance, and, after hovering for a few moments over their killed or
wounded companions floating in the water, would return to the same
part of the cliff from which they had just been startled. They were
not heard to utter any notes, and the silence with which they would
suddenly appear out of the cliff* and then return and vanish again in
its gloomy face inoduced an uncanny eft'ect.

BIRDS OF THE TRER MARIAS ISLANDS. 27

Colonel Grayson found tliem nesting on the north end of Isabel
Island in April, 1869, and states that they were breeding in communi-
ties on shelving rocks beneath overhanging cliffs. The nests were
placed close together, but were inaccessible. A single egg was pro-
cured, however, which was white, with scattering brownish blotches,
most numerous about the larger end. This is undoubtedly the teru
which Colonel Grayson reports as replacing the sooty teru ou the
Kevillagigedo Islands, and which he described as being black, with a
hoiiry forehead.

Puflinus cuneatus Salvia. Wedge-tailed Shearwater.

J'ti^inua cuneatus Salvin, Ibis, 5tb ser., VI, p. 353, July, 1888.
rnjjinus huudseni Stejn., Proc. U. S. Nat. Mus., XI, p. 93, Nov. 8, 1888.

During our trip to and from the islands we saw 100 or 200 wedge-
tailed shearwaters. They were usually seen singly skimming along over
the sea, at an elevation af a few yards, making widely sweeping circuits
and pausing occasionally to pick up bits of food. When about midway
between Isabel Island and the Tres Marias we encountered several
schools of small porpoises of 150 or more individuals, which traveled in
close array, frequently gamboling about and playfully leaping high in
the air. A swarm of sooty terns followed the porpoises, and twice
when they passed near us I saw considerable numbers of these shear-
waters among the terns. Judging from the numbers, they must be
rather common in these waters, but none were seen near the islands.

This species was first described from specimens taken ou the Kru-
senstern Islands, in the Marshall Group, and Mr. A. W. Anthony made
the first record of its occurrence on the American side of the Pacific,
at the Revillagigedo Islands, during the summer of 1897 (Auk, XV,
Jan., 1898, p..39). As it is a species new to North America, a detailed
description of our specimen is given, in order to facilitate identification
in case other examples should be taken.

Description.— ^o. 156678, U. S. Nat. Mus., Dept. of Agriculture Goll.
Ad. S , off Maria Mandre Island, May 2, 1897. Collected by E. W. Nel-
son and E. A. Goldman.

Top and sides of head and neck grayish-brown; forehead, lores, and
space from latter area back beneath eyes and along sides of neck paler
or more ashy, thus edging the darker area of the crown and upper neck
with lighter. Back, including rump and upper tail coverts, mainly dark
brown, but mixed with numerous feathers of a decidedly grayish, almost
ashy, shade. These latter feathers undoubtedly indicate the color of
dorsal surface in fresh ]>.. imaged birds. Wings and tail blackish-brown.
Entire lower surface of body white, shaded with dingy ashy, darkest
on sides and palest along median portion. Under tail coverts mixed
dark brown and grayish-brown. Border of the wing along under side
brown or grayish-brown; under coverts white with a little flecking
of pale gray ou some feathers. In the flesh this specimen had a horn-
blue bill with flesh-colored feet and tarsi.

28 NORTH AMERICAN FAUNA."

Dimensions. — Wing, 293; tail, 135 (lengtli of middle pair of tail feath-
ers beyoud lateral pair, 47); culmen, 42; tarsus, 50; middle toe, with
claw, 59.

Hahiiat. — The range of this species is now known to extend across
the middle North Pacific from Japan to the west coast of Mexico.

Oceanodroma melania (Bonap.)- Black Petrel.

Common between Isabel and the Tres Marias. Black petrels were
by far the most numerous of the petrels seen, and outnumbered all the
others two to one. Three, and possibly four, other species were seen
on the way to and from the islands, but this was the only one secured.
They circled about in all directions, sometimes coming very near, but
nothing peculiar in their liabits was observed. They were quick to
see little fragments of fat thrown overboard while we were skinning
other waterfowl, and when the morsels were small enough ate them
greedily.

Phaethon aethereus Linn. Red-billed Tropic Bird.

Tropic birds are readily distinguished on the wing by their graceful
ternlike flight and long filamentous tail feathers. Many of them breed
on Isabel Island and in suitable places on rocky islets near San Bias
and about the Tres Marias. The last of April fresh eggs and nearly
grown young were found on Isabel, and by the last of May the young
on the Tres Marias had taken wing and few were to be seen, although
we were told by the tortoise shell hunters that many breed there earlier
in the season.

Soon after landing on Isabel, a tropic bird was found sitting on its sol-
itary egg at the end of a little hole in the rock close to the beach. The
hole was only about 15 or 18 inches across and about 3 feet deep, so that
there was no dififlculty in taking the bird by hand after a little maneu-
vering to avoid its sharp beak. During a stay of about twenty-four
hours on this island at least 20 nests containing eggs or young were
examined. A single Q,gg is laid directly on the rough rock or loose
dirt forming tlie floor of the nesting site, which is always located under
the shelter of over arching rock, but varies greatly in situation. The
inner ends of holes in clifl's facing the sea were favorite i^laces, but as
the number of such situations was limited, the birds were forced to
utilize small caves and even rock shelters. In one locality five or six
nests were placed on loose earth at the bottom of rock shelters so situ-
ated that I could walk directly up to them and pick up the birds.
Whenever a nest was approached the parent screamed and fought
viciously, rufiied its feathers and looked very fierce, but made no attempt
to escape. They protested with beak and voice when pushed about,
but as soon as I went away a few yards they would shuffle back to
resume their former position over the egg. The young, even when
quite small, were eciually fierce in resenting any intrusion. One nest
was found on the beach under the edge of some great rocks that had
fallen from the adjacent clift'. It was only 5 or G feet above high tide

BIRDS OF THE TRES MARIAS ISLANDS. 29

and would have been overlooked but for the angry cries of the old
bird when she heard me walking- over the roof of her habitation. At
sunrise the old birds were found sitting side by side at the mouths of
their nesting places waiting to enjoy the first rays of sunlight. Half
an hour later one of each pair started out to sea while the other resumed
its place on the nest. When disturbed on the nest their cries are very
shrill and strident, consisting of a series of short, harsh, clicking or
rattling sounds something like the noise of an old-fashioned watch-
man's rattle. The young are covered at first with fluft'y white down.
Before they are one-third grown the first plumage begins to appear,
and is very similar to that of the adults, except that the black barring
on the back is broader.

Sula -websteri Rothschild. Webster's Booby.

Sula hassana Grayson, Proc. Boston Soc. Nat. Hist., p. 302, 1871.

Sula wehsteri Rothschild, Bull. Brit. Orn. Club, VII, No. LIV, p. LII, 1898.

This booby is no doubt the Sula hassana reported from Isabel Island
by Grayson but not seen by us. Mr. Anthony found it the most abun-
dant species breeding on the Eevillagigedo Islands during the summer
of 1898.

Sula brevrsteri Goss. Brewster's Booby.

Sula hretvsteri Goss, Auk V, p. 242, 1888.

Brewster's boobies were very numerous on a small hill at one side
of the little bay where we landed on Isabel Island April 22, but there
were no signs of their breeding.- They came in from sea during the
first half of the afternoon and sat about on rocky parts of the shore
until nightfall. Scattered individuals were also seen about the ledges
and tops of the cliffs facing the sea. The following morning at day-
break they were congregated on the little hill already mentioned which
is probably their regular roosting place. About half an hour after
sunrise they began to start out to sea singly and by twos and threes
until all were off on the day's fishing expedition. A few were seen
about the rocks just off San Bias, and were said to breed ou the large
rock (Piedra Blanca) midway between Isabel and San Bias. Only a
few of these boobies were seen about the Tres Marias until an islet
was visited off the northwest shore of Maria Cleofa. This islet rises
from 150 to 200 feet above the sea, with cliffs on all sides. The sum-
mit is mainly rolling, with an elevated, sloping bench on one end At
this time. May 30, many thousands of boobies were breeding ou the bare
top of this rock. The eggs were laid directly on the surface, with no
sign of a nest. The sun was intensely hot and heated the rocks so that
they were uncomfortably warm to the touch. The birds did not sit
ui>on the eggs during the hottest hours, but while standing to avoid
contact with the heated rocks kept in such position that the eggs or
young were shaded from the sun, and thus had their vitality preserved.
While trying to secure photographs of this breeding ground a few of
the old birds flew away and it was surprising to see how quickly the

30 NORTH AMERICAN FAUNA. T

newly hatched young succumbed to the heat when the parents left them
exposed to the rays of the sun. The nests were spaced at intervals of
4 or 5 feet, so that the old birds were safely out of reach of one another.
Although so gregarious in their breeding habits, they appeared to
have but little regard for one another. It was amusing to see the sav-
age way in wbich the nest owners assisted intruders of their own kind
out of their territory. While we were walking among them some of
the birds would often waddle off to one side, and in so doing neces-
sarily trespassed on their neighbors. The latter at once raised a
hoarse shrieking and set upon the outsiders with wicked thrusts of
their beaks, which continued until the victims took wing and escaped.

We were also subjects of this proprietary rage, and had our legs nipped
every now and then, despite all efforts to walk circumspectly. Our
progress over the breeding ground was accompanied by a wave of
hoarse, nasal cries that sometimes became almost deafening. Many of
the birds were valiant upholders of their rights and sturdily refused
to leave their nests, which they defended vigorously, all the time utter-
ing loud cries of rage.

These birds show very little individual variation in color. As the
species is not well known the following descriptions are appended from
specimens taken on Isabel and Maria Cleofa.

Adult male. — Nearly entire head white, shading gradually on poste-
rior portion into drab of neck and then insensibly into dark, sooty brown
of back. On lower side of neck the drab becomes darkest at poste-
rior border, where it ends abruptly against the i)ure white of lower
parts. Bill light horn colorj gular pouch in life livid blue; feet green-
ish yellow — the latter varying in intensity.

Male in immature plumage. — Dorsal surface uniform dark brown,
slightly paler than back of adult; entire lower surface still paler and
more dingy brown. Feathers over much of body, especially about
head, neck, and lower parts, narrowly edged with grayish brown, giving
a faint wavy barring. Bill bluish horn color, with darker shade of
same about base and on gular pouch; feet and tarsi dull fleshy yellow;
iris greenish gray.

Adult feinale. — Head, neck all around, and back sooty brown; ven-
tral surface below neck white. Bill light horn color; a spot of leaden
bluish on lores; ba>>e of bill, gular pouch, feet, and tarsi grayish yel
low; iris pale grayish.

Average measurements of these birds from Isabel Island are as fol-
lows: S (5 specimens), wing 384.4; tail 189.6; culmen 93.6; tarsus 45.4;
$ (5 specimens), wing 416.6; tail 192.8; culmen 96.6; tarsus 48.8.

Nestlings a few days old are covered with fluffy white down. A male
bird of the previous year, which still retained the immature plumage,
was taken at Isabel on Aj^ril 23, and several others were seen.

BIRDS OF THE TRES MARIAS ISLANDS. 31

Sula nebouxii Milne-Edwards. Blue footed Booby.

Sula ])iiicaior Grayson, Proc. Boston Soc, Nat. Hist., XIV, p. 302, 1871; Lawr.,

Mem. Boston Soc. Nat. Hist., II, p. 316, 1874.
Sula nebouxii Milne-Edwards, Ann. Sci. Nat., Paris (Zool.), 6vme s^r., XIII, Art.

4, p. 37, pi. 14, 1882 (Chile).
Sula gossi Goss (ex Ridgway MS.), Auk, V, p. 241, July, 1888 (Gulf of Calif.).

Sida 7iehouxn is the most abundant species of booby occurring on
Isabel and the Tres Marias. On April 22 they were breeding abun-
dantly on the beaches and on a low flat area that covers a part of the
former island. They were common on the grassy beach at the landing
and thence back among the scrubby trees and bushes which form a
scanty growth over the flat. They were most numerous on the open
beach a little above high-water mark, but dozens of them were seen
with their eggs farther back among the bushes. Like the preceding
species, they fought and screamed savagely when approached. The
males usually flew away, but the females remained to give battle over
the nests, which were mere hollows in the earth, sand, or gravel. Not
a single young one was seen in the hundreds of nests on Isabel.

The sun- was excessively hot the morning of our arrival, and while
the men were landing the outfit, ropes were fastened between the tops
of some scrubby trees close to the beach and a piece of canvas spread
for an awning, under which the baggage was placed. An old booby
had her eggs in the sand within 3 feet of the edge of the sheltered area
and stood her ground unflinchingly while the men were at work,
keeping a wary eye on their movements and making vicious dabs
whenever a leg came incautiously within reach. Having arranged
camp, I went out exploring for an hour or so and returned with various
specimens, including the egg of a tropic bird, and found that one of the
boatmen had driven off the booby and thrown away her eggs. Wish-
ing to test the bird's discrimination, I placed the reddish-brown egg of
the tropic bird in the hollow where the two greenish- white eggs of the
booby had been, and sitting under the awning began to prei)are
specimens. In the course of half an hour the owner of the despoiled
nest returned and alighted 10 or 15 steps away near another deserted
nest, gave a look at the eggs in it, walked to still another, looked at it,
and then proceeded directly to her own nest and stopped. She looked
about and then down at the nest. The presence of the single
reddish- colored egg appeared to surprise her; she looked at it with
one eye and then with the other as if in doubt. An instant later the
feathers on her head and nape ruffled up and with a loud squawk of
rage she suddenly dashed her beak again and again into the strange
egg, breaking it to fragments in a moment. As soon as the egg was
demolished she took wing and disappeared out to sea. There was no
intention to sacrifice the tropic bird's egg in this experiment, so the
booby carried oft" the honors.

About 10 o'clock the following night a visit was paid to the nesting-
boobies. The night was calm, and taking a lighted candle I walked

32 NORTH AMERICAN FAUNA.

out a short distance to an opening in the bushes where there were
twenty or thirty nests. The females were found on their eggs with the
males standing close beside them. When the strange visitor appeared
in their midst the birds set up a continuous series of hoarse cries and,
like so many moths, seemed to become fascinated by the light. They
started up on all sides, and trooping within the circle of bright light,
began to run around me in a ring about 20 feet in diameter. They ran
in single file from right to left and presented a most ludicrous sight.
Occasionally one fell on its breast, whereupon the others scrambled
over the fallen bird until it regained its feet and rejoined the proces-
sion. One of the number was suddenly j^ossessed with a desire to run
around one of my legs, and, although seized by the head several times
and tossed out among its companions, persisted in returning to the
same place and continuing its gyrations. The next morning at day-
break the birds were seen standing in pairs by their eggs and remained
in this position until about sunrise, or a little after, when all of the
males went out to sea — usually in little parties of two to five or six.
They returned between 1 and 3 o'clock in the afternoon, and a number
of them flew directly to their mates and disgorged nl^me^ous small
fishes which the females ate greedily. These observations seemed to
show that the females did the incubating and the males provided the
food. As the neighboring waters do not abound in small fishes, the
boobies have to go in many cases from 10 to 30 miles to obtain their
daily supply. During a visit to San Juanito Island, the latter part of
May, many blue-footed boobies were found breeding on sandy beaches
at the south end of the island; many of the young were hatched and
some were more than half grown. Like the young of the tropic birds,
the young boobies uttered angry cries and fought savagely when
approached.

This species is found in Chile, on the Galapagos Islands, and north to
the island of San Pedro Martir in the Gulf of California. The type of
Sida nebouxii was obtained on the coast of Chile, and the tyj)e of S.
gossi came from San Pedro Martir.

The sexes are alike in color, but when standing together the males
may be readily distinguished by their smaller size and slenderer form.
In life the bill is leaden horn color, with its base and the gular pouch
leaden blue; the feet are bright blue. The downy young are pure
white. Four specimens measure as follows: S (1 specimen), wing, 410;
tail, 219; culmen, 108; tarsus, 51. ? (average of 3 specimens), wing,
438; tail, 219; culmen, 109.6; tarsus, 56.

Phalacrocorax sp. Cormorant.

Two or three cormorants seen at a distance were the only ones noted
during the trip. None were seen near Isabel nor on the rocks near
San Bias.
Pelecanus californicus Ridgway. California Brown Pelican.

A few pairs of brown pelicans were breeding on Isabel Island the
last of April. The nests were made of sticks and placed in the dense

BIRDS OF THE TEES MARIAS ISLAND.s. 33

tups of the scrubby trees growing- on the rocky inner slopes of the islaud.
They were found about all of the Tres Marias, but usually occurred singly
and were nowhere common. Two or three were seen fishing in the
breakers alongshore at the north end of Maria Madre.

Fregata aquila (Linn.)- Man-o'-war Bird.

Hundreds of man-o'-war birds were breeding on Isabel in April, and
on approaching the island many were seen soaring over the rocky sum-
mit. The first shot caused hundreds of others to take wing, and in a
few moments the air was swarming with them. They soared in con-
stantly intersecting circles, until the sky seemed covered with their
silhouetted outlines. So few had been seen alongshore near San Bias
that it was an interesting and unexpected sight. Upon landing, numer-
ous large, oval, and brilliantly red objects were seen in the tops of the
dark-green bushes along the slopes. These proved to be the gular
pouches of old male man-o'-war birds inflated to the size of a man's head,
the brilliant red color of the distended membrane making them very
conspicuous objects. It appeared to be a common custom of the birds to
sit quietly on the top of a tree for a long time Avith the pouches thus
distended and evidently serving as sexual ornaments. A few birds
were seen circling high overhead with their pouches fully inflated, but
as a general rule, when soaring, the pouches were closed.

The nests were built of sticks and placed in the tops of low trees
and stout bushes from near sea level to the summit of the island.
Three or four of these platform-like structures were found together in
some of the larger bushes. Many of the young were hatched and, when
able to stand alone, would do their best with voice and beak to resent
our approach. The young are covered with white down until nearly
half grown.

A few of these birds were also seen about the Tres Marias, and are
said to breed on San Juanito, but were not common.

Ardea herodias Linn. Great Blue Heron.

A few solitary individuals were seen at various times during IVIay
along the beaches, and it is probable that a few pairs may breed on the
islands. They were also noted by Colonel Grayson.

Ardea egretta Gmel. American Egret.

The American egret is another species noted by Colonel Grayson,
which we did not see. It must occur merely as a straggler from the
mainland.

Ardea candidissima Gmel. Snowy Heron.

Ivecorded by Colonel Grayson as a straggler, but not seen by us.
Nycticorax violaceus (Linn). Yellow-crowned Night Heron.

A dozen or more night herons were noted and a young bird was shot
on a rock close to Isabel Island, where it must have strayed from the
mainland. As Colonel Grayson found theju on the islands and took
13950_No. U 3

34 NORTH AMERICAN FAUNA.

specimens iu immature plumage, it is very probable that they breed
there iu small numbers.

Totanus flavipes (Gmel.). Yellow-legs.

A single specimen was shot by Prof. C. L. Herrick on Maria Madre
the middle of May.
Actitis macularia (Linu.). Spotted Sandpiper.

A few were seen along the shore on all of the islands, where they
probably breed.

Several small flocks of another sandpiper were seen along shore on
the islands; but although considerable energy was expended iu their
pursuit we failed to secure a specimen.

^gialitis semipalmata Bonap. Semipalmated Plover.

Colonel Grayson took a single specimen of this plover. It was not
seen by us and must occur only as a straggler.

Haematopiis palliatus Teiumiuck. American Oyster-catcher.

Rather common on the shores of the Tres Marias and Isabel, as well
as along the coast of the mainland near San Bias. A series of teu
oyster- catchea:s were secured and have been compared with four speci-
mens of Hamatopus galapagensis, one of H. frazari, and a number of
typical JET. palliatus (from the Atlantic coast of the United States and
the West Indies).

All of the birds from the Tres Marias, Isabel Island, and the adjacent
mainland were found to be surprisingly close to typical imlliatus. As
the Tres Marias are not far south of Lower California, the birds from
the islands might naturally be expected to be nearly typical repre-
sentatives of //. frazari. In reality about the only sign of gradation
toward the latter form is the mixed black and white across the lower
border of the black neck area. Some specimens from a single small
flock on Maria Cleofa had the line of demarkation between the black and
white areas on the breast as sharply defined as iu palliatus, while others
had the mixed black and white areas, as in frazari and (jalapagensis.
Some of the birds have a white spot on the under eyelid, which is
absent iu others, but otherwise the color is the same as in tjincal pal-
liatus. Birds from the Tres Marias and the mainland coast to the
south have an average shorter bill and tarsus than tm^ palliatus, and
in this character approximate /ra2;ari and galapagensis. Mr. Eidgway
has already called attention to the close general similarity existing
between the two latter species. The series from the Tres Marias and
adjacent coast agree with specimens iu the National Museum from
various points along the Pacific coast of Mexico, Central, and South
America in being very close to typical palliatus, thus showing pretty
conclusively that this is the resident bird along the coast and adjacent
islands south of Lower California.

So far as can be judged from specimens at hand, B. galapagensis is
distinct from frazari, although the birds resemble one other more closely

BIRDS OF THF: TRES MARIAS ISLANDS.

35

than tliey do representatives of palliatus from the adjacent mainland.
A series of specimens from the southern end of Lower California will
probably show intergradation between palliatus ^ud. J razari. The fol-
lowing measurements show the comparative sizes of birds from various
localities :

Measurements of Hcematopus palliatus, H. frazari, and H. galapaciensls.

Name.

Locality.

Sex.

Num-
ber of
speci-
mens.

Wing.

TaU.

Cul-
men.

•
Tarsus.

Hsematopua palliatus..

Atlantic coaat, United
States and West Indies.

?

3

260

102.3

90

63

Haematopus palliatus. .

'i'res Marias and Isabel
Islands.

d

6

256

100.1

75.8

58.1

Hasmatopus palliatus..

Trea Marias and adjacent
coast.

?

5

262.2

104

83.8

58.4

Hnematopus palliatus..
Hiumatopus frazari

?
?

o

263

97.5

84

61

Coast of Lower California . .

4

250.2

116.8

74.4

56.9

Haematopus galajia-

gensis.
Ha?matopu8 g a 1 a p a-

gousis.

2

253

98

83 5

54 5

3

246 6

101 6

82

57

Columba flavirostris madreiisis N'elaon. Ties Marias Pigeon.

Columha flai-irostris Grayson, Proc. Boston Soc. Nat. Hist., XIV, p. 274, 1871;

Lawr., Mem. Boston Soc. Nat. Hist., II, p. 304, 1874.
Columha flavirostris madrensis Nelson, Proc. Biol. Soc. Washington, XII, p. 6, 1898.

These handsome birds were rather common on Maria Madre and
Maria Magdalena, ranging to the summits of the islands, and they
probably live also on Maria Cleofa. On Maria Madre they were most
numerous along the wooded sides of a canyon some distance back from
the coast, where they usually perched among the higher branches of
the trees or were seen tiying about by twos and threes. Early in the
morning a few could be found among the smaller trees on the bases of
the foothills near the settlement, but later in the day they retired farther
inland to the more heavily wooded slopes. On Maria Magdiilena they
were numerous in some trees near a group of deserted houses and in
old clearings a short distance back from the shore. They came to these
trees to feed ui)on the ripening fruit, but were rather shy. When one
becomes startlfed and takes wing it makes a loud flapping noise that
alarms its companions, and then all dash swiftly away. They were less
conliding than most of the birds on the islands, but were not so shy as
their representatives on the mainland. Wild figs and the small fruit
of a tree, probably a species of Psidium^ or wild guava, were favorite
articles of food. Their loud cooing note is uttered at short intervals
and is one of tne characteristic sounds in the forests they frequent.
They are essentially arboreal in habits and are rarely seen near the
ground.

3G NOHTH AMERICAN FAUNA.

Zenaidura niacroura (Linn). Mourning Dove.

A single mouriiiiig dove was taken on May 5, ou Maria Madre, and a
few others w^ere seen on the ishmd during the first half of the month.
They were found for a short time about an old field near the shore, and,
like several other species, were probably stray migrants.

Leptotila fulviventris brachyptera (Salvadori). Wbite-fronted Dove.

There is a typical specimen of this bird in the National Museum col-
lection, which was taken on the Tres Marias by Colonel Grayson. It
was undoubtedly a straggler from the mainland, as it shows no approach
toward the characters distinguishing the resident insular species.

Ijeptotila capitalis Nelson. Tres Marias Dove.

Leptoptila alhifrons Grayson, Proc. Boston Soc. Nat. Hist., XIV, \i. 274, 1871 (part) ;

Lawr., Mem. Boston Soc. Nat. Hist., II, p. 305, 1874 (part).
Leptotila capiialis Nelson, Proc. Biol. Soc. Washington, XII, p. 6, 1898.

Very common on Maria Madre and Maria Magdalena, and prob-
ably occurs also on Maria Cleofa. They run about on the ground
under the shade of the forest with motions like those of a quail. Dur-
ing the morning and evening hours quiet trails leading through the
forest are their favorite resorts. When walking along these trails one
sees them for a moment, sometimes running and sometimes on the wing
close to the ground, as they disappear around the next bend. ]f one
is walking slowly the birds will frequently keep ahead for some distance,
but if pressed they either run or fly to one side into the sheltering
woods. They have a loud cooing note, which is heard at short intervals
wherever the birds are common. During the hot hours of the day they
retire to the shadiest recesses of the forest and usually perch in some
thick-topped tree. While resting in these retreats they have the pretty
custom of uttering mellow call notes, as if in response to one another.
During the breeding season they are seen in pairs, keepiugclose together,
but at other times are solitary. When forced to take wing, they do so
with a loud whirring sound and dart away through the intricate mazes
of the dense forest with wonderful (juickness. Their agility in flying
at full speed among the network of trunks and branches is extraordinary
and equalled by few birds If they take wing without being alarmed,
their flight is almost noiseless.

Melopelia leucopteia (Linn). White-winged Dove.

White-winged doves were rather common residents'on both Maria
Madre and i\Iaria Magdalena, and a few were seen on Maria Cleofa.

Colonel Grayson does not mention having seen this bird during any
of his visits to the islands. It seems quite improbable that so con-
spicuous a si)ecies should have been present and overlooked, and I
am inclined to believe that it has become a resident of the islands since
his visits. It is now a conspicuous and widely spread species and one
of the two resident land birds found by us that are not in Grayson's
list. In habits and appearance the white- winged doves of the islands
are identical with those on the mainland, where the species is very

BIRDS OF THE TRES MARIAS ISLANDS. 37

iiumeioiis. Two specimens were taken on Maria Madre, May 7, and
a single suecimen was shot on Isabel Island, April 22; tlie latter was
undoubtedly a straggler from the mainland, since Isabel is a waterless
island.

Columbigallina passerina pallescens (Baird). Mexican Ground Dove.

These pretty little doves were common on Maria Madre and Maria
Magdalena, but were most numerous about old fields and in the settle-
ment on the former island. The series of specimens taken on Maria
Madre appears to be identical with the birds of the adjacent mainland;
seven males from the islands average as follows: Wing, 87.4; tail, 61.8;
culmen, 11.6; tarsus, 16.6.
Cathartes aura (Linn.). Turkey Vulture.

Generally distributed, and very common about the settlement on
Maria Madre.

Buteo borealis fumosus Nelson. Tres Marias Red-tailed Hawk.

Buteo horcalis var. moniaua Grayson, Proe. Boston Soc. Nat. Hist., XIV, p. 268, 1871

(part).
Buteo ho7'eaUfi var. calurus Lawr., Mem. Boston Soc. Nat. Hist., II, p. 301, 1874

(part).
Btdeo horealis fitmosiis Nelson, Proc. Biol. Soc. Washiugton, XII, p. 7, 1898.

Colonel Grayson records this as a common species. We found a
few living along the canyons tliat score the slopes of Maria Madre.
They were very sparsely distributed and only some twelve or fifteen
individuals were noted; two or three were seen on Maria Magdalena
and none on Maria Cleofa. They were not at all shy, and whenever
found perched on a tree were readily approached within gunshot.
They feed mainly upon iguanas and rabbits, both of which are common
on the two larger islands. Nothing distinctive was noted about the
habits of these hawks. They are uniform in color, and differ more from
the mainland forms than does B. horealis socorroensis, although the
latter is from an island much fartlier out at sea.

The adult female of ,/MWo.s^^s• has some heavy shaft streaks of dark
brown on the chest, but these are not heavy enough to form a well-
defined dark area as is often the case in calurus.

Description of an im mature male ( ?) from Maria Madre : Upper surface
almost uniform blackish brown; tail mainly of same color, but crossed
by nine narrow, irregular lighter bands; a light area on tlie throat,
where the feathers have narrow blackish shaft lines and broad, dull
white borders; breast and sides of neck dnll, dark brown, with dull,
rusty edgings to feathers on latter area ; middle of breast paler ; feathers
on lower breast and flanks blackish brown, with irregular whitish spots;
abdomen and lower tail coverts dull brownish, jialer tlian flanks, with
pale butty barring; some feathers of tibia bufty or heavily barred with
butty, but mostly like those of lower breast.

38

NORTH AMERICAN FAUNA.

lielow are averages showing the relative size of the two islaud forms:

Measurements of Buteo borealis fumosus and Buteo b. socorroensis.

Name.

Locality.

Sex.

Num-
ber of
speci-
mens.

Wing.

Tail.

Cul-
men.

Depth
of bill.

Tarsus.

Buteo borealis fumo-
sus.

Maria Madre Island .
do

Ad. cT

Ad 9
Ad, ,:r
Ad. 9

3
1
2
1

X73 3
412
387.5
425

207.3
214
207.5
221

26
30
25
30

18.5
20
17.5
20

81.3
81

sus.
Buteo borealis socor-
roensis.

Socorro Island

do

80
86

roensis.

Falco peregrinus anatum (Bonap. ). Duck Hawk.

Falco peregrinus var. nigriceps Grayson, Proc. Boston Soc. Nat. Hist.. XIV, ]). 268,
1871.

A single duck hawk was taken by Colonel Grayson, who mentions
that it was shot while in close jjursnit of a sparrow hawk. We did
not see this species during our visit to the islands.

Paico sparverius Linn. Sparrow Hawk.

The sparrow hawk was recorded from the islands by Colonel Gray-
sou, but we did not see a single individual, and it probably occurs
merely as a straggler.

Falco columbarius Linn. Pigeon Hawk.

Colonel Grayson records this species as very common upon the
islands. Not a single individual was seen by us, and it probably
occurs only as a winter visitant or stray migrant.

Falco albigularis Daudin. Wbite-tbroated Falcon.

Hypotriorchis rufigularis Grayson, Proc. Boston Soc. Nat. Hist., XIV, ]>. 269. 1871;
Lawr., Mem. Boston Soc. Nat. Hist., II, p. 301, 1874.

Colonel Grayson took a specimen of this fine little falcon on the
islands. None were seen by us, and it must, no doubt, be classed as
one of the uumerous accidental visitants from the mainland.

Polyborus cheri^w^ay pallidus Nelson. Tres Marias Caracara.

Polyhorus audubonii Grayson, Proc. Boston Soc. Nat. Hist., XIV, p. 268, 1871,

■(part).
Polyborus tharus var. auduboni Lawr., Mem. Boston Soc. Nat. Hist., II, p. 308,

1874, (part).
Polyborus cheriway pallidus Nelson, Proc. Biol. Soc. Washington, XII, p, 8, 1898.

Very abundant about the settlement on Maria Madre and ratber
commonly distributed elsewhere over the island. They were also
rather common on the other islands, including Sau Juanito. The old
log roads and dry bottoms of the canyons on Maria JMadre were
favorite resorts. The birds were met in many unexpected places, and
were frequently seen perched in tree tops in the midst of the unbroken

BIRDS OF THE TRES MARIAS ISLANDS. 39

forest. Iguanas were excessively numerous, and furnislied tlie main
supply of food for the caracaras ; the birds were on the alert, however,
for anything in the form of carriojQ that turned up along shore or in
the forest.

Pandion haliaetus carolinensis (Gmel.) American Osprey.

Several osprey s were seen along the shores of all the islands, where
tliey ])robably nest. Colonel Grayson found a nest on a rock adjacent
to the northern shore of Maria Madre and another iu a large cactus.
Mr. Forrer obtained an immature bird during his visit to this island.

Strix pratincola Bonap. American Barn Owl.

Colonel Grayson states that he heard the well-known hissing scream
of this bird ;tt night on the Tres Marias. We were told of the pres-
ence of barn owls on the islands, but did not see them, and failed to
learn whether they occur as residents or stray migrants.

Speotyto cimicularia hypogaea (Bonap. ) Burrowing Owl.

Colonel Grayson says of the burrowing owl, " A few individuals of
this species inhabit the Marias Islands, perhaps wandered from the
m. inland." We neither saw nor could learn anything of them, and
those seen by Colonel Graysou were probably winter stragglers.

? Micropallas Sp.

The first evening after landing on Maria Madre, Mr. Goldman saw
and heard a little owl about the size of Glancidium phakvnoides on an
old log road in the forest. It was very near, and when he had moved
back far enough to shoot without destroying the bird it became too
indistinct to see and so escaped. Its notes were not like those of the
ferruginous owl. This was the only one seen or heard during our stay.
If a resident, it is very uncommon, for special but unsuccessful efforts
were made to find others.'

Amazona oratrix Ridgway. Double Yellow-headed Parrot.

Chrysotis levalUaniii Grayson, Proc. Boston Soc. Nat. Hist., XIV, p. 271, 1871;

Lawr., Mem. Boston Soc. Nat. Hist., II, p. 296, 1874.
Amazona oratrix Ridgway, Man. N. Am. Birds, p. 594, 1887.

The yellow-headed parrot is a common resident and always nests in
holes high up iu large trees iu the forest. Half-grown young were
found the middle of May. The birds were usually seen flying about
the forest in pairs, but congregated in flocks of from six to twenty or
more at their feeding places. They feed upon the fruits of various
trees, and during May the fleshy pods of Pithecolohium dulce formed
their staple article of diet. These low trees, from 15 to 35 feet high,
were growing around the houses of the settlement on Maria Madre and
were scattered thence along the coast, especially about the borders of

'During the visit to Maria Magdalena Island a larger owl was seen in the forest
along the bottom of a steep canyon, but was not sufficiently near to identify, and
escaped before it could be obtained. This was probably Ciccaba sqitamiilatn ( Bonap. ),
a species which is not rare on the mainland.

40 NORTH AMERICAN FAUNA.

old clearings and in scrubby second growth on the lower slopes. A
number of yellow-headed parrots came down every day to feed in the
trees, even among the houses, and did not i)ay the slightest attention
to passing people. As these birds readily learn to talk, they tire highly
prized as pets, and are sold to visitors, or sent to towns on the main-
laud; the birds taken while young being most highly prized on account
of their docility. The men search for their nests, and when one is
located tlie hunter strikes the base of the tree several sharp blows with
a stone or ax, and then places his ear against the trunk and listens.
He can tell whether the young are old enough to remove, by the strength
of the cries they utter in reponse to the blows on the tree. Being satis-
lied of the presence of his game, the hunter climbs the tree, and if
necessary cuts into the nest with his machete. Each brood contains
two young, which are carried to the ground inside the hunter's shirt.
By means of a noose on the end of a long cane, like a fishing rod, many
old parrots are captured while feeding. An old woman had twenty birds
which she had taken in this manner while they were feeding in the
top of a small rithecolobium tree by her door. The hunters search for
regular feeding places in the forest and wait under the trees for the
birds to come. When the birds arrive, the end of the rod is slowly and
cautiously pushed up through the branches, the noose slipped over the
bird's head and drawn about its neck with a quick jerk, after which
the victim is hauled down and thrust into a cage. A favorite resort
for the parrots on Maria Madre was a group of trees about half a mile
from the settlement. The birds were very unsuspicious, and one could
walk up within UO or 25 yards in full view and watch them without
their paying much attention. The parrots were constantly cliatteiing,
and the greatest good comradeship seemed to prevail. Mates kept
close together and showed their attachment by caressing and feeding
one anotlier at short intervals. The i>roficiency in speaking which some ■
of these birds attain is remarkable. The daughter of the customs
inspector <»u Maria Madre had one wliich afforded much amusement
by the variety of its remarks and their fre(]ueiitly absurd appropiiate-
uess. Colonel Grayson supposed these birds to be peculiar to the Tres
Marias, as he did not chance to tind them on the mainland. In reality,
they are widely distributed on both coasts of Mexico.

After comparing the series taken on the islands with specimens
from both coasts of the Mexican mainland certain slight differences
are noticeable, but are too poorly defined to be worthy of subspecific
recognition. The island birds are usually apple green on the dorsal
surface, and mainland specimens are more of an oil green; the ventral
surface has a more decided bluish wash ; there is also tendency to a
richer suifusion of orange and orange red on the yellow feathers about
the necks of old birds.

The following measurements show that the island birds are a little
larger than those of the mainland, with proportionately smaller bill and

BIRDS OF THE TRES MARIAS ISLANDS.

41

shorter tarsus. Averages of 9 specimens from the islands (both sexes) :
Wing, 233.3; tail, 133.6; oulmen, 34.3; tarsus, 24.7. Averages of 7
specimens from both coasts of the mainland (both sexes) : Wiug, 222.5;
tail, 120.1; culmen, 34.4; tarsus, 24.5.

Some old residents on Maria Madre reported that they had occa-
sionally seen stray individuals of another parrot which was a little
smaller than the yellow head, probably Amnzona finsehi, which is
abundant on the mainland.

Psittaculainsularis Ridgway. Tres Marias Lovebird.

Psiitacula eyanopyga Grayson, Proc. Boston Soc. Nat. Hist., XIV, p. 271, 1871;

Lawr., Mem. Boston Soc. Nat. Hist., II, p. 297, 1874 (part).
Psiitacula iiisiilaris Ridgway, Proc. U. S. Nat. Museum, X, 1887, p. .541 (Aug. 1888).
Paittacula ciianopygia Salvador!, Cat. Birds Brit. Mus., XX, p. 249, 1891 (part).

Lovebirds, or 'catarinas,' as they are called by the Mexicans, are
common on Maria^ Madre and Maria Magdalena, and probably occur on
Maria Cleofa, although none were seen on the latter island. They
were usually seen in flocks, from a few pairs up to 30 or 40 individuals,
and in May were feeding on small, sweet, wild figs, common on the lower
slopes. While feeding they keep up a constant chattering, which noti-
fies one of their presence. When flying over the top of the forest they
keep in compact flocks and move steadily forward with rapid wing
beats, suggesting a flock of cedar birds. They are very gentle, affec-
tionate little creatures and quickly become tame and greatly attached
to their owners.

Salvadori considers P-sittacula insiilaris a synonym of P. eyanopyga.
The series of eight adults from the islands and seven fiom the main-
land show that the island birds can be readily distinguished. The
two series show no signs of intergradation in color, and tlieir specific
distinctness is well defined, although the average measurements show
but slight difierences in size.

Description of Psitfacula insularis, ^ ad.,
Maria Madre, May, 1S97 :

Checks^ lores, forehead, and crown,
back to line between orbits, rich green,
decidedly clearer than lu eyanopyga, and
much more sharply contrasted with sur-
rounding colors.

Middle of crown green, shading
abruptly into dull bluish green, which
extends thence over neck, middleof back,
scapulars, tertials, and lesser wing
coverts.

Rump patch, axillars, and greater wing
coverts vivid cobalt blue.

Upper tail coverts Ijrighter green than
middle of back, but darker than in
cyauopyga.

Upper surface of tail dark green, darker
than in cyanopgga.

Description of Psittacula eyanopyga, $ ad.,
Tepie, Mexico, April, 1S97 :

Cheeks, lores, and forehead bright
green, more yellowish than m insularis,
and shading gradually into surrounding
colors.

Crown, from between orbits, neck, mid-
dle of back, sca]Miiars, tertials, .and
lesser wing coverts, dark green without
bluish.

Knmi> patch, axillars, and greater wing
coverts bright turcpioise blue.

Upper tail coverts clearer green than
back.

Ul)per surface of tail rich green.

42

NORTH AMERICAN FAUNA.

Secondaries and outer vanes of prima-
ries near base, dark blue; primaries with
outer vanes along distal half and at tips
edged with dark green; inner Avebs of
primaries brown.

Exposed under surface of primaries
and secondaries dull blue.

Lower parts, back to crissum, dingy
blue with a dull greenish wash; blue
brightest on abdomen, and contrasting
abruptly with green of crissum.

Crissum rich dark green.

Secondaries and outer vanes of inner
primaries dark blue ; rest of outer vaues
and tips of primaries dark green; inner
webs of primaries brown.

Exposed under surface of primaries
and secondaries dull bluish green.

Lower parts, back to crissum, dull
green, richest on abdomen and shading
insensibly into color of crissum.

Crissum brighter green.

Description of rsittacula insularis, § ad.,
Maria Madre, May, 1897.

Entire dorsal surface dingy green ;
brightest on sides of head, forehead,
rump, and upper tail coverts, with n dull
bluish shade on middle of back and
wings.

Lower surface dull green, back to
crissum ; the latter brighter green.

Description of rsittacula cyanopyga, 9 ad.,
from Tepic, Mexico, April, 1S97.

Dorsal surface dark green ; brightest
on forehead, rump, and upper tail coverts,
with an olive shade on middle of back
and wings.

Lower surface nearly uniform light
green, with a yellowish shade; crissum
nearly the same.

Average measurements of Fsittacula insulaiis and F. cyanopyga.

Name.

Locality.

Sex.

Num-

^^''".'^ Wing.
speci- ' »

mens.

Tiiil.

Oul-
men.

Tarsus.

ad. d
ad. ?
ad. cT

6 91.8
2 ni f;

45.8
46.5
45.0
41.5

13.9
13.5
12.8
13.1

12.9

do

13

Psittacula cyanopyga. .

3
4

90. C
89.2

13 1

do

ad. ?

12.7

Coccyzus minor (Gmel.) Mangrove Cuckoo.

This bird is rather common among' the inaugroves and other trees
bordering the salt lagoons near San Bias. A single specimen was
taken on May 8 near the shore of Maria Madre, but no others were
seen, and it probably occnrs there only as a straggler.

Trogon anibiguus goldmani Nelson. Gohlman's Trogon.

Trogon atnhiguns Grayson Proc. Boston Soc. Nat. Hist., XIX, p. 272, 1871 ; Lawr.,

Mem. Boston Soc. Nat. Hist., II, p. 290, 1874 (part).
Trogon ambiguits goldmani Nelson, Proc. Biol. Soc. Washington, XII, p. 8, 1898.

Goldman's trogon was common in the more heavily wooded parts of
Maria Madre and Maria Magdalena, and occurs no doubt on the less
heavily wooded Cleofa. On Maria Madre they were found from the coast
up nearly to the summit in suitable timber and especially along tlie sides
and bottoms of heavily wooded canyons. Their habits were similar
to those of the mainland bird. They sit quietly for a time on a branch

BIRDS OF THE TRES MARIAS ISLANDS. 43

and then tly, with an undulating motion, to another tree in the vicinity.
Their notes are limited to a short succession of unmusical sounds,
which are frequently heard. They were reported to nest in hollow
trees. Unlike most of the birds of these islands, the trogous were
nearly as shy as their representatives on the mainland, In life they
have light yellow bills and bright red eyelids.

Ceryle alcyon (Linn.). Belted Kingfisher.

Colonel Grayson records that during each of his visits one or two
belted kingfishers were observed sitting on rocks along the seashore.
None were seen by us.

Dryobates scalaris graysoni (Baird). Grayson's Woodpecker.

Picus scalaris var. graysoni Baird, Hist. N. Am. Birds, II, pp. 515, 517, 1874; Lawr.,
Mem. Boston Soc. Nat. Hist., II, p. 294, 1874.

This is the only woodpecker found on the Tres Marias. It is com-
mon on all of the islands and generally distributed, except in the most
heavily wooded areas. It seems to prefer second-growth thickets and
other places where shrubs and scrubby trees form low and rather thin
forests, and was usually seen hunting for food along the trunks of large
shrubs or small trees. It was a common practice for them to alight on
tree trunks near the ground and work slowly to the top, and after
remaining there quietly for a time to make a shoct flight to another
tree. Like its relative of the mainland, it is a very quiet bird, rarely
uttering any call notes and making little noise while searching for food.
From its habit of j)eering into crevices of the bark and doing only a
small amount of pecking it is evident that it finds most of its food on
or near the surface. Old nesting sites were seen in the trunks of both
living and dead trees and in the flower stems of large magueys. The
holes were usually between 5 and 10 feet from the ground. Colonel
Grayson found a nest about 12 feet from the ground in the green flower
stem of a large maguey (Agave) near the seashore in April.

A compari-son of a series of these island woodpeckers with other forms
shows that the dorsal surface, including the crest of the males, is most
like typical D. scalaris from the plains of Puebla. On the ventral
surface it may be distinguished from all the other races by its whiter
color and scantier and smaller black markings along the sides. These
markings are usually in the form of small rounded spots instead of
more or less elongated streaks, as in the other races, and the white
markings on the greater and lesser wing coverts are decidedly larger
and more conspicuous, i). s. graysoni averages a little smaller than
D. 6'. scalaris. The darker dorsal surface and unmarked bases of outer
tail feathers distinguish it from J), s. bairdii. It has a shorter, stouter
bill than D. s. lucasanus, with considerably more barring on the outer
tail feathers. From I), s. sinaloensis it is distinguished mainly by its
larger size, darker dorsal surface, and paler, less marked lower sur-

44

NORTH AMERICAN FAUNA.

face. The following averaj;es show the relative size of {/raysoni and
sinaloensis :

Measurements of Dryohates s. graysoni and Dryobates s. sinaloe^isis.

Kame.

Locality.

Sex.

Num-
ber of
speci-
mens.

Wing.

Tail.

Cnl-
nien.

Tarawa.

Dryobates scalaris

graysoni.
Dryobates scalaris

graysoni.
Dryobates scalaris

sinaloensis.

ad. cT
ad. 9
ad. cT

6
3
3

98.7
96.6
94.6

59.6

57

53.6

22.1
19.5
19.8

IS 6

do

Sinaloa and Tepic

17.5
17.1

An adult female in the National Museum collection from Mazatlau,
while having the normal bill and tarsus of sinaloensis, agrees with
birds from the Tres Marias in its long wings and tail. Its measure-
ments are as follows: Wing, 9G; tail, GO; culmen, 18.5; tarsus, 16.5.

Nyctidromus albicollis insularis Nelson. Trcs Marias Parauquc.

Nyctidromus alhicollis Grayson, Proc. Boston See. Nat. Hist. XIV, p. 273, 1871;

Lawr., Mem. Boston Soc. Nat. Hist., II, p. 291, 1874.
Nyctidromus alhicollis insularis Nelson, Proc. Biol. Soc. Washington XII, p. 9, 1898.

On the mainland the parauques are rarely seen while the sun is
above the horizon, but when night falls they come out of the dense
thickets where they have passed the day and sit in dusty trails and
other open places. On Maria Madre they were among the commonest
birds frequenting old log roads through the forest and shady canyon
bottoms until late in the morning and coining out again at 3 or 1 o'clock
in the afternoon. Of late years these plnces have been so completely
given over to solitude that when a human being chances to stray into
tliem he is looked upon with little fear. The wood folk seem to con
sider him harmless and only a strange creature of their own kind.

Parauques were among the most confiding birds found in these quiet
retreats and permitted a close approach before taking wing and moving
away. In the early dusk they were frequently seen hawking for insects
among the low trees Several came about cainp at the north end of
Maria Madre just after sunset, and flew very swiftly back and forth
with the same erratic course and vigorous wing strokes that are so
characteristic of the night-hawk. In fact, I mistook one of these birds
for a night hawk until it was secured. Their notes remind one slightly
of the whip-jioor-will's, but are not so loud and far-reaching. The
regular call is made up of two and sometimes three syllables, besides
which they have various little clucking and purring notes.

Curiously enough the parauques of the Tres Marias bear a much
greater resemblance, in size and color, to y. alhicollis merrilli of the
Rio Grande Valley than to the ordinary birds of the adjacent mainland.

BIRDS OF THE TRES MAUIAS ISLANDS. 45

Chordeiles acutipennis texensis (Lawr.) Texas Nighthawk.

A single specimen was taken May 5 on Maria Madre, and several
others were seen during tlie first half of the month, after wliich time
they disappeared. These birds were probably stray migrants, for there
was nothing to indicate that they were residents.

Amazilia graysoni Lawr. Grayson's Humming Bird.

Amazilia (/rayaoni Lawr., Ann. Lye. Nat. Hist., N. Y., VIII, p. 404, 1867.
Pyrrhophcrna (/rayaoni Grayson, Proc. Boston Soc. Nat. Hist., XIV, p. 283, 1871;
Lawr., Mem. Boston Soc. Nat. Hist., II, p. 292, 1874.

Very common on the islands. They were seen almost everywhere
darting about the less luxuriant parts of the forest hunting for tlowers,
and now and then stopping on a twig in some low tree top to rest or
arrange their plumage. They were among the numerous feathered
visitors to the little flower garden at the custom-house on Maria Madre
wliere they were very confiding, and would carry on their search for food
among the flowers quite iiidifterent to one's presence. These hum-
ming birds are very pugnacious, as the following incident recorded by
Colonel Grayson well illustrates : " Sometimes combats between them
become of a desperate nature. One day while watching a number of
them in active motion around some tobacco flowers (of which they seem
to be very fond) two fine males, after darting at each other for some
time, at length came to a deathly struggle, high above my head; they
finally clinched each other, each having one of the mandibles of the
other in his mouth, at the same time scratching with their little claws,
and using their wings with the greatest force, and in this situation,
whirling round and round, they fell to the ground near my feet. During
this terrible conflict, in which liassioii and desperation were exhibited,
I observed them for a few seconds and then gently i)laced my hat over
both. Even after they were thus captured, and I held one iu each
hand, they evidenced a desire to continue the war.''

The same author records having seen these hummers dart upon and
capture little flies in the manner of a flycatcker, and found their crops
full of minute insects. This I can corroborate from my own observations.

U])on comparing a series of 8 si)ecimens of Amazilia graysoni with an
equal iiumber of A. cinnamomea the general style of coloration is seen
to be very similiar, yet the difterences between the two forms are so
constant it seems advisable to consider them specifically distinct. The
dorsal surface of A. graysoni is a more dingy green with less coppery
iridescence than in cinnamomea; the bronze tips of the tail feathers
are less uniform, and the extreme points sometimes terminate witli a
little cinnamon spot; the cinnamon of the lower surface is considerably
darker. There is also a well marked and constant difference in size —
graysoni being the larger, as shown by the following averages:

46

NORTH AMERICAN FAUNA

Measurements of AmazUia r/rayfiuni and J. cinnamomea.

Name.

Locality .

Sex.

Num-
ber of
speci-
mens.

Wing.

Tail.

Cul-
raen.

ad. d
ad. ?

ad. (f
ad. 9

4
4
7
1

68.8
65.9
57.9
53

42.6
43.4
36.6
32.5

25

do

25.4
22

do

23

laclie lavyrencei Ridgway. Lawrence's Humming Bird.

Circe latirostris Grayson, Proc. Boston Soc. Nat. Hist., XIV, p. 282, 1871.
lache lawrencei (Berlepsch, Ms.) Ridgway, Man. N. Am. Birds, p. 320, 1887.

Like the preceding species this is a commou and generally distrib-
uted bird, and was found on all the islands. Like Grayson's hum-
miug birds, they were common about tlie flower garden at the custom
house on Maria Madre. Colonel Grayson found its nest on Maria
Madre and describes it as follows: "The elegant little structure I
found attached to a slender twig, and shaded with its leaves, about
5 feet from the ground. Tlie situation was fronting the sea, but a few
paces from the water's edge, where tbe first beams of the morning sun
dissolved the dews. Its form is cup shaped, and composed of the down
of the silk cotton tree {Eriodendron) intermingled with the down of
other plants and spider webs, the whole exterior neatly studded with
diminutive lichens; it contained two newly hatched young, but little
larger than Hies."

In general appearance J«c7ie lawrencei cioselyresemhles I. latirostris.^
The dorsal surfaces of the males are nearly the same color, but the
upper tail coverts of lairrencei are grayish instead of green, as in lati-
rostris-, their lower surface is a darker, duller green 5 the blue-throat
j)atch is nearly obsolete and replaced by an extension of the green of
the neck; the under-tail coverts are darker brown.

The females of lawrencei differ mainly from those of latirostris in the
clearer, brighter green of the dorsal surface and darker under-tail
coverts. As the difierences between the two birds seem to be fairly
constant," although not very striking, it is perhaps advisable to treat
them as species until more material jiroves the contrary.

The type of I. latirostris formed part of the Bullock collection and
probably came from the southern end of the table-land near the Valley
of Mexico. The following measurements show the differences in size:

' lache latirostris undoubtedly occurs on the islands as a straggler. About midway
on our return to the coast a hummer passed close to the side of the boat, coming
from the direction of San Bias and heading in a direct line for the islands. As this
wanderer passed 1 had a close view and identified it as /. latirostris.

BIRDS OF THE TRES MARIAS ISLANDS.

47

Measurements of laehe laivrencei and I. latirosiris.

Name.

Locality.

Sex.

Num-
ber of
speci-
mens.

Wing.

Tail.

Cul-
men.

ad. cf
ad. 9
ad. cf
ad. $

3
4
5
1

52.3
51.2

54.2
52

32.6
29.2
33
33

18.8

do

20 2

Southern table-lands, Mexico

do

21.9

23

Platypsaris aglaiae insularis (Ridg.) Graysou's Becard.

Hadrosiomus aglaiw v:ir. affinis Grayson, rroe. Boston Soc. Nat. Hist., XIV, p. 279,

1871; Lawr., Mem. Boston Soc. Nat. Hist., II, p. 289, 1874.
Platypsaris insularis Ridgway, Man. N. Am. Birds, p. 325, 1887.

The rose-tbroated becard was iiot commou and only three specimens
were taken, all on Maria Madre. They were found in the heavier
forest ou the slopes well back from the coast, and nothing unusual was
noted in regard to their habits. They probably occur on Maria Mag-
dalena and perhaps on Maria Cleofa. A considerable series of sj)eci-
mens from various parts of Mexico shows that a single species of rose-
breasted becard ranges over a large part of Mexico and has developed
four geographical subspecies. The ranges of these four forms may be
defined as follows :

Platypsaris aglaiw (Lafr.). Eastern Mexico from northern Tamau-
lipas south along basal slopes of the Cordillera of Vera Cruz and
Tabasco to arid parts of Yucatan. (Tyj)e from vicinity of Jalapa, Vera
Cruz.)

Platypsaris aglaice sumichrasti I^^elson. Humid lowlands of Vera
Cruz, and thence southward in similar country nearly or quite to Guate-
mala. (Type from Otatitlan, Vera Cruz.)

Platypsaris aglaice albiventris (Lawr.). West coast of Mexico from
the Isthmus of Tehauntepec to southern Arizona, ranging along river
valleys into the interior of western Mexico. (Type Irom Plains of
Colima.)

Platypsaris aglaiw insularis (Kidg.). Tres Marias Islands. (Type
from Maria Madre Island.)

Typical specimens of insularis are much darker than typical examples
of albiventris. Specimens from the coast lowlands about San Bias are
intermediate in color. The island birds, however, may usually be dis-
tinguished by their smaller bills. Back from the coast of Tei^ic,
esijecially in the arid river canyons at Bolaiios and near Guadalajara,
only typical specimens of albiventris were found. On the eastern side
of Mexico these two forms are paralleled by the pale bird of the foot
hills and adjacent interior {aglaiw) and the darker one of the coast
lowlands [sumichrasti). The color of extreme specimens of albiventris
is very different from that of aglaice and insularis, but among the series
from western Mexico, where albiventris has its home, are various inter-
mediate stages, some sijecimeus approaching very closely to both the

48

NORTH AMERICAN FAUNA.

forms just named. . Some specimens of insularis are nmcli nearer
typical ar/laiw in color than albiventri,s. Females of msularis are more
distinct from those of aglcvkc than the males, owing- to their generally
grayer backs, but even this is not a constant character. The only
(jharacter of insularis that is fairly constant is the smaller bill; a curi-
ous development, since there is a general tendency to an increase iu
size of bill among Tres Marias birds.

Measurement? of Phitypsaris aglaia: and its races in Mexico.

Tyrannus melancholicus couchi (Baird.) Couch's Kingbird.

On Maria Madre Island ten or a dozen of these birds were seen and
appeared to be resident. Two or three were noted on Maria Magdalena,
and others on Maria Cleofa, where they were most numerous. Tliey
were always found near the seashore. The sj)ecimens obtained seem
to be identical with those from the adjacent mainland. As a rule birds
from northeastern Mexico, the type locality, are lighter than those from
western Mexico, but this difference is not constant.

Myiarchus niexicanus magister Kidgway. Arizona Crested Flycatcher.

Although resident on all the islands, birds from the Tres Marias are
almost identical with those from the mainland, and nothing distinctive
was noted in their habits. They frequent the thinner parts of the
scrubby forests which cover most of the slopes, but were most numer-
ous within a mile or two of the sea. They kept among the low trees,
usually ])erching on tops of bushes or on branches within 10 or 15 feet
of the ground, where they watched for passing insects.

Myiarchus la-wrencei olivascens Ridgway. Olivaceous Flycatcher.

This was much more abundant than the preceding species, and one
of the commonest and most generally distributed resident birds on the
islands. Like the preceding, it was most numerous in the scrubby

1 Num-

Name.

Locality.

Sex.

ber ot
speci-
mens.

Wing.

Tail.

Cul-
men.

Tarsus.

Platypsaris aglaise. . .
Platypsaris aglaise
sumichrasti.

ad. cf

3

94.3

72.3

16

21.8

ad. cT

3

80.6

68.3

16.3

22

Platypsaiis aglai»

Plains of Colinia and Bolafios,

ad. cf

4

90.5

67.5

15.8

21.6

albiventris.

Jalisco.

Platypsaris aglai;«

Maria Madre Island

ad. cT

3

87

66.8

14.3

22.1

insularis.

Platypsaris aglaise,
insularis.

do

ad. ?

3

88

66.6

15

22.3

Intermediates be-

Coast near San IJlaa, Tepic

ad. rT

3

89.8

64.3

15.3

22.3

tween P. a. albi-

.

ventris and P. a.

insularis.

BIRDS OF THE TRES MARIAS ISLANDS.

49

forest within a mile or two of the seashore and kept among- the branches
of trees and bushes within 10 or 15 feet of the ground. Its habits
were the same as on the mainland.

A careful comparison of series from the islands and the mainland
shows but little difference. The island birds are slightly grayer on the
back, the bills average a little longer and wider, and the tarsus is longer,
but these differences are too slight to warrant subspecific recognition.
Unfortunately the type of this subspecies is based on a winter specimen
from Santa Eflgenia, Oaxaca, near the border of Chiapas. This leaves
the summer range of typical birds in doubt.

Below are averages of specimens from the islands and the mainland:

Measurements of Myiarchus lawrencei olivasvens.

Name.

Locality.

Sex.

Num-
ber of
speci-
mens.

Wing.

Tail.

Cul-
men.

Tarsus.

Myiarchus lawrencei

olivascens.
Myiarchus lawrencei

olivascens.

Trea Marias Islands

Northwest Mexico, southern
Arizona.

ad. ?
ad. $

7
7

76.3
7G.8

74.8
74.7

16.8
16.1

19.7
18.8

Contopus richardsonii (Swains). Western Wood Pewee.

During the first ten days of May these birds were not uncommon in
some of the denser growths of small trees along the lower slopes of Maria
Madre. Kone were seen on the other islands and they all disapjieared
a little later, showing that they were merely stray migrants. The
single specimen taken is identical with others from the western United
States.

Empidonax difdcilis Baird. Western Flycatcher.

These birds were very sparingly distributed on the Tres Marias.
They were seen on Maria Madre soon after our arrival, and on May 27
cue was taken on Maria Magdalena. They were usually found in dense
thickets and along shady canyons. Colonel Grayson records them
as common, but they undoubtedly occur only as winter visitants and
migrants.

Ornithion imberbe (Sclater). Beardless Flycatcher.

Two specimens were taken and a few others seen early in May on
Maria Madre; none were seen later, and they probably occur merely as
stragglers during migration. They move about like small vireos in the
tops of low trees and in large bushes, searching the outer twigs for
insects and flying out every now and then to capture one on the wing.

A series of specimens from various localities on both coasts and the
interior of Mexico, the Tres Marias Islands, and southern Arizona fails
to show any tangible characters to distinguish O. imherbe ridgicayi
from typical 0. imherbe.
13950— Ko. U 4

50 NORTH AMERICAN FAUNA.

Myiopagis placens (Sclater). Golden Crowned Flycatclier.

In the National Museum collection is a typical specimen of M.
2)laeens, taken by Colonel (ji ayson in January, 1865, on the Tres Marias
Islands, which agrees perfectly in size and color with the large dark
birds from the mountains of Jalisco, on the adjacent mainland. Eight
specimens of both sexes from various localities in Mexico and Central
America average as follows: Wing, 68.2; tail, 65.8; culmen, 11.2;
tarsus, 19.1.

Myiopagis placens minimus Nelson. Little Golden Crowned Flycatcher.

Elainea i)lace\)s Lawr., Proc. Boston Soc. Nat. Hist., XIV, p. 279, 1871; Mem.

Boston Soc. Nat. Hist., II, p. 286, 1874.
Myiopaais placens minimus Nelson, Proe. Biol. Soc. Washington, XII, p. 9, 1898.

Early in May a few of these birds were seen in the low scrubby
forest near the shore on Maria Madre, but by the middle of the mouth
they had retired to the heavily wooded canyons and slopes above 600
or 800 feet. They were common among the trees bordering the head of
a large canyon in the middle of the island, where a small spring formed
the center of attraction for many birds. Like their mainland relative,
they frequent the tree tops, where they may be seen running in and
out among the ends of the branches and fluttering about the tips of
twigs in the manner of some warblers. They frequently hop from twig
to twig, with their tails uptilted like gnatcatchers, but their dull garb
is usually suflicient to identify them. When high up in the tops of tall
trees, however, their form and habits are so warbler like that they can
not be distinguished from other small birds. A few individuals were
seen on Maria Magdalena, but none on Maria Cleofa.
Corvus mexicanus Gmelin. Mexican Crow.

Two residents on Maria Madre reported that at long intervals they
had seen stray crows on the island. My informants were familiar with
the bird on the mainland, where it is extremely numerous about San
Bias, and there is no reason to doubt the correctness of their identifi-
cation.

Cissolopha beecheyi (Vigors). Beechey's Jay.

A specimen of this bird in the National Museum collection, obtained
by Xantus, is labeled "Tres Marias Islands, October 15, 1859". This
record, however, is doubtful.

Icterus graysoni Cassin. Grayson's Oriole.

Icterus graysoni Cassin, Proc. Acad. Nat. Sci. Phila., p. 48, 18fi7; Mem. Boston Soc.
Nat. Hist., II, p. 280, 1874.
These beautiful birds are very common on all of the islands. Although
more numerous about the settlement on Maria Madre than elsewhere,
they were common in the thin, low forest all about the lower parts of
the islands and were very unsuspicious During my excursions through
the woods they came again and again and alighted on low branches of
shrubs or trees beside the old log roads and peered at me with inno-

I

BIRDS OF THE TRES MARIAS ISLANDS.

51

ceutly inqiiiriDg eyes as if wondering at the strange creature newly
arrived in their haunts, but evidently quite unconscious of any feeling
that the newcomer might be dangerous. Such confidence made it very
trying work to collect many of these birds.

They came familiarly about the houses and yards at the settlement
on Maria Madre. A number of them made several visits each day to
the verandas and shrubbery about the custom-house, and added greatly
to the attractive surroundings by their bright colors and frank uncon-
cern. They searched for insects among the shrubs and small trees in
the ijatio or court, came to the veranda railing, down upon the floor,
and along the walls, where i>lump spiders furnished many choice morsels.
Several bags of corn piled against the wall on one side of the veranda
were infested with weevils, which could be found creeping about on the
outside of the bags. A pair of orioles was in the habit of regularly
visiting the veranda and soon discovered these insects. They walked
all over the bags, sometimes upside down or on one side like a nuthatch,
and inied into every spot likely to contain a little beetle. They were
frequently seen also clinging to the stems of the giant cactus ( Cereus)
and feeding on the juicy fruit.

As Colonel Grayson has recorded, the nests of these orioles are about
a foot in length and of the usual purse shape. They are made of fibers
of grass or maguey plants, lined with silk cotton and swung near the
end of some slender branch overhanging a clear space, usually from 18
to 35 feet above the ground.

Grayson's oriole is evidently an offshoot from the wide ranging
Icterus pustulatus of the adjacent coast, but has become siitticiently
distinct to rank as a species. Like so many of the island birds, it is
larger than its mainland relative. The yellow is much lighter than
on /. pustulatus and lacks most of the intense orange that is so
conspicuous on many of the latter birds. Some adult males of graysoni
have the back entirely bright yellow, while the backs of others are
marked with a few narrow black shaft streaks. The females of gray-
soni are more greenish-yellow and have but faint traces of the orange
shade present in ty\)\cai 2)ustulatus.

The following averages show the relative dimensions of the two
species :

Measurements of Icterus (jraysoni and I. pustulatus.

Name.

Locality.

Sex.

Num-
ber of
speci-
mens.

"Wing.

Tail.

Cul-
men.

Tarsus.

Icterus graysoni

ad. d
ad. ?
ad. d
ad. ?

4

4
4
4

104
96.7

100.2
91.7

89.7
84.2
91
81.5

25.4
25.1
21
20.2

26 7

do

26.2

Icterus pustulatus

25.6

do

24 5

52 NORTH AMERICAN FAUNA.

Quiscalus macrourus Swiiinson. Gre.it-t.iiled Grackle.

Two of these grackles were shot the latter part of May on a level
bit of ground bordering the shore in front of the settlement on Maria
Madre. They were the only ones seen and were undoubtedly strag-
glers from the mainland where tliey are abundant and resident near
San Bias.

Astragalinus psaltria mexicanus (Swainsoii). Mexicau Goldfinch.

Rather common and apparently resident, but nothing distinctive was
observed in their habits. On Maria Madre they were usually found on
the lower slopes and were most numerous about the settlement. Ten
specimens fail to show any characters distinguishing the island birds
from those of the mainland.

Cardinalis cardiualis niariae Nelson. Tres Marias Cardinal.

Cardinalis rirginianus Lawr., Proc. Boston Soc. Nat. Hist., XIV, p. 281, 1871.
Cardinalis vir<jinianus var. ignens Lawr., Mem. Boston Soc. Nat. Hist., II, p. 275,

1874.
Cardinalis cardinalis maria; Nelson, Proc. Biol. Soc. Washington, XII, p. 10, 1898.

Cardinals were very common on Maria Madre ami not uncommon
on the rest of the group. No one ever molests them, and they were
especially abundant about the settlement, where they came into the
yards and around the houses in the most familiar way. Several pairs
could be found at any time during a short walk in the scrubby thickets
along the lower slopes of the island. While we were hunting in the
low woods it was a common occurrence for them to come very near, and
after looking at the intruders with mild curiosity for a short time, to
move off through the bushes in quiet pursuit of their usual occupations.
At other times, while engaged in search of food among the fallen leaves
they would scarcely notice one as he walked slowly by within three or
four paces.

Piranga ludoviciana (Wilson). Louisiana Tanager.

During the first half of May these tanagers were not uncommon near
the settlement on Maria Madre, but were not seen on the other islands.
Those shot the first of the month were in fair condition, and, several
pairs being seen, it was at first considered a resident species. Later,
when others were secured, it was noted that they were more find
more emaciated, until those killed about the middle of the month were
so excessively thin, it was surprising that they had continued to live.
About this time the last ones disajjpeared, no doubt dying from star-
vation. From these observations it appeared that the birds must
have strayed to the island during migration, about the last of April
or first of May, and were unable to find a proper food supply. At the
same time they feared to start over the sea for an invisible shore and so
perished. Another member of the genus, Firanga Mdentata JIammea, is
resident in large numbers on the islands and found an abundant food
supply, as was shown by their being among the fattest birds collected
during the time that F. ludoviciana was dying of starvation.

BIRDS OF THE TRES MARIAS ISLANDS. 53

Piranga bidentata flammea (Ri(lji;way). Tres Marias Taoager.

Pyrantja hidentala Lawr., Proc. Boston Soc. Nat. Hist., XIV, p. 281, 1871; Mem.

Bostou Soc. Nat. Hist., II, p. 274 (part), 1874.
Firamja flammea Ridgway, Man. N. Aqi. Birds, p. 457, 1887.

Several species of birds were very much at liome about the settle-
ment on Maria Madre, and among these the brilliant Tres Marias taua-
ger was one of the most numerous. Like Grayson's oriole, they came
daily to the veranda railing and investigated the shrubs and small
trees in the court and iiower garden at the custom-house. These birds
were common and generally distributed in the scrubby forest on the
lower parts of Maria Madre and Maria Magdalena, and probably occur
on Maria Cleofa, although none were seen there. Their habits were
very much like those of Piranga bidentata on the mainland. On the
island, however, these tanagers were most numerous within a few hun-
dred feet of sea level, while their relatives of tlie mainland inhab-
ited oak forests at an altitude of 2,000 or 3,000 feet. They have a
short warbling song, which is similar to, but less musical than, that of
the mainland bird. They were seen hunting for food in the small tree
tops of the scantier forest growths rather tlian in the more densely
wooded areas and were very fat.

P. hidentata was described by Swainson from a specimen in the
Bullock collection, taken at Temascaltepec, southwest of tlie Valley of
Mexico, on the Pacific slope of the mountains. It was described as
having the 'head, neck, and under parts golden'. This style of colora-
tion is shown in specimens from various localities in Jalisco, Sinaloa,
and the Tres Marias Islands. Judging from specimens in the National
Museum and from the results of recent work, tanagers of this descrij)-
tion are only found north of the Isthmus of Tehuantepec, on the arid
western slope of Mexico, and are not common. The Tres Marias
tanager is closely related to typical P. hidentata, and the males are so
closely alike in color that it requires careful scrutiny to find distin-
guishing characters. In P. fiamniea the white tips of the greater
and lesser wing coverts are larger and clearer white than in P.
hidentata, thus rendering the two wing bands more consj)icuous. The
white spots on the outer rectrices are smaller and confined to the inner
webs, except at the extreme t\^; in P. hidentata these marks occupy
most of the terminal third of the feathers. In general color of the
body the two forms are indii^stinguishable. The bill of P. flammea
averages longer and is decidedly more swollen, especially toward the
tip; this difference is one of the most important characters of the
island form. The female of P. flammea can be distinguished only by
the larger bill and the restriction of the white spot on the outer pair
of tail feathers.

54

NORTH AMERICAN FAUNA.

The following measurements give the averages of the two forms:
Measurements of Piranga bidentata and Piranga b. flammea.

Name.

Locality.

Sex.

Num-
ber of
speci-
mens.

Wing.

Tail.

Cul-
men.

Tarsus.

Piranga bidentata flammea.
Piranga bidentata liammea.

Piranga bidentata

Piranga bidentata

Maria Madre Island . .
do

Jalisco and Sinaloa . . .
do

ad. cT
ad. ?
ad. cf
ad. ?

6
4
3
2

98
95
98
96

81

78.7
79.3
79

18.1
18.4
17.3
17

23.7
22.5
21.1
21.5

Hirundo erythrogaster Bodd. Barn Swallow.

Soon after our arrival on Maria Madre a few swallows, supposed to
be this species, were seen by my assistant, but none were taken. They
were undoubtedly stray migrants, for none were seen afterwards.

Vireo flavoviridis forreri (Madarasz). Forrer's Vireo.

Vireo forreri Madarasz, Term^szetrajzi Fiizetek, IX, pt. I, p. 85, 1885.

Although Forrer's vireo is one of the most abundant and widely dis-
tributed species on the islands, yet it does not appear in Grayson's
list. It was very common in the small trees in the patio of the custom-
house and elsewhere about the settlement on Maria Madre. Like its
mainland relative, its habits are very similar to those of the red-eyed
vireo. Its favorite range was in the smaller growth of forest along the
lower slopes, from near the sea u^) to an altitude of 600 or 700 feet, but
some were seen up near the summits of Maria Madre and Maria
Magdalena. Kext to the Tres Marias warbler, Forrer's vireo was prob-
ably the most abundant bird on Maria Madre, and its restless habits
while tlutteiing and peering about in search of food among the small
tree tops added greatly to the animation of the forest.

Vireo forreri is evidently only a geographical race of Vireo flavovi-
ridis. It has the same color pattern, but the ashy crown is paler and
the dusky supraorbital stripe usually obsolescent; the latter is one of
the main characters upon which forreri was originally based, but is not
constant. Some specimens from the i.slands have this strii)e as strongly
marked as dull-colored individuals of flavoriridis proper, although none
have it so strongly marked as some of the latter. The two forms are
alike on the underparts, and the greater size of forreri is the most
constant and striking character.

Average measurements of 17 adult mates of Vireo flavoviridis forreri:
Wing, 84.3; tail, 59.3; culraen, 15.1; tai'sus, 20.1. Averages of Vireo
flavoviridis (from mainland of Mexico): Ad. $ (9 specimens), wing, 79.2;
tail, 55.1; culmeu, 11.3; tarsus, 18.7. Ad. 9 (3 specimens), wing, 7G.6;
tail, 50.6; culmeu, 14.1; tarsus, 18.5.

Vireo hypochryseus sordidus Nelson. Tres Marias Vireo.

Vireo hypochryseus Grayson, Proc. Boston Soc. Nat. Hist., XIV, p. 281, 1871 ; Lawr.,

Mem. Boston Soc. Nat. Hist., II, p. 272, 1874.
Vireo hypochryseus sordidus Nelson, Proc. Biol. Soc. Washington, XII, p. 10, 1898.

A few of these vireos were seen in the thin forest on the lower slopes
of Maria Madre, but were not cooinion. They were especially numer-

BIRDS OF THE TRES MARIAS ISLANDS. 55

ous among the trees and tall bushes about the few springs and little
streams near the summit. A few were also seen in similar places on
Maria Magdalena. Vireo f. forreri occupies the lower slopes, while
sordidus occurs mainly higher up, the ranges of the two birds being
complementary. The Tres Marias vireo is usually found at a medium
height among the foliage of thick-topped trees, rarely ascending to the
extreme top. It was also often seen in the dense, tall undergrowth
near water.

Compsothlypis insularis" (Lawr.). Tres Marias Parula.

Panda insularis Lawr., Auu. Lye. Nat. Hist., N. Y., X, p. 4, 1871; Grayson, Proc.
Boston Soc. Nat. Hist., XIV, pp. 278, 300, 1871; Lawr., Mem. Boston Soc. Nat.
Hist., II, p. 269, 1874.

These pretty little warblers were the most abundant of the land birds
on the Tres Marias. A few of them were also founo on Isabel Island,
and the only Compsothlypis taken on the mainland at San Bias belongs
to this species. They frequent the thin forest of the lower slopes on
the Tres Marias, and dozens of them were seen during every visit to the
woods, and they were seen in smaller numbers on the higher slopes.
Many also came familiarly into the small trees and shrubbery about the
houses at the settlement. They were always busily at work in pursuit
of insects among the branches, and searched the bark of small shrubs
near the ground as well as the branches at the tops of large trees.
They were rather common in the scrubby growtli of stunted trees on
Isabel, and were very abundant in the tree tops of the heavy forest on
the mainland between San Bias and Santiago. Their song is weak and
lisping and not at all musical.

There is little doubt that a good series of specimens will demonstrate
that Compsothlypis pitiayumi of northern South America is represented
in Central America and Mexico by a number of geographical races
rather than by the closely related species now recognized — C. inornata,
C. pulchra, C. nigrilora^ and G. insularis. Even the imperfect series at
hand shows signs of intergradation, but treating C. insularis as a
species for the i)resent, its differences from its nearest relative, C.
pulchra^ are set forth in the following notes. C. pulchra was the only
form found on the mainland back of the low coast plain, on the tropical
or subtroi)ical sloj)es of the mountains. This species was described
I'rom Chihuahua, and appears to be a resident of the lower slopes of
the Sierra Madre, ranging from Chihuahua to Tei)ic, while G. insularis
is characteristic of the hot lowlands on the coast near San Bias and the
outlying ishmds,

G. insiddris is larger than C. pulchra^ with a heavier shading of
brown along the Hanks; the yellow of the under parts is duller and
more generally suffused with dull orange brown; the white spots on
outer tail feathers are deciderlly larger, and the bluish of the dorsal
surface is grayer, in the small series examined, difference in size
seems to be the most constant character. Following are average meas-
urements of the two species :

56

NORTH AMERICAN FAUNA.

Measurements of Compsothlypis insalaris and ('.pulchra.

Name.

Locality.

Sex.

Nnm-

ber of
speci-
mens.

Wing.

Tail.

Cul-
men.

Tarsus.

Compsothlypis insularis ...

Maria Madre Island . .
do

ad. cf
ad. ?

6
4
3
1

CO

55.7
55.6
52

49.3
47.5
42.3

41

10.4
10

9.6

?

19.9,
19

Compsotlilyi)is pulchra

Compsotlilypis pulchra

Jalisco and Sinaloa . ..
.do

art. d
ad. ?

17
17

Dendroica aestiva rubiginosa (Pallas). Alaskan Yellow Warbler.

Several of these birds were taken and others seen about the settle-
ment ou Maria Madre. TLey were evidently stray migrants, and most
of them left before the end of May.

Dendroica aestiva niorconii Coale. Western Yellow Warbler.

Among the yellow warblers taken on Maria Madre during the first
half of May were two specimens referable to Dendroica astira viorcomi.
Like ruhiginoso^ they were stray migrants which had wandered out of
their course while en route to their more northern breeding grounds.
They were found about weed patches and shrubbery in the settlement.

Dendroica auduboni (Townsend). Audubon's Warbler.

Two of these birds were seen during the first half of May about the
settlement on Maria Madre, and May 30 a specimen was taken on
Maria Cleofa. Like the yellow warblers, they occur merely as stray
migrants and were seen only near the seashore.

Dendroica towrnsendii (Townsend). Townsend's Warbler.

Two or three of these warblers were seen at the settlement on Maria
Madre between the 8th and 20th of May. They kept about the weed
patches and yards for several days, and were stray migrants like the
preceding species.

Granatellus fraiicescae Baird. Tres Marias Chat- Warbler.

Gratiatelhis francescw Bairl, Rev. Am. Birds, p. 232, 1865; Grayson, Proc. Boston
Soc. Nat. Hist., XIV, p. 278, 1871 ; Lawr., Mem. Boston Soc. Nat. Hist., II, p. 270,

1874.

These beautiful birds were seen only on Maria Madre, but they prob-
ably occur also on Maria Magdalena, where the conditions are equally
favorable. They were far from common, and inhabited the forest on
the higher slo^ies, but two or three individuals, evidently wanderers,
were encountered in the scrubby forest near the shore. They were
usually seen on the ground searching for food among low underbrush
and weeds. In such places they ran about among the thick stems of
plants and matted undergrowth, springing up every now and then to
a twig or weed stalk a foot or two from the ground, and then perhaps
flitting along from stem to stem to another feeding place a few yards
away. When thus passing through the undergrowth, they are very
conspicuous and attractive objects, owing to their beautifully contrasted

BIRDS OP' THE TRES MARIAS ISLANDS.

57

black, white, aud rose-colored plumage. Their habit of carryiDg the
tail up-tilted and more or less widely si)read renders them still more
conspicuous. It is doubtful if they ascend into the tops of trees, as
they are even more terrestrial than their relatives the chats.

The color pattern of this species is much like that of G. venustiis, but
the black collar on the lower side of the ueck in the males is nearly
obsolete, being represented only by a few black feathers, the red or
rose colored area on the breast and chest is paler and more restricted,
the postocular white stripe larger and extending across the nape as an
indistinct nuchal band, the bluish of the dorsal surface grayer, aud the
white on the tail more extended. The females are browner above and
paler below. G. franceHCAV is larger than G. venustus, as shown by the
following averages :

Measuremevts of (TranateUus francescw and fr.renustus.

Name.

Locality.

Sex.

Num-
ber of
speci-
mens.

Wing.

Tail.

Cul-
men.

Tarsus.

Granatellus francescaj

Maria Madre Island . .
... do

ad. cf
ad. ?
ad. cf
ad. ?

5
2

1

65.8

63

61.5

58

76.5

74.5
66.5
67

12.2
12.2
12

12

21

21 5

Granatellus Yennstiis

Guerrero and Oaxaca .
do

19.7
20.5

Wilsonia pusilla pileolata (Pall.), Pileolated Warbler.

The only one seen was taken on Maria Cleofa May 30. Tt was in
some bushes by a little stream near the seashore and was evidently a
straggling migrant.

Minius polyglottos (Lian.). Mocking Bird.

A few mocking birds were seen on Maria Madre, where they are prob-
ably resident in small numbers. They were found only on the lower
slopes near the sea. The two specimens secured appear to be identical
with others from the adjacent mainland.

Thryothorus law^rencii (Ridgway). Maria Madre Wren.

Thryothorus felLc Grajson, Proc. Boston Soc. Nat. Hist. XIV, p. 278,1871 (part);

Lawr., Mem. Boston Soc. Nat. Hist. II, p. 268, 1874 (part).
Thrijothorua felix fi lawrencii Ridgway, Bnll. Nutt. Orn. Club, III, p. 10, Jan., 1878.

The song of this wren was one of the most constant and pleasing of
the woodland notes heard on Maria Madre. The bird was extremely
abundant everywhere in the undergrowth ranging from the shore up
to the higher slopes. Like its near relatives, it is a restless little crea-
ture, constantly climbing and i^eering about in the thickets. The male
stoi)s every now and then to utter his song and then continues insect
hunting. When in a musical mood he takes a position in some small
shrub, sometimes on its summit but oftener on a branch at one side,
aud there pours out his song again aud again at short intervals. Like
many other birds on these islands, the wren was vsry familiar and un-

58

NORTH AMERICAN FAUNA.

sus]]ricious, aud many came every day to the feuces aud shrubbery around
the bouses at the settlement.

A series of Thryothorus felix from the mainland, including one speci-
men from the region of the type locality, and a series of T. laivrencii
from Maria Madre, show sufficient differences to warrant giving specific
rank to lairrencii. The latter difters very constantly in several respects
from birds of the mainland, but has much the same color pattern. The
series I'rom San Bias is nearer lawrencii than is the specimen from near
the type locality of felix, but there appears to be no crossing of the
gap between the two.

The following measurements show the relative sizes of the two
species :

Measurements of Thryothorus lawrencii aud T. felix.

Name.

Locality.

Sex.

Num-
ber of
speci-
mens.

Wing.

Tail.

Cul-
men.

Tarsus.

Thryolhorna lawrencii

Maria Madre Island . . .
.... do

ad. cf
ad. ?
ad.cT

ad.?

3

7
2

3

60

57.1

57.5

54.3

55.6

54

55.5

50.6

17.2
16.8
16

U.6

22
21.4

Thryothorus felix

Santiago, Tepic to

Ometepec, Guerrero.

do

21.5
20.5

Thryothorus lawrencii magdalenae Nelson. Magdalena Wren.

Tliryothorus felix Grayson Proc. Boston Soc, Nat. Hist., XIV, p. 278, 1871 (part);

Lawr., Mem. Boston Soc. Nat. Hist., II, p. 268, 1874 (part).
Thryothorus lawrencii mat/dalena' Nelson, Proc. Biol. Soc. Washington, XII, p. 11,

1898.

The habits aud distribution of this wren on Maria Magdalena are
the same as those of T. lawrencii on Maria Madre. No one lives on
Maria Magdalena, and the wrens are even tamer than on Maria Madre.
Their confidence was shown very prettily by one encountered by 3Ir.
Goldman in the dark bottom of a narrow rocky canyon overhung with
heavy forest. He saw the little fellow busily searching for food among
the fallen leaves along the base of a low cliff, and as the bird seemed
very fearless he approached quietly but in full view, and succeeded
in closing his hand over the tiny creature, which had continued its
search without i)aying the slighest attention. The bird showed but
little fright, and its captor, after holding it a few moments, stooped
and gently opened his hand to let it escape. The wren hopped away a
few feet, arranged its plumage, aud then continued feeding with the
utmost unconcern. Mr. Goldman watched it for a few minutes aud
again approached slowly. As before the bird paid uo attention until
he was within a yard, but when another attempt was made to pick it
up, hopped away a few feet and again resumed its occupation. This
was repeated three or four times with the same result, until finally the
bird was left in its solitude.

BIRDS OF THE TRES MARIAS ISLANDS. 69

Melanotds caerulescens longirostris Nelson. Ties Mari.is Blue Mockingbird.

Melanotis ccerulescens Grayson Proc. Boston Soc. Nat. Hist., XIV, p. 275,1871;

Lawr., Mem. Boston Soc. Nat. Hist., II, p. 266, 1874 (part).
Melanotis ccerulescens longirostris Nelson, Proc. Biol. Soc. Washington, XII, p. 10
1898.

These fine songsters are very common on the Tres Marias. They keep
in the thickets and low trees and bushes like a catbird and were
especially numerous and familiar about the settlement on Maria Aladre.
In one yard, among a few fruit trees, a trough was kept full of water,
where scores of blue mockingbirds came daily to drink and would
almost allow themselves to be caught by liand. Their numbers and
general distribution make them among the most noticeable birds on the
islands, and they frequently follow one with much curiosity. Their song,
although rich and varied, was not so clear and musical as that of their
relatives on the mainland. The birds on Maria Madre show a marked
tendency to albinism, which usually appears in the form of grayish or
whitish bars on the wings and tail. In addition to the barring on the
primaries and secondaries, the alula is often similarly marked and some
sj)ecimens have lighter spots on the tips of the wing coverts, produc-
ing well-defined wing bands. The markings are usually symmetrical,
but vary in amount and intensity with the individual. In some they
are barely distinguishable and in others very conspicuous. More rarely
the albinism api^ears on other parts of the body, occasionally in asym-
metrical areas of pure white, but these spots also are sometimes regu-
lar. One specimen has the entire under surface white, except some
blue feathers along the flanks, and the rump is white mixed with blue.
This bird has a striking general resemblance to the Central American
Melanotis liypoleucns. At least 2 or 3 per cent of the birds on the
islands are albinistic, and the constant recurrence of the same light
barring on the wings and tail seems to indicate the possible evolution
of a form in which these markings will be constant.

Myadestes obscurus iusularis Stejueger. Tres Marias Solitaire.

Myiadesies obscurus Grayson, Proc. Boston Soc. Nat. Hist., XIV, p. 277, 1871;
Lawr., Mem. Boston Soc. Nat. Hist., II, p. 273, 1874.

Myadestes obscurus var. iusularis Stejneger, Proc. U. S. Nat. Mns., IV, pp. 371, 373,
1882.
This is a common bird in the heavy forest about the heads of can-
yons on Maria Madre and Maria Magdalena. They v:ere not found
anywhere in the scrubby growth of the lower slopes, and if they occur
there at all it must be only as stragglers. They are shy birds, remain-
ing silent when approached, but when undisturbed flitting through
the tree tops like wandering spirits of melody uttering their sweet
strains from the mysterious depths of the forest. Their song was
heard from the tops of tall trees where the birds sat amid the heavy
foliage, rarely coming down to lower levels except in the morning or
evening, or to drink at midday. Many were seen about a spring near
the top of Maria Madre where they came to drink at noon.

60

NORTH AMERICAN FAUNA.

Although Myadestcs ohscurns insnlaris is very closely related to occi-
dentalis, yet it may be distinguished by several slight but coustant
characters, such as the greater extension and paler shade of ashy from
the ne(;k over the forward part of the back. The lower parts also are
paler, especially on the throat and abdomen. The white tips to the tail
feathers, mentioned by Dr. Stejneger as characteristic of this form, are
equally common on specimens of occidentalis.

The following measurements show the relative size of the two forms:

^feasurements of Mijadestes ohscuriis insularia and Mi/adesfen o. occidentalis.

Name.

Locality.

Sex.

Num-
ber of
speci-
mens.

Wing.

Tail.

Cul-
men.

Tarsus.

Myadestes obscurns insii-
laris.

Myadestes obscums iiisu-
laris.

Myadestes obsciirus occi-
dentalis.

Maria Madre Island . .
do

Jalisco and Sinaloa . . .
do

ad. cT
ad. 9
ad. d"
ad. ?

5
3
3
2

102.7
98.6
104
100.5

102.6
95.6
102. 3

92

12.2
11.5
12.5

12

22.5

22

22.1

21.7

dentalis.

Hylocichla ustulata (Nuttall). Russet-backed Thrush.

A typical specimen of this si)ecies, taken on the islands by Colonel
Grayson in the winter of 1865, is in the National Museum. In his
notes Colonel Grayson says: "I found this little thrush in the mouth
of January quite abundant in the thickest of the woods of the Tres
Marias. It is very timid and shy, more so than any bird I saw ui^on
the islands ; it frequently uttered a low, jdaintive whistle, and seemed
solitary in its habits." We saw none of them on the islands in May,
and it is safe to class them as winter visitants.

Hylocichla ustulata swainsonii (Cabaiiis). Olive-backed Thrush.
Hylocichla ustulata alma Oberholser, Auk, XV, p. 304, October, 1898.

Two specimens of this thrush were taken on Marie Madre, one on
May 5, the other on May 19. They were found in the heavj^ forest back
from the coast, and evidently occur only as stray migrants.

Mr. Oberholser mentions these specimens as typical examples of his
subspecies, which is considered a synonym of Hylocichla u. swainsonii
by the American Ornithologists' Union.

Menila graysoni Ridgway. Tres Marias Roliiii.

Turdus Jlavirosiris Grayson, Proc. Boston Soc. Nat. Hist., XIV, p. 27fi, 1871 (part);

Lawr., Mem. Boston Soc. Nat. Hist., II, p. 266, 1874 (pan j.
Merula flavirostris (iraysoni Ridgway, Proc. U. S. Nat. Mus., V, p. 12, 1882.

Grayson's robin is one of the most abundant and widely spread resi-
dents and takes the place of M. flavirostris of the mainland, which it
closely resembles in habits and general appearance. Although a char-
acteristic bird of the islands, yet occasional stragglers reach the main-

BIRDS OF THE TRES MARIAS ISLANDS.

61

laDtl, as is sliowu by a perfectly typical specimen (a female in worn
plumage) taken at Santiago, Territory of Tepic, Jnne L*0, 1897. On the
islands it was found from the shore to the forests of the higher slopes
and was also very plentiful and familiar about the settlement. It had
a variety of notes, among them a rich Avarbling song and a character-
istic clear, mellow, whistling call. While among the trees, or during
their search for food upon the ground, these birds closely resemble the
common robin in habits and general appearance. At the time of our
visit a species of wild fig was in fruit, and the tops of the trees were
swarming with these robins, tanagers, orioles, lovebirds, and trogons,
all eagerly feeding upon the figs.

Merula graysoiii is another of the Tres Marias birds which are evi-
dently oifshoots from species now resident on the adjacent mainland,
but with differences sufficiently pronounced and constant to warrant
their recognition as separate species. Merula flavirostris, the main-
land representative of the Tres Marias robin, is much more richly
colored than f/raysoni, and the differences mentioned by Mr. IJidgway
are constant and well shown in the present series. The following aver-
age measurements show the relative dimensions of the two species:

Measuremejits of Merula graysoni and M. flavirostris.

Merula graysoni . . .

Merula graysoni

Merula flavirostris .
Merula flavirostris .

Locality.

Maria Madre Island . .

do

West coast Mexico . . .
do

Sex.

Num-
ber of •
speci-
mens.

Wing.

Tail.

Cul-
nien.

ad. cf

4

127

99.7

24.4

ad. $

5

125.6

98.8

24

ad. cf

i

125

99.7

21

ad. ?

5

124.4

98

2.?

34.5
34.5
32.6
32.4

BIEDS ERRONEOUSLY ATTRIBUTED TO THE TRES MARIAS.

Among the birds sent to the Smithsonian Institution from western
Mexico by Mr. John Xantus are five species of humming birds named
below which were not found on the Tres Marias either by Colonel Gray-
son or myself, and which are not known even from the adjacent parts
of the mainland. These specimens are now in the National Museum,
all labeled "Tres Marias, July, 1861." The improbability of their cap-
ture on the Tres Marias is very great, and the fact that species from
such widely separated areas should be credited to these islands during
a single month can be accounted for in only one way. Probably Mr.
Xantus purchased these specimens from some one who misled him con-
cerning their origin. That this could be done very easily I know from
personal experience. Some years ago I purchased a small collection of
birds from a San Francisco dealer, who claimed that they came from

62 NORTH AMERICAN FAUNA.

La Paz, Lower California, but which proved to be made up of species
found near Mazatlau, Siualoa.
Thalurauia luciee Lawr.

Thahirania liiciw liSbwr., Anu. Lye. Nat. HiBt., N. Y., VII, p. 2, 1867; Proc. Boston
.Soc. Nat. Hist., XIV, p. 284, 1871 ; Mem. Boston Soc. Nat. Hist., II, p. 291, 1874.
Described as new from the specimen sent in by Xautus, but proved
to be Thalurania glaucopis, a resident of southeastern Brazil.

Florisuga iiiellivora (Liun.).

Lawr., Proc. Boston Soc. Nat. Hist., XIV, p. 284, 1871; Mem. Boston Soc. Nat.
Hist., II, p. 291, 1874.

A well-known species of the humid tropics from southern Mexico to
South America. There is no authentic record for it in western Mexico,
and it is safe to say it has not been taken on the Tres Marias.

Uranomitra guatemalensis (Gould).

Lawr., Proc.' Boston Soc. Nat. Hist., XIV, p. 284, 1871; Mem. Boston Soc. Nat.
Hi8t.,II, p.202, 1874.

A species which ranges from Guatemala and British Honduras
southward. There is no authentic Mexican record.

Pstasopiiora thalassina (Swainaon).

Lawr., Proc. Boston Soc. Nat. Hist., XIV, p. 284, 1871; Mem. Boston Soc. Nat.
Hist., II, p. 292, 1874.

This humming bird ranges from the highlands about the Valley of
Mexico southward into Central America, but there appears to be no
authentic record for western Mexico.

Chlorostilbon insularis Lawr.

Chlorostilbon insularis Lawr., Anu. Lye. Nat. Hist. N. Y., VII, p. 457, 1867; Proc.
Boston Soc. Nat. Hist., XIV, p. 284, 1871; Mem. Boston Soc. Nat. Hist.,

II, p. 292, 1874.

This bird was described by Mr. Lawrence from a Xantus specimen,
but proved to be Chlorostilbon pueherani of southeastern Brazil.

Merula grayi Lawr.

Merula grayi Lawr., Proc. Boston Soc. Nat. Hist., XIV, ]». 276, 1871; Mem. Bos-
ton Soc. Nat. Hist., II, p. 266, 1874.

Grayson's notes on Merula grayi on the Tres Marias refer to pale
S]>ecimens of M. graysoni, and his record of 31. grayi at the city of
Tepic, on the adjacent mainland, refers to M. fristis. Merula tristis is
a common and widely spread species in suitable localities in western
Mexico and is the only Merula sent in by Grayson from the city of
Tepic.

Merula grayi, on the contrary, does not appear to occur anywhere in
western Mexico north of the Isthmus of Tehuantepec, for no specimens
were taken by Grayson nor, during our own work at many localities
between the Isthmus and Mazatlan, has a single individual been noted,
and there appears to be no authentic record of its occurrence there.
This thrush is a species of the humid tropics, ranging along both coasts
of Central America north to the Isthmus of Tehuantepec, and thence
northward its range is limited to the humid region of the Gulf coast
and adjacent mountain slopes of eastern Mexico,

REPTILES OF THE TRES MARIAS AND ISABEL ISLANDS.

By Lp^onhard Stejneger,
Curator, Division of Reptiles and Batrachians, U. S. National Afiiseidn.

The present pai)er is based upon the collection made on the Tres
Marias and Isabel Islands in April and May, 1897, by E. W. Nelson
and B. A. Goldman.

The surprising fact that the two expeditions which have collected
systematically in the Tres Marias brought home the same number of
species, Forrer only collecting one snake, Diplotropis diplotropis, which
Nelson did not collect, and Nelson also oollecting only one snake which
Forrer did not obtain, viz. Boa imperator^ seems to indicate that not
many more species than the 16 here enumerated are to be found in
these islands.

It will thus be seen that the reptile fauna is an exceedingly poor one
and very disappointing in several respects. Thus most of the species
are common on the opposite mainland and generally distributed over
tropical Mexico and Central America. Then, again, it seems as if the
species are practically identical on all the islands of the group. This
would indicate a comparatively recent severance of the islands from
each other as well as from the opj)osite mainland of Mexico.

It is worthy of note, perhaps, that there is absolutely no indication
of relation to the Cape Saint Lucas fauna of Lower California. The
only species occurring in both places is Fhyllodactylus tuherculosus, a
gecko of wide distribution, the presence of which is of absolutely no
moment in determining zoogeographical relations.

The only species whch seems to be peculiar to the islands is Cnemi-
dophorus mariarum. As will be explained more fully under the head
of this species, I have never seen a specimen from the mainland, and
those which have been recorded from there I regard as wrongly identi-
fied. However, the herpetology of the regions in question is too little
explored in detail to incline one to be dogmatic on a point like this,
but I may call attention to the fact that the swift which occurs on the
little Isabel Island, about halfway between the Tres Marias and the
mainland, is most certainly the same form which inhabits the latter,
viz, Gnemidophorus gularis mexicanuSj imd not C. mariarum, to which
it bears only a superficial resemblance. The species collected on Isabel
Island are referred to in the following i)aper without any number pre-
ceding the specific names. Mr. Nelson has contributed field notes on
some of the species, and these notes are given in brackets with his
initials at the end of the jjaragraph on the species to which they refer.

63

64

NORTH AMERICAN FAUNA.

TESTUDINATA.

[The tortoise-shell turtle frequents the sea about the Tres Marias,
approaching- the shores to mate and deposit eggs in May and June
eacli year. At the same time the large green sea turtle abounds along
these shores, where they congregate for the same purpose. — E. W. N.]

Kiiiosternon integrum Lecontu.

I have no hesitation in endorsing Boulenger's view (t)at. Chel. Brit.
Mus., p. 4L') that the Tres Marias mud turtles are K. integrnm and not A".
hirtipes, as held by Giinther (Biol. Oentr.-Am., Kept,, j). 15, pis. xii-xiv).
They have the broader bridge and broader plastron of the former and
agree with undoubted specimens from the mainland. The island speci-
mens, of which there are four adults and one young, do not differ from
those from Colima, Guanajuato, Cuernavaca (Morelos), Acaponeta
(Tepic), Guadalajara (Jalisco), Presidio, and Mazatlan (Sinaloa), from
all of which localities I have examined specimens. K. hirtipe.s I believe
to be confined to the eastern side of Mexico.

List of specimens of Kinosttrnon inteffruin.

U. S. Na-
tional
Museum
number.

Collect-
ors' num-
ber.

Locality.

Date.

24606

712
713
714
715
716

May 15,1897
May 15,1897
May 15,1897
May 15,1897
May 15,1897

24607

do

24608

do

24609

do

24610

do

LORICATA.

Crocodylus americanus Laiir.

ISTo specimens were secured, but Mr. Nelson assures me that the
crocodile occurs on Maria Magdalena Island. There can be but little
doubt that it is the present species which is distributed all along the
coast of Central America, Mexico, the West Indies, and southern
Morida.

[The unmistakable furrow in the mud where a crocodile had hauled
up on the border of a brackish lagoon on the eastern side of Maria Mag-
dalena, the sight of a small head in the water, and the testimony of
the people on Maria Madre established the fact of their occurrence.
They appeared to be limited to Maria Magdalena. — E. W. X.J

SQUAMATA.

SAURI.
Phyllodactylus tuberculosus Wiegm.

This species is distributed over Mexico and Central America, and
has also been collected in the Cape Saint Lucas region of Lower Call-

REPTILES OF THE TKES MARIAS ISLANDS.

65

foruia, the specimeus from the latter locality haviug been described by
Cope as Phyllodactylus xanti.

List uf specimens of rhi/lludactiilus iiihircalonus.

IT. S. Xa-

tioual
Mu.seuiii
number.

Collect-
ors' IIUIU-

ber.

Locality.

Date.

24611
24612
24613

669
'686

700

Maria Madre Island May 21, 1897

do May 28, 1897

.Maria Cleofa Islaud May 30, 1897

' No. 686 was taken iu an old house.

Auolis nebulosus Wicgm.

All the specimeus from the three islands are normally colored and
alike, except No. 092, which has a wide whitish dorsal band originating
on the occiput and extending down the upper surface of the tail. It is
edged with dusky, and a narrow broken line of the same dusky color
in the white band near the edge on each side extends from neck to
rump. This specimen is small and without gular pouch; but No. 691,
from the same island, which equals it in these respects, is colored like
the larger specimens. Both specimens a])pear to be females, having no
enlarged postanal scales.

This species is widely distributed over Mexico, and has been collected
in the Tres Marias Islands not only by Forrer but also by Capt. William
Lund, specimens from the latter being in the museum of the California
Academy of Sciences iu San Francisco (Van Denburgh, I'roc. Phila.
Acad., 1897, p. 4G0).

List of specimens of Anolis nehulosus.

IT. S. Xa-

tioiial
Museum
uuinber.

Collect-
ors' num-
ber.

Locality.

Date.

24614

036

041

1083

'684

'685

688

090

091

692

May 3, 1897
May 4, 1897
May 28,1897
Mav 28, 1897

24615

do

24610

24617

... do

24618

do

Mav 28, 1897

24619

do

May 28,1897
May 29,1897
Mav 29, 1897

24620

24621

do

24022

. ...do

May 29,1897

' No.s. 683-685 wore found living in au olil liouse.

Ctenosaura teres (Harlan). Black Iguaua.

The material at hand is very unsatisfacitory inasmuch as all the
full-grown specimens are of the same sex and iu rather poor state of
preservation, while the younger specimens afford no characters for
13950— No. U 5

66

NORTH AMERICAN FAUNA.

satisfactorily separating tbe various forms which naturally group
themselves around Ctcnosaura teres. They are therefore left under
that general name for the present, the writer hoping some day to be
able to review the whole genus. The chief difficulty now lies in the
lack of typical specimens of C. teres from Tampico and from the east-
ern coast of Mexico generally, and until a series of full-grown speci-
mens of both sexes is obtained from tliat region it will be futile to
attempt to straighten out the nomenclature of these lizards. As far as
I can make out from my defective material the Tres Marias and Isabel
specimens differ sufficiently from specimens from Oolima and Tehuan-
tepec to warrant their subspeciflc recognition, but whether identical
with the Mazatlan form or not I am not able to say. There are certainly
several pretty well defined races of this species; but more adult speci-
mens and a direct comparison with the types of many of the old names
in various foreign museums will be necessary before the intricate ques-
tions involved can be settled.

[The females were burrowing in the gravel in dry washes and flats
on the islands the last half of May. The burrows were from 2 to 3
or 4 feet deep, and after the eggs had been deposited at the lower
end, the female scraped in loose gravel until the hole was filled, and
frequently raised a little mound over the entrance. — E. W. oST.]

TJ. S. Na-
tional
Museum
nmulier.

Collect-
ors' num-
ber.

Loealitj-.

Date.

24623

655
656
659
660
■ 662
(bis) 662
676
693
630
631
632

May 14,1897
May 14,1897
May 15,1897
May 15,1897
May 15,1897
May 17,1897
May 24,1897
May 29,1897
Apr. 23,1897
Apr. 23,1897
Apr. 23,1897

24624

do

24625

do

24626

do

24627

do

24628

do

24629

do

24630
24631
24632

Maria Cleof:. Island

Isabel Island

do .

24633

..do

Uta lateralis Boukmi^er.

Mr. i^elson remarks that this species lives on stones and driftwood
near the border of the woods along the sea beaches.

Uta lateralis was based by Bouleuger in 1883 upon specimens from
the Tres Marias and from Presidio, near Mazatlan, collected by Forrer,
and specimens from both localities are designated as 'types' in the
' Catalogue of Lizards in the British Museum.'

REPTILES OF THE TRES MARIAS ISLANDS.

67

V. S. Na-
tional
Museum
number.

Collect-
ors'num-
ber.

Locality.

Date.

24034

635
642
643
653
670
671
672
673
674
675
678

Maria Madro Island

May 3, 1897
May 4, 1897
May 7,1897
May 13,1897
May 21,1897
May 21,1897
May 21,1897
May 21,1897
May 21, 1897
May 21,1897
May 25,1897
May 25,1897

24635

do

24636

do

24637

do

24638

do

24639

do

24640

do

24641

do

24642

do

24643

do

24644

do

24645 679

do

Sceloporus boulengeri Stejneger.
N. Am. Fauna No. 7, 1893, p. 180, pi. I, figs. 5a-c.

This species appears to be smaller than iS. clarMi, of wbicb it is the
southern representative. A full-grown mal^ (No. 634e) measures only
72mni from snout to vent.

Van Denburgh's belief that iS. boulengeri "is the same form as Cope's
8. oligoporus^^ (Proc. Phila. Acad., 1897, p. 463) is not well founded. The
latter is easily distinguished by having only 2 to 3 femoral pores, besides
other differences. It is probably identical with S. horridus.

U. S. Na-
tional
Museum
number.

Collect-
ors' num-
ber.

Locality.

Sex.

Number
of pores.

Date.

24646

634
634 a
634 &
634 c
634 d
634 e

d ad....
? jun...
d" adol..
? ad....

?

d ad....

8
9
7
9
9
9

Apr. 23,1897
Apr. 23,1897
Apr. 23,1897
Apr. 23,1897
Apr. 23,1897
Apr. 23,1897

24647

do

24648

do

24649

do

24650

do

24C51

do

Cnemidophorus mariarum Giinther.

Cnemidopliorus mariarum Giinther Biol. Cunt. -Am., Kept. p. 28, pi. XX, April, 1885;
Boulenger, Cat. Lizards, Brit. Mus., p. 368, 1885.

The swifts from the Tres Marias are essentially alike. Those from
Maria Madre are the largest and possibly also most distinctly marked;
those from the small detached rock oft' the west side of Maria Cleofa
as well as the one from the main island of that name are somewhat
smaller. According to Mr. Nelson's observation those from the detached
islet, which is a bare rock, the nesting place of numerous sea birds,
appeared to him paler when alive than those on the other islands, but
now, in alcohol, the difterence, if any, is very slight.

This species, which was originally described by Giinther from speci-
mens collected by Forrer on the Tres Marias seems to be confined to
this group of islands. If so^ it is the only species of reptile hitherto

68

NORTH AMERICAN FAUNA.

collected which is peculiar to these islands. The species has been
recorded from the inaiuland(by Van Deiiburgh, Proc. Phila. Acad., 1897,
p. 4G3, who identifies "a large number of lizards from Mazatlan, Sau
Bias, and Tepic" with Giiuther's species), but I am satisfied that these
records are based n[)on specimens of C. gularis mexicanus (Peters)
which superficially very much resemble the island species. The mis-
identification is probably due to the fact that Cope, in his monograph
of the genus, overlooked the different keeling of the caudal scales
which is the essential character of this species.

List of /specimens of Cnemidophorus marianivi.

U. S. Na-
tional

Museum
number.

CoUect-
ors' num-
ber.

24652

637

24653

638

24654

639

24655

640

24656

644

24657

645

24658

646

24659

647

24660

687

24661

701

24662

702

24663

703

24664

704

24665

705

24666

706

Locality.

Date.

Maria Madre Island.
do

-do.
.do.
.do.
.do.
.do.
-do.

rock) .

Maria Magdalena Island

Maria Cleofa Island (outlyin

do

do

do

do

Maria Cleofa Island (main island)

May

3, 1897

May

3, 1897

May

4, 1897

May

4, 1897

May

7, 1897

May

7, 1897

May

7, 1897

May

7, 189T

May

28, 1897

May

30, 1897

May

30, 1897

May

30, 1897

May

30, 1897

May

30, 1897

May

31, 1897

Cnemidophorus gularis mexicanus (Peters).

The Isabel Island swifts are identical with the mainland form, two
specimens of which were brought from San Bias. They are quite dis-
tinct from the species on the Tres Marias, which is well characterized
by the smaller femorals and the parallel caudals. It is strange that
Cope, having had the latter character clearly pointed out by Boulenger,
should have referred G. mariarum to G. gularis as a subspecies.

List of apedmens of Cnemidophorus gularis mexicanus.

U. S. Na-
tional
Mu.seum
number.

Collect-
ors' num-
ber.

Locality.

Date.

24607
24068
24669
24670
24671

633

633a

6336

633c

633d

Isabel Island .

do

do

do

do

April 23,1897
April 23,1897
April 23,1897
April 23, 1897
April 23,1897

REPTILES OF THE TRES MARIAS ISLANDS.
SERPENTES.

69

Boa imperator Daudin.

This is the first record of this species from the Tres Marias. The
species is generally distributed through southern Mexico and Central
America.

Scale rows 73.

List of specimens of Boa imperator.

V. S. Na-
tional
Museum
number.

Collect-
ors' num-
ber.

Locality.

Date.

24672

648

Maria Madre Island

May 12,1897

Oxybelis acuminatus (Wietl).

A coiuiuou species occurring all through tropical America from
Guaymas, Mexico, south.

List of specimens of Oxyhelis acuminatus.

U. S. Na-
tional
Museum
number.

Collect-
ors' num-
ber.

Locality.

Date.

24673

677

May 25,1897

Diplotropis diplotropis (Giinther).

This species seems to be confined to western Mexico. It was not
collected by Mr. Nelson, but there are two specimens in the British
Museum collected by Forrer on the Tres Marias {Leptophis diplotropis
Boulenger, Cat. Snakes Brit. Mus., II, p. 110).

Drymobius boddaerti (Seetzen).

A common species distributed over tropical America.

In No. 681 the fourth labials on both sides are divided horizontally,
so as to suggest a subpreocular. This is an adult male, and is uni-
formly colored above, without any markings. The adolescent specimens
are uniform above, with a few scales tipped with black; the anterior
part of the underside has square blackish spots. The two young ones
have above brown, dark-edged, squarish spots, separated by narrow
light-colored interspaces. They are marked underneath like the adoles-
cent specimens.

No. G81, male ad. — Scale rows, 17; ventrals, 183; anal, h ; caudals, |||;
supralabials, 9. ■ ,

70

NORTH AMERICAN FAUNA.

List of specimens of Drymohius hoddaerti.

U. S. Na-
tional
Museum
number.

24674
24675
24676
24677
24678
24679

Collect-
ors' num-
ber.

652 juv.
654 ad.
658 ad.
661 ad.
663 juv.
681 ad.

Locality.

Maria Madre Island

do

do

do

do

Maria Magdalena Island .

Date.

May 12,1897
May 13,1897
May 14,1897
May 15,1897
May 18,1897
May 27,1897

Bascanion lineatum Bocourt.
This species is apparently confiued to western Mexico.

List of specimens of Bascanion lineatum.

U. S. Na-
tional
Museum
number.

Collect-
ors' num-
ber.

Locality.

Date.

24680

850
651
660

May* 12, 1897
May 12,1897
May 16,1897

24681

... do

24682

... do

Drymarchon corais melanurus (Dum. &. Bibr.)

Scale rows, 19; veutrals, 205; anal, 1; caudals, f|, siipralabials, 8.
Adult male with the characteristic coloring of this subspecies, which
seems to be confined to Mexico and Central America.

List of specimens of Drymarchon corais melanurus.

V. S. Na-
tional
Museum
number.

Collect-
ors' num-
ber.

Locality.

Date.

24683

664

May 18,1897

Lampropeltis micropholis oligozona (Bocourt).

Scale rows, 23 ; veutrals, 230; anal, 1; caudals, f|; temporals, 2 + 3.
Adult male. Thirteen annuli on body, separated by wide, red inter-
spaces, without black spots, both on back and belly; all the annuli
complete, including that on neck and throat, which does not touch the
parietals; snout white, with black on rostral and anterior nasal. From
Boulenger's account it appears that Forrer's specimens from the Tres
Marias are identical. (Oat. Snakes Brit. Mus., II, p. 204.)

Distributed over Mexico and Central America.

List of specimens of Lampropeltis micropholis oligozona.

U. S. Na-
tional
Museum
number.

Collect-
ors' num-
ber.

Locality.

Date.

24684

661

May 16,1897

REPTILES OF THE TRES MARIAS ISLANDS.

71

Agkisti-odon bilineatus (Giinther).
Scale rows, 23 ; veil trals, 138; anal, 1; caudals, 21 + ff . Adult male.
Southern Mexico and Central America to Nicaragua.

List of specimens of Agkistrodon bilineatus.

U. S. Na-
tional
Museum
number.

Collect-
ors' num-
ber.

Locality.

Date.

24685

707

May 15, 1J>97

Crotalus sp. ?

No rattlesnake was collected on the Tres Marias by Foner, nor by
Nelson, but the latter informs me that he was told of the occurrence of
a rattler on Maria Masdalena Island.

NOTES ON THE CRUSTACEA OF THE TRES MARIAS ISLANDS.

By Mary J. Ratiibun,
Second Assistant Curator, Dirinon of Marine Invertebrates, U. S. National Museum.

Of the four species of Crustacea taken by E. W, Nelson and E. A.
Goldman on the Tres Marias Islands in May, 1897, two are identical
with forms inhabiting Lower California, one is found in all the warm
countries of the world, while the fourth, a fresh-water shrimp, is dis-
tributed throughout tropical America.

Gecarcinus digueti Bouvicr.

Gecarcinus dif/ueli Bonvier, Bull. Mna. Hist. Nat., Paris, I, 8, 1895.
Maria Clcofa, Island. May 30. One large male (Collectors' No. 717).

The type and only specimen hitherto collected is from Lower Cali-
fornia, and is in the Paris Museum. This species differs from others
found on the Pacific coast in its wider carapace, narrower front, longer
legs, and in the form of the abdomen of the male.

Measurements.

o

o

CD '

o

= 1-

•r

3

s

.D

2 o

3

t-)

o

Specimen.

i

3

s-3

a

u

1

a

a

'3

a

.a

Xi

"3

o

o 3

o

o

o

.a

o

o

o

.a

Cm

O

Ml

a

g

.<!>

n o

'd

r3

a

r3

a

•a

h^

'i

W

q

M

t-J

^

iJ

^

i-J

■^

Hi

^

Type, cf, Lower Cali-

46.3

70

09
104

25
37.5

9
13

33
46.3

9.7
13.4

16
22.5

8
11.5

•17
21.5

7.3
10

24.2
33.5

3.7

cT, Maria Cleofa Island..

5

The measurements of the legs are exclusive of the large spines, and
the length given is that of the anterior or superior margin. The penult-
imate segment of the abdoinen of the male is very wide. Length and
distal width, 12 mm.; xjroximal width, 21.5.

Mr. Nelson says of these crabs :

On the Tres Marias we foimd them only on Maria Cleofa, Avhero they -R-ere very
nnnierons above high-vrater mark on the sandy beaches of the low eastern part of the
island. They were also living very al)nndnntly in burrows in the soft soil almost
everywhere on the slopes of Isabel Island. They are nocturnal in habits, and caused

73

74 NORTH AMERICAN FAUNA.

some annoyance by walking over us at night while we were camped iu their haunts.
They began to como out of their burrows as soon as it became twilight in the even-
ing. In both localities most of their burrows were found among the scrubby bushes.
On Isabel Island they were often seen during the day sitting in the burrows a foot
or so from the entrance, but scuttled back to a safe depth when I approached.

Ocypode ' occidentalis Stimpsou.

Oci/poda occidentalis Stimpsou, Ann. Lye. Nat. Hist. N. Y., VII, 229, 1860.
Maria Magdaleua Island. May 28. One female (No. 689).
Maria Cleofa Island. May 30. One male (No. 699).

This much neglected species is distiuct, it seems tome, from 0. huhlii
de Haau, of which Miers made it a variety. According to the descrip-
tion of 0. Jcuhlii given by de Man (Notes Leyden Mus., Ill, 250, 1881),
who had the type before him, 0. occidentalis differs from it in having a
narrower carapace, in the outer orbital angle directed inward and not
outward, iu the shorter hand, the length of the upper margin of the
palm being less than the width, and iu having from 18 to 21 tubercles
in the stridulating ridge (de Man gives 8 or 10 for kuJiUi, while Miers
figures 17). The form of the abdomen of the male furnishes excellent
characters for the determination of the species of Ocypode. In 0. occi-
dentalis the penultimate segment is much wider at its middle than at
its proximal end.

It is singular that this species is not mentioned in the revisions of
the genus by Kiugsley, 1880, or by Ortmann, 1897.

Dimensions of a type specimen^ U. S. National Museum. — Male: Length,
40.5 mm.; epibranchial width, 48; exorbital width, 41; length of supe-
rior margin of palm, 22.8; entire length of propodus, 43.5; greatest
width, 24.

^ctwr/c— Type locality, Cape St. Lucas. Also taken at Turtle Bay
and San Jose del Cabo, Lower California, by Mr. A. W. Anthony, in
1896 and 1897.
Grapsus grapsus (Linnjeus).

"This crab was very abundant on the rocks along the water's edge
on the Tres Marias as well as on Isabel Island." (Nelson.)

The species is distributed throughout the tropics.

Bithynis jamaicensis (Herbst).

Maria Magdalena Island. May 27. One adult, 7 young (No. 709).
Maria Cleofa Island. May 30. One adult, 3 young (No. 710).

"These shrimps were very numerous in a small stream among the
hills in the interior of Maria Magdalena, and were also numerous in
streams iiowing through the hilly parts of the adjacent mainland."
(Nelson.)

The species is found on the Pacific slope of the continent from Lower
California to Ecuador, and on the Atlantic slope from Texas to Rio
de Janeiro. The following localities, not before recorded, are repre-
sented by specimens in the U. S. National Museum: On the Pacific

Wcypode, not Ocypoda, Fabricius, Entom. Sys., Suppl., 312 and 347, 1798; also
Entom. Sys., emend, et auct., IV, index, 115, 1796.

CRUSTACEA OF THE TRES MARIAS ISLANDS. 75

coast, La Paz, Lower California; Rio Presidio, Siualoa; Eio de Alica,
Tepic; Barranca Ibarra, Rio Santiago, Jalisco, and Rio Arnieria,
(Jolima, Mexico; Riode los Platanales and Quebrada Cbavarria Golfito
(both tributary to the Gulf of Dulce), Costa Rica; River David, Chiri-
qui, United States of Colombia, 4,000 feet elevation; Guayaquil,
Ecuador. On the Atlantic coast, San Antonio, Tex. ; Las Moras Creek,
Kinney County, Tex. ; Brownsville, near mouth of Rio Grande, Tex. ;
Amixtlan, and Zacatlan, Puebla, Mexico; Escondido River, 50 miles
from Bluefields, Nicaragua; Port Castries, St. Lucia, West Indies.

The west African form, B. vollenhovenli (Herklots) is no more than
a subspecies of B.jamaicensis. It differs only in the slenderer second
pair of feet, the carpal and meral joints of which are subequal. The
relative lengths of the rostrum and the antennal scales and peduncles
agree with those in some specimens of jamaicensis. The two forms are
considered identical by Dr. Ortmann.

According to Dr Edward Palmer, B. jamaicensis is much eaten at
Colima, and is offered in the market there as a choice article of food,
especially on Fridays and Sundays.

PLANTS OF THE TRES MARIAS ISLANDS.

By J. N. Robe,
Assistant Curator, Dirision of Plants, U. S. National MuHetimJ

Tlie Tres Marias, lyiug about 65 miles off the west coast of Mexico
ill about 22"^ north hititude, are amoug the last of the west coast islands
to be studied. All the others have yielded valuable botanical results,
but almost nothiuj? has been known of the flora of these islands
except in a commercial way. Several botanical expeditions had been
planned to explore the islands,- but heretofore none had succeeded in
reaching them. They are out of the line of traffic, although some of
the smaller steamers stop now and then for fuel, and small boats
occasionally ply between the islands and San Bias. They are usually
visited during the dry season, as it is dangerous to attempt the passage
during summer and autumn.

Mr. I^elson visited the islands at the very close of the dry season,
when the vegetation is at its poorest, and this accounts for the small
number of species collected. His collection contains 154 numbers (i^os.
4179 to 4333) and 136 species, mostly from Maria Madre, the largest
of the islands, and only a few from Maria Magdalena and Maria
Cleofa. In the subjoined list the j)lants are from Maria Madre unless
otherwise stated.

There are no cultivated plants on the islands, except one or two
grasses. Fithecolohium dulce, perhaps introduced, is common and
much prized for its delicious fruit. The exportation of Spanish cedar
{Gedrcla .sp.) has long been the chief source of income for the islands,
but the available supply of this timber is now nearly exhausted. Tlie
Hora is purely tropical and does not differ essentially from that of the
adjacent mainland. Many of the species have not been reported from
the mainland opposite, but this is doubtless because the flora is not
well known, since these species have been collected either farther north
or south. One hundred and twelve species are named below, of which
11 are new. Many of them have a wide distribution in tropical
America; all but 6, except the new si^ecies, have heretofore been
reported from Mexico; 24 range northward into the United States;
64 extend into Central America; 61 into South America; 44 into the
West Indies^ and 21 are found in the Old World.

' Published by permission of the Secretary of the Smithsonian Institution.

77

78' NORTH AMERICAN FAUNA.

The following report must be regarded as a i)reliminary one. The
sijecimens ui)on which it is based are simply those in fruit or flower at
the close of the dry season, a considerable number of which have not
been determined specifically and a few not even generically. As will
be seen from the list below, mostly trees and shrubs were collected,
while the herbs, which spring up in great variety during the rainy
season, are scarcely represented.

The Gamopetalte and Apetalai have been named by Mr. J. M. Green-
man, GramineoB by Prof. F. Lamson-Scribuer, and Filices by George E.
Davenport.

The following new species and varieties are based on this collection:

j3Sgi2)hila jxicijica Greenuiau. Euphorhia suhcccrttlea iresmariw Millsp.

BeJoperone nehoni Greenman. Gilibertia insula7-is Rose.

Buxu8 puhescens Greenman. Pilocarpus insularis Rose.

Cordia insularis Greenman. Ternstrocmia malthya Rose.

Erythrina Janata Rose. Zanthoxylum insularis Rose.

Euphorhia nehoni Millnpangli. Zanthoxylum nelsoni Rose.

ANNOTATED LIST OF SPECIES.

Cissampelos pareira L.

Common in Mexico and other tropical countries. May 3 to 25 (Nos.
4233 and 4262).

Argemone ochroleuca Sweet.

Widely distributed throughout Mexico. Maria Magdalena Island,
May 2G to 28, 1897 (No. 4318).

Capparis cynophallophora L.

Found along the coast of Mexico, South America, and the West
Indies. May 3 to 25 (No. 4302).

Capparis breynia L.

Common in Mexico, South America, and the West Indies. May 3
to 25 (No. 4219).

Crataeva tapia L. ?

Perhaps this is the species which has been reported from Acapulco
and Mazatlan. May 3 to 25 (No. 4274.)

Ternstroemia maltbya Rose, sp. nov.

Tree 3 to 9 meters high; leaves obovate, entire, obtuse, glabrous,
thickish, not black-punctate beneath, 5 to 10 cm. long; flowers solitary;
peduncles 2.5 to 3.5 cm. long becoming curved, bracteate a short dis-
tance below the calyx; sepals 5, orbicular, 8 to 10 mm. in diameter;
petals united at base, acute; stamens numerous; fruit (immature) ovate,
20 mm. long, two-celled; seeds red.

This species is in all i^robabilitj^ Seemau's No. 2148, collected on
the road from Mazatlan to Dnrango and enumerated in the Biologia
Centrali-Americana without specific name,

PLANTS OF THE TRES MARIAS ISLANDS. 79

Collected on Maria Madre Island, May, 1897, by T. S. Maltby (No.
105) and E. W. Nelson (No. 4242); by J. N. Eose near Colorao, Sinaloa,
July, 1897 (No. 1675).

Wissadula hirsutiflora (Presl) Rose.

The type of tbis species came from Acapulco. It is probably com-
mon on the west coast, although its distribution and specific limits are
not well known. May 3 to 25 (No. 4250).

Abu til on reventum Watsou.

This species extends as far north as Arizona. May 3 to 25 (No. 4203).
Hibiscus tUiaceus L.

A common tree in most tropical couiitries. Maria Magdalena Island,

May 26 to 28 (No. 4328a).

Melochia tomentosa L.

Common throughout tropical America. May 3 to 25 (No. 4205).
Guazuma ulmifolia Laui.

Common throughout tropical America. Maria Magdalena Island,
May 26 to 28 (No. 4325).

Heteropterys floribunda H. B. K.
Common in Mexico and Central America.
Maria Magdalena Island, May 26 to 28 (No. 4323).

Guaiacum coulteri ? Gray.

Seemingly common on the west coast of Mexico. Island specimens do
not corresjiond with the form found on the mainland and may represent
an undescribed species. May 3 to 25 (No. 4180).

Zanthoxylura insularis Rose, sp. nov.

Tree 6 to 20 meters high, thornless; leaves oddly pinnate; leaflets
6 to 7 pairs, opposite, sessile, obovate to spatulate, obtuse or retuse,
2 to 3.5 cm. long, creuate, with large pellucid dots between the teeth
and small scattered dots over the surface, glabrous; flowers unknown;
fruit small, in a rather compact i^anicle; pedicels very short; stipe
short and thick.

Collected by E. W. Nelson on Maria Madre Island. May 3 to 25, 1897
(No. 4278).

Zanthoxylum iielsoui Rose, sp. nov.

Tree 7.5 to 20 meters high, thornless(?); leaves oddly pinnate; leaf-
lets about 6 pairs, distant, opposite, shortly petioled, 5 to 11 cm. long,
rounded at base, long-acuminate, crenate, glabrous on both sides,
thickly set with pellucid dots; inflorescence in small compact panicles;
perianth complete; petals 4 ( ?) ; fruit large in dense head-like clusters,
not stipitate.

A very peculiar species, unlike any Mexican one known to me. Col-
lected by E. W. Nelson on Maria Madre Island. May 3 to 25, 1897
(No. 4279),

80 NORTH AMERICAN FAUNA.

Pilocarpus insularis Rose, sj). nov.

Tree 3 to 6 meters liigb, glabrous throughout; leaflets usually in
threes, some solitary or in rows, 5 to 7.5 cm. long, retuse at apex,^
cuueate at base, in the lateral ones more or less oblique; mid vein
prominent, lateral veins indistinct below, not very prominent above ;
racemes short and compact, 5 to 10 cm. long; fruiting pedicels hori-
zontal, 16 mm. long; ovary deeply 4 or 5-lobed or parted, 1 to 4 lobes
not maturing.

This species is near P. longipes of Mexico, but with somewhat diflereut
leaves, more compact inflorescence, etc. Collected by E. W. Nelson on
Maria Madre Island. May 3 to 25, 1897 (Ko. 4307).

Amyris sp.

May 3 to 25, 1897 (No. 4237).
Picramnia sp.

A tree 4.5 to 7.5 meters high; flowers said to be greenish, but none
with si^ecimeus. Much resembling the South American species i'. ciUata
Mast., but without flowers or fruit. Exact identification is doubtful.
May 3 to 25 (No. 427G).

Ochna sp.

May 3 to 25 (No. 4238).
Bursera gummifera Jacq.

Common throughout tropical Mexico, Central America, the West
Indies, and extending into Florida. May 3 to 25 (No. 4227).

Guarea sp.

May 3 to 25 (Nos. 4222 and 4230).
Trichilia spondioides Swartz.

Common in tropical America. May 3 to 25 (Nos. 4214 and 4309).
Xhneuia aniericana L.

Common in most tropical countries. May 3 to 25 (No. 4224).
Schoepfia schreberi Gmel.

Seemingly rare, but has been collected in Mexico and South America.
May 3 to 25, 1897 (No. 4271).

Hippociatea sp.

Maria Magdalena Island, May 20 to 28 (No. 4320). Maria Madre
Island, May 3 to 25 (No. 422G).

Colubrina arborea Braudegec.
Reported from Lower California and the west coast of Mexico. ^lay

3 to 25 (No. 4213).

Cissus sicyoides L.

A common species in tropical America. May 3 to 25 (No. 4198).
Serjania mexicana Willd.

A common species in tropical America. May 3 to 25 (No. 4231).

PLANTi> OF THE TRES MARIxVS ISLANDS.

81

PauUinia sessilifloia Radl.

Heretofore only known from the type specimens collected by Dr.
Edward Palmer in the State of Oolima, Mexico. May 3 to 25 {No.
4310).

Urvillea ulmacea H. B. K.

Common in Mexico and northern South America. May 3 to 25 (ISTo.
4277).

Cardiospermum corindum L.
A widely distributed, species. Maria Maj,'dalena Island, May 2G to 28

(No. 4328).

Crotalaria lupulina ? H. B. K.

Perhaps this species, which is common in Mexico, and extends into
the United States. May 3 to 25 (No. 4248).

Tephrosia sp.
May 3 to 25 (No. 4193).

Desmodmm sj).
May 3 to 25 (No. 4287).

Erythrina lanata Rose, sp. iiov.

A small tree, 4.5 to 7.5 meters hig^h, with a trunk 10 cm. in diameter;
branches glabrous, bearing mostly single infrasti[)uhir spines; leaflets
triangular, shortly acuminate, 5 to 10 cm. long, 5 to 7.5 cm. broad,
glabrous or nearly so.

/

Fig. 1 Erythrina lanata; a, calyx; &, banner; c, keel; rf, wing; e, stamena; /.ovary.

Inflorescence unknown; calyx lanate becoming glabrate, tubular, 10
to 13 mm. long, truncate, one-toothed; banner 68 mm. long, folded,
densely white-lanate, rounded at apex; wings (9 mm. long) and keel
(10 mm. long) included within the calyx; ovary densely lanate; legume
glabrous, 12.5 to 15 cm. long, strongly constricted between the seeds,
long-stipitate, attenuate at tip; seeds small (for the genus), nearly
orbicular, G to 8 mm. long, bright scarlet, with a dark spot at the
micropyle.

13950— No. 14 G

82 NORTH AMERICAN FAUNA.

The type of this species is Dr. Edward Palmer's No. 120, from
Acapulco, Mexico, collected in 1894-95. To this species I would refer
specimens collected by Frank Lamb near Villa Union, State of Sinaloa,
January, 1893 (No. 428), and flowering specimens by W. C. Wriglit
from the head of Mazatlan Eiver, January, 1889 (No. 1292), and also
those collected by J. N. Kose at liosario, Sinaloa, July 10, 1897 (No.
1592), and July 22 (No. 1822). The latter two specimens are not in
flower and their reference here is attended with some doubt. The
seeds are larger and the pods less constricted between the seeds. Here
also belongs E. W. Nelson's No. 4303 from the Tres Marias, collected
May, 1897. I have tentatively referred to this species E. W. Nelson's
No. 2099, taken at an altitude of 480 meters, near Santo Domingo, State
of Oaxaca, June 18, 1895. It has similar pods, but is described as
being but 0 to 12 cm. high and has more bluntish leaflets.

Dr. Palmer says this tree flowers in January, and is often used for
hedge fences. It differs from all other Mexican species which I have
seen in its white lanate banner. Its one-toothed calyx suggests E,
rosea, but in the latter the calyx is described as obli(i[uely truncate.

Phaseolus sp.
Maria Magdaleua Island, May 26 to 28, 1897 (No. 4319).

Canavalia gladiata DC.

A species of wide distribution, perhaps throughout tropical America.
May 3 to 25 (No. 4190).

Rhynchosia minima DC.

A common Mexican species extending into South America and the
United States. May 3 to 25 (No. 4200).

Rhynchosia precatoria ( ?) (H. B. K.) DC.

This species has been reported from Acapulco and Panama. May 3

to 25 (No. 4179).

Lonchocarpus sjj.

May 3 to 25 (No. 4310).
Ateleia (?) sp.

Without flowers or named specimens for comparison it is impossible
to name this plant definitely. If it belongs to the genus Ateleia it is
perhaps A. pierocarpa, the only species known from Mexico. A shrub
or small tree 3.5 to 10.5 meters high. May 3 to 25 (No. 418()).

Cassia emarginata L.

Common in Mexico, South America, and the \Vest Indies. May 3
to 25 (Nos. 4192 and 4297).

Cassia bifloia L.

Common in trojncal America. May 3 to 25 (Nos. 4194 and 4190).
Cassia atomaria L.

Common in Tropical Mexico and South America. Maria Magdaleua
Island, May 20 to 28 (No. 4321).

PLANTS OF THE TRES MARIAS ISLANDS. 83

Bauhiiiia sp.

Apparently belonging- to the genus Bauhinia, but very unlike any of
the Mexican species with which I am familiar. A vine G to 9 meters
long; only iu fruit. May 3 to 25 (No. 4300).

Acacia si).

This appears to be an undescribed species, of whicli J collected
specimens on the mainland. May 3 to 25 (No. 4188).
Albizzia occidentalis Braudegee.

Probably the above species, which is found in Lower California and
has been reported from western Mexico. May 3 to 25 (No. 4252).

Pithecolobium dulce Bentli.

Common in tropical Mexico and South America. Often cultivated.
May 3 to 25 (No. 4285).

Pithecolobium ligustrinum Klotzscli.

Common in tropical Mexico and northern South America. IMaria
Magdalena Island, May 26 to 28 (No. 4314).
Conocarpus erectus L.

Common throughout tropical America extending into Florida and
reported from tropical Africa. May 3 to 25 (No. 4220).

Psidium sp.

Tree 6 to 9 meters high; flowers white; called 'palo prieto.' This
species is not represented in the National Herbarium. May 3 to 25
(No. 4306).
Casearia corynibosa ( ?) H. B. K.

The Tres Marias specimens should probably be referred to this species
although our herbarium material seems to represent more than one
species. This form is common on the west coast of Mexico and Central
America. May 3 to 25 (Nos. 4270 aiul 4308).

Casearia sylvestris Swartz.

Widely distributed throughout tropical Mexico, South America, and
the West Indies. May 3 to 25 (No. 4241).

Casearia sp.
Maria Magdalena Island. May 26 to 28, 1897 (No. 4326).

Passiflora sp.
May 3 to 25 (No. 4249).

Opuutia sp. _
May 3 to 25 (Nos. 4263 and 4286).

Gilibertia insularis Rose, sp. uov.

Tree 6 to 12 meters high; leaves 25 to 35 cm. long, including the
slender petioles (7 to 18 cm. long), 9 to 20 cm. broad, entire or 3-lobed,
oblong, rounded at base, rounded at apex or with a short acumination,
glabrous, 3-nerved at base; fruiting inlioresceuce a short dense panicle;

84 NORTH AMERICAN FAUNA.

rays 2 to 3 cm. long; pedicels 4 to 8 mm. long; fruit white, 6-lobed,
4 nun. liigb ; styles short, connate to near the top.

Collected on Maria Madre Island May 3 to 25 (No, 4282).

Portlandia pterosperma Watson.

A sj^ecies recently described by Dr. Watson, the type coming from
near Guaymas, Sonora. May 3 to 25 (No. 4211).

Eupatorium sp.

May 3 to 25, 1897 (No. 4225).
Eupatorium sp.

May 3 to 25, 1897 (No. 4244).
Eupatoriuni coUinum DC.

Common in Mexico and Central America. May 3 to 25, 1897 (No.
4199).

Mikauia cordifolia Willd.

Keported from Central and South America. May 3 to 25, 1897 (No.
4299).

Conyza lyrata H. B. K.

lieported from Mexico, Central and South America. May 3 to 25
(Nos. 4290 and 4312).

Baccharis glutinosa Pers.

A common Mexican and Central American plant. May 3 to 25, 1897
(No. 4291).

Pluchea odorata Cass. •;

Widely distributed in Mexico and South America. May 3 to 25,
1897 (No. 4181).

Partheniuni hysterophorus L.

Common in Mexico, South America, and in the southern United
States. May 3 to 25, 1897 (No. 42G7).
Perityle microglossa Benth.

A common Mexican plant. May 3 to 25, 1897 (No. 4266).
Porophyllum nummularium DC.

Restricted to Mexico. May 3 to 25, 1897 (No. 4292).
Trixis frutescens P. Browu.

A common Mexican and Central American plant. May 3 to 25, 1897
(Nos. 4191), and Maria Cleofa Island, May 30, 1897 (No. 4331).
Jacquinia niacrocarpa Cav.

Species not represented in the National Herbarium, but reported
from Mexico, and Central and South America. May 3 to 25, 1897 (No.
4208).
Goiiolobus sp.

Fruit only. May 3 to 25, 1897 (No. 4313a).
Buddleia verticillata H. B. K.

A common Mexican species. May 3 to 25, 1897 (No. 4183)*

PLANTS OF THE TRES MARIAS ISLANDS. 85

Cordia sonorae Rose.

A recently described, species from Sonora. May 3 to 25, 1897 (No.
4207).

Cordia insularis Greenman.

Cordia insularis Greenman, Proc. Amer. Acad. 33 : 483. 1898.

The original description,is as follows : "Shrub 3 to 5.5 m. high ; stems
and branches glabrous, reddish brown, conspicuously dotted with
numerous whitish lenticels; the extreme branchlets covered with hir-
sute pubescence; leaves scattered, elliptic ovate or sometimes slightly
obovate, 1.5 to 3 cm. long, 1 to 1.5 cm. broad, narrowed below into a
short petiole, obtuse, the upper portion more or less deeply crenate-
dentate, occasionally sharply toothed, entire toward the base, hispid
above, spreading hirsute-pubescent beneath, especially on the midrib
and veins; inflorescence capitulate; heads small (after the corolla has
fallen, about 5 mm. in diameter) ; j)eduncles, during anthesis, 1 cm. or
less in length, covered witb a spreading hirsute pubescence; calyx 2
mm. long, 5-dentate; teetli short, acute; corolla 3 mm. long, nearly
cylindrical, with short recurved lobes, externally glabrous, pubescent
inside along the line of the filaments, stamens included; style a little
exserted. Collected by E. W. Nelson on Maria Madre Island of the
Tres Marias group of islands. May 3 to 25, 1897 (No. 4296)."

Tournefortia Candida Walp.

Not previously in herbarium. May 3 to 25, 1897 (Nos. 4217 and
4229).

Tournefortia cymosa L.
I have only seen specimens from Guatemala. May 3 to 25, 1897

(No. 4189).

Tournefortia velutina H. B. K.

Eeported from the west coast of Mexico and Guatemala. May 3 to
25, 1897 (No. 4209).

Heliotropium indicum L.

Common in Mexico and most tropical countries. May 3 to 25, 1897
(No. 4253).

Heliotropium curassavicum L.

Common in Mexico and South America as well as in the Old World.
Keported in the United States as far north as Oregon and Virginia.
May 3 to 25, 1897 (No. 4313).

Ipomoea bona-nox L.
A common tropical plant extending into Florida. May 3 to 25, 1897

(No. 42G9).

Ipomoea peduncularis Bertol.

Common in Mexico and Central America. May 3 to 25, 1897 (No,
4235).

86 T^ORTH AMERICAN FAUNA.

Jacqiiemontia violacea ( "boiay.

Eeported from Mexico, Central and South America, and the West
Indies. May 3 to 25, 1897 (No. 4251).

Solanum uigrum L.

A widely distributed species. May 3 to 25, 1897 (No. 42(m).
Solanum lanceaefolmni Jacq.

A common tropical plant. May 3 to 25, 1897 (No. 4240).
Solamim callicarpaefoliuni Kiiutli A: Bouchd.

Common in south Mexico and northern South America. Maria
Magdalena Island, May 26 to 28, 1897 (No. 4322).

Solanum torvum Swartz.

Common in Mexico and Central America. May 3 to 25, 1897 (No.
4185).

Solanum verbascifolium L.

Only reported hitherto from one station in southern Mexico. May
3 to 25, 1897 (No. 4216).

Physalis pubescens L.

A common tropical plant. May 3 to 25, 1897 (No. 4255).
Bassovia donnell-smithii Coulter.

A recently described South American and Cuatemalan species. May
3 to 25, 1897 (No. 4232).

Datura discolor Bernb.

Reported from Mexico and West Indies. May 3 to 25, 1897 (No.
4197).

Nicotiana trigonophylla Dnn.

Common in IMexico. May 3 to 25, 1897 (No. 4212).
Russelia sarmentosa Jacq.

A common Mexican and Central American siiecies. (May 3 to 25,

1897 (No. 4289).

Capraria biflora L.

A widely distributed plant, extending into Florida. May 3 to 25,
1897 (No. 4195).

Bignonia aequinoctialis L. (li. sarmen1o»a Bertol. )

Recently collected at Acapulco by Dr. Edward Palmer. It is com-
mon in Central and South America. May 3 to 25, 1897 (No. 4301), and
Maria Magdalena Island, May 26 to 28 (No. 4324).

Beloperone nelsoni Greenman.

Beloperone nehoni Greenman, Proc. Amer. Acad. 33 : 488. 1898.

" It is nearest B. comosa Nees, in DC. Prodr. 11 : 416, but differs very
markedly in the size of the flower and the character of the lower lip." —
Greenman in litt. May 3 to 25, 1897 (No. 4246).

The original description is as follows: "Erect; stems branching, sub-

PLANTS OF TITE TREft MARIAS ISLANDS. 87

terete, covered with a spreading or slightly retlexed grayish pu-
bescence; leaves ovate-lanceolate or obloug'-lanceolate, 5 to 10 cm. long,
2 to 4 cm. broad, obtuse at the apex, entire, narrowed below into a
slender i)etiole, densely lineolate above, pubescent on either surface,
especially on the veins, later becoming glabrous; petioles about -5 cm.
in length; inflorescence terminatin.; the stem and branches in rather
close bracteate spikes; bracts oblong or obovate; bracteoles linear,
nearly 1 em. long, exceeding the calyx ; calyx about 5 mm. long, deeply
5-parted; divisions nearly equal, lanceolate, acute, ciliate; corolla 2 to
2..") cm. long; tube exceeding the limb; upper lip shortly 2-lobed, the
lower more deeply 3 lobed, rather broad, somewhat plaited in the
throat; capsule 10 to 12 mm. long, pubescent. Collected by E, W. I^el-
son on Maria Madre Island of the Tres Marias grou]) of islands, 3-25
May, 1897 (Xo. 424G).

"A species closely resembling B. comosa Nees, but with a much
shorter corolla, and broader lower lip. The leaves are also somewhat
larger, longer-petioled, and much less pubescent. It may be that fur-
ther material will prove tliis to be a variety of B. comosa Nees, but as
the material at hand shows no sign of intergradation, it seems best for
the present at least to regard Mr. Nelson's plant as a distinct species."

Lantana horrida H. B. K.

IJeported from Ijoth northern and southern Mexico. Ma.>- 3 to 25,
1897 (No. 4187).

Citharexylum affinis D. Don.

This is a rare Mexican species which has been "compared witli the
Prodromus specimen at Geneva by C3. De Candolle'' — J. M. (r. May 3
to 25, 1897 (No. 4311).

.fflgiphila pacifica Greenman.
^l^giphila pacifica Greenman, Pioc. Amer. Acad. 3.3:485. 1898.

The original description is as follows : " Shrub 2.5 to 7 m. high ; stems
and branches terete, covered with a grayish brown bark and dotted
here and therewith lenticels, glabrous; branchlets terete, somewhat
compressed at the nodes, fulvous-pubescent; leaves opposite, oblong-
ovate, 5 to 15 cm. long, 3.5 to 7.5 cm. broad, more or less acuminate,
entire, rounded or rather abruptly narrowed at the slightly unequal
base, glabrous, or at least glabrate above, with scattered, tawny, sub-
appressed hairs beneath, especially upon the midrib and veins; petioles-
less than 1 cm. in length; inflorescence terminating the stems and
branches in rather close paniculate cymes; peduncles, x>edicels, the
subulate bracts and calyx covered by a fulvous suba])pressed ijubes-
cence; calyx about 4 mm. long, 4-lobed; lobes broader than long,
submucronate, greenish; corolla tubular, 10 to 12 mm. long, glabrous;
tube somewhat ampliated above; lobes oblong-elliptic, about 4 mm.
long, obtuse; stamensequal or rarely unequal, exserted; filaments pubes-
cent below, glabrous above; drupe yellow, obovoid, 8 to 10 mm. long, G

88 NORTH AMERICAN FAUNA.

to 8 mm. in diameter, one-lialf or luore exserted from the persistent
coriaceous subcrenately lobed, cup-sLaped calyx.— Collected by E. W.
Nelson on Maria Madre Island of the Tres Marias group of islands,
3-25 May, 1897, No. 4245 (in flower) and No. 4254 (in fruit)."

Hyptis albida H. B. K.

Several times reported from Mexico. May 3 to 25, 1897 (No. 4223).

Salvia aliena Greene.
A Mexican species. May 3 to 25, 1897 (No. 4247).

Stachys coccinea Jacq.

Common in Mexico and Central America, extending into Texas and
Arizona. May 3 to 25, 1897 (No. 42G5).
Iresine interrupta Bentb.

Reported from western and central Mexico. May 3 to 25, 1897 (No.
4234).
Phytolacca octandra L.

May 3 to 25, 1897 (No. 4293).
Stegnosperma halimifolia Beutli.

Common along the west coast of Mexico. May 3 to 25, 1897 (No. 4184).

Batis maritima L.

Extending from Florida and California to Brazil and the West Indies
and also reported from the Sandwich Islands. Magdalena Island, May
20 to 28, 1897 (No. 4327).

Coccoloba leptostachya Benth.

This species has not been heretofore found in Mexico, but has been
reported from Central America and South America. Maria Magdalena
Island, May 26 to 28, 1897 (No. 4315).

Antigonon leptopus Hook. & Am.

A very common vine on the west coast of Mexico. May 3 to 25, 1897
(No. 4204).
Aristolochia pardina Ducli.

A little-known i^lant collected at Colima many years ago by Ghies-
brecht, and recently at the same place by Dr. Edward Palmer. May
3 to 25, 1897 (No. 4304).

Piper aduncum L.

Reported from Mexico, Central and South America, and the West
Indies. May 3 to 25, 1897 (No. 4283).

Euphorbia sp.

May 3 to 25, 1897 (No. 42G8).

Euphorbia subcaerulea tresmariae Millspangh, var. nov.

"In the characters present in the specimens collected, this agrees well
with U.suhcacruleaHoh. and Greenm. (Pringle No. G265, Oaxaca), except
in the hairy involucre more regularly toothed involucral lobes, and in

PLANTS OF THE TliES MARIAS ISLANDS.

89

buviuf? the styles bifurcate, to the middle only, and flat spreading with
no tendency to reflexion or peltation as in the other species. The
fruits may prove this to be a distinct species. May 3 to 25, 1897 (Xos.
4298 and 4202)."— Mill spaugh MSS.

Euphorbia sp.

Specimens are indeterminable from lack of characters.
1897 (No. 4215).

Euphorbia nelsoni Millspaugh.

Eitphorhia velsoni Millspaugh, Bot. Gaz.26:2fi8. 1898.
May 3 to 25, 1897 (No. 4294, not 4284, as published).

May 3 to 25,

Fig. 2.—Eupliorl/ia nelsoni.

The original description is as follows: " Fruticosa, glabra, longe et
corymbosa ramosa, ramis teretis, iuternodiis longis, cortex maculatis,
maculic oblongis roseus. Foliis iuferioris fasciculatis, petioliis longis
filamentosis, pagina tenuis ovato-cuneatis, obtusis, apiculatis, foliis
floralibus oppositis, orbiculatis petiolis limbum sequantis. Involucriis
terminalibus corymbosis, pedunculatis, campauulatis glabris, lobis latis
truncatis irregulariter G-8 fimbriatis, glandulis 5, transversis oblongis
integris, appendicibus minutis vel nullus. Stylis longis revoluto-cir-
cinalis. Capsuhe luridjc profunde tri-sulcatoe, semine sub-globosis
pallide-fuscis, scrobiculatis, linea media nigra geminatis, rugjc anasto-
mosantis tuberculatis 2 mm. long, 1.9 mm. lat."

Several Euphorbias were collected on the islands in too imperfect
condition to determine, and it has been thought advisable to reproduce
the cut^ of the present species for the purpose of assisting future study
of the iiora.

' Through the kindness of the editors of the Botanical Gazette I am permitted to
use this illustration.

90 NORTH AMERICAN FAUNA.

The main figure shows a cluster of leaves. To the right is a flower
cluster and to the left a dissected flower with end and side views of the
seed.

Garcia nutans Rohr.

Found in Mexico and South America. May '^ to 25, 1807 (No. 422S).
Croton ciliato-glandulosus Ort.

May 3 to 25, 1897 (No. 4218).
Acalypha sp.

May 3 to 25, 1897 (No. 4200).
Celtis monoica Ilenisley.

May 3 to 25, 1897 (No. 423G).

Buxus pubescens Greenman.

Jlnxus piiheseenn Greenman, Proc. Anier. Acad. .S3 : 481. 1898.

The original description is as follows: " Shrub or small tree, 4.5 to
8 m. high; stems and branches covered with a grayish bark; the
branchlets and younger shoots provided with a soft, spreading pubes-
cence; leaves opposite or occasionally subalteruate, sessile or nearly so,
rhombic-ovate to oblong-ovate, 2 to 5 cm. long, 1.5 to nearly 3 cm.
broad, 3-nerved, obtuse or acntish, mucronate, cuneate at the base,
entire, ciliate, soft-pubescent beneath, more sparingly pubescent and
glabrate above, showing the reticulate venation on the upper surface;
inflorescence of axillary short-pedunculate much contracted subrace-
mose pubescent clusters; staminate flowers pedicellate; pedicels 3 mm.
long, about twice exceeding the ovate acute bracts; calyx deeply 4-
parted; divisions ovate, acute, 2 mm. long, the inner divisions slightly
broader than the outer ones; the rudimentary pistil somewhat quatre-
foil or X-shaped; fertile flowers about 5 mm. long, single, sessile,
terminating the inflorescence; ovary glabrous ; fruit not seen. — Col-
lected on Maria Madre Island by E. W. Nelson, 3-25 May, 1897,
No. 4221.

"A species apparently endemic in the Tres Marias Islands, and most
nearly related to the West Indian B. j)ulcliclla Baill."

Ficus radulina Watson.

A recent species of Dr. Watson's from northern Mexico. May 3 to
25, 1897 (No. 42G1).

Picus fasciculata Watson.

Only known from western Mexico. May 3 to 25, 1897 (No. 4288).
Ficus sp.

May 3 to 25, 1897 (No. 4182).
Myriocarpa longipes Liebra.

Found in Mexico and Central America. May 3 to 25, 1897 (No. 4275).
Agave sp.

Six meters higli, leaves 9 to 18 dm. long; marginal teeth small, dis-
tant; end spine short, stout, i)ungent; capsules oblong, large, 7 cm.
long.

PLANTS OF THE TRES MARIAS ISLANDS. 91

This species belongs to the subgenus Eiigave and the Rigidae group
of ]Mr. Baker's revision. It is near A. vivqjara, and perhaps not dis-
tinct. Mr. iS"elson's plant does not seem to differ from specimens col-
lected by me on the mainland. May 3 to 25, 1897 (No. 4204).

Cyperus ligularis L.

Eeported from Mexico, Central and South America, and West Indies,
as well as Africa and Australia. Maria Cleofa Inland, May 30, 1897
(No. 4330).
Cyperus incompletus Link.

Eeported from Mexico and Brazil. May 3 to 25, 1897 (No. 4259).

Panicum brevifolium L.

May 3 to 25 (No. 4257).
Eleusine indica Gaertn.

May 3 to 25, 1897 (No. 4305).
Dactyloctenium aegyptiacum Willd.

May 3 to 25, 1897 (Nos. 4295 and 4256); Maria Magdalena Island,
May 2G to 28 (No. 4317).

Arundo donax L.

Maria Cleofa Island, May 30 (No. 4332).
Zamia loddigesii ( ?) Miq.

Reported from Mexico. Maria Cleofa Island, May 30, 1897 (No. 4329).

Pteiis longifolia L.

Maria Madre Island, May 3 to 25, 1897 (No. 4201).
Aspidium trifoliatum Swartz.

Maria Madre Island, May 3 to 25, 1897 (No. 4280).
Aspidium patens Swartz.

A widely distributed species. Maria Magdalena Island, May 20 and
28, 1897 (No. 4316). .
Adiantum concinuuni H. B. K.

Maria Madre Island, May 3 to 25, 1397 (No. 4273).
Adiantum tenerum Swartz.

Maria Madre Island, May 3 to 25, 1897 (No. 4281).
Gymnogramme calomelanos Kaiilr.

A widely distributed species. Maria Cleofa Island, May 30, 1897
(No. 4333).

PARTIAL BIBLIOGRAPHY OF THE TRES MARIAS ISLANDS.

By E. W. Nelson.

1703. Dampiek, William. A New Voyage round the World, 5th cd., I, pp. 26b-264.
Notes the presence of seals abont the Tres Marias. These notes are quoted
l)y Allen and Alston.

1865. Baird, Spencer F. <^Revievv of American Birds, p. 232. Description of

Granatcllun fra7icesca'.

1866. Allen, Harrison. Notes on the T'esjKT<(7J0Hirfa' of Tropical America. <^Proc.

Acad. Nat. Sci. Phila. 1866, p. 285. Description of Bhoijirssa parvida.

1867. Cassin, John. A third study of the Icteridie. <^Proc. Acad. Nat. Sci. Phila.

1867, p. 48. Description of Icterus fjraysoni.
1867. Lawrence, George N. Descriptions of Six New Species of Birds of the

Families Hirmidinlda', Forniicarichv, Ti/rannidw, and Trovhilidw. <^Ann. Lye.

Nat. Hist., N. Y., VIII, pp. 404-405. Description o{ Amazilia graysoni.
1871. Grayson, Andrew Jackson. On the Physical Geography and Natural His-

torj' of the Islands of the Tres Marias and Socorro off the Western Coast of

Mexico. Edited by Geo. N. Lawi'euce. <Proc. Bost. Soc. Nat. Hist., XIV,

pp. 261-302. In addition to Grayson's notes, Lawrence gives a few remarks

on certain birds said to have been taken on the Tres Marias by Xantus and

describes Pyrrhophana graysoni {^Amazilia graysoni) and Sterna fuliginosa

crissalis (ex Baird MSS.).
1871. Lawrence, George N. Descriptions of New Species of Birds from Mexico,

Central America, and South America, with a note on liallus longirostris.

<Ann. Lye. Nat. Hist. N, Y., X, pp. 1-21. Description of Parula insularis

{=^CompsotMypis insularis).
1874. Baird, Spencer F. <[History of North American Birds, II, pp. 515, 516.

Description of Dryobates svalaris graysoni.
1874. Lawrence, George N. Birds of Western and Northwestern Mexico. <Mcm.

Bost. Soc. Nat. Hist. II, pp. 265-319. Contains extracts from Grayson's untes

on various species of birds of the Tres Marias.

1876. Wallace, Alfred Russell. Geographical Distribution of Animals, II, pp.

59-60. Summary of the fauna of the Tres Marias, comprising 52 species of
birds, 3 mammals, and several species of snakes and lizards.

1877. Allen, Joel Asapii. <Mou. N. Am. Rodentia, 347-348. Description of Lepns

graysoni.
1877-79. Grayson, Andrew Jackson. Historia Natural de las Islas de las Tres
Marias y Socorro. <LaNaturalezaIV, pp. 159, 203, and 252. A Spanish
translation, by Senor Don Aniceto Moreno, of Colonel Grayson's pai»er
originally published in the Annals of the Lyceum of Natural History,
New York, X, 1871.

1878. Ridgway, Robert. Description of a New Wren from the Tres Marias Islands.

<Bull. Nutt. Orn. Club, III, p. 10. TItryothorus felix lawrencii {= T. lawrencii).
1879-82. Alston, Edward R. <Biologia Ceutrali-Americana. Mammalia. 1879-
1882. Notes on Veaperttgo parvulus {=:Bliogecsa parvula), p. 21, Lepus gray-
soni, p. 177, and a seal, p. 210, on the Tres Marias.

93

94 NORTH AMERICAN FAUNA.

1879-98. Salvin, Osbert, and Godman, F. DuCane. <^Bi()logia Centrali-Americana'
Aves I aud II, 1879-1898. Coiitain.s notes on various species of birds of
till' Tres Marias, based on the work of Grayson aud Forrer.
1880. Allex, Joel Asaph. History of North American Pinnipeds, 1880, p. 290. Misc.

publication No. 12, U. S. Geol. aud Geog. Survey Terr. Quotation of Dauijiier's

notes on seals about the Tres Marias.
1882. Riugway, Robert. Description of Several New Races of American Birds.

<;Proc. U. S. Nat. Mus., V, p. 12, 1882. Description of Merula flavirostris

graysoni.
1882. Ste.jneger, Leonhard. Description of TviTo New Races of Myadestes ohncarm

Lafr. <Proc. U. S. Nat. Mus., IV, p. 373, 1882. Description of Myadestes

obscurus insularis.
1882. Thomas, Oldfield. Biologia Ceutrali-Americaua, Mammalia, Supplement,

1882. Notes on various species of mammals found on the Tres Marias:

VespertKjo parindn>i, \\. 203; Atalapha noreburaceusiti, p. 205; Vespertilio niyri-

t'«HS, p. 206; Macrotus waterhoiisii, p. 207; Charonycteris mexivaiia l=^(llosi>o-

phaija miitica^, p. 207; I'rocyon cancrivoriis, p. 208; Lepiis (/raysoni, p. 211.
1885. Madauasz, Julius von. Oinithologiai Kozlem6nyek A Magyar Nemzeti

Muzeum Gylijtemeuyebol. <^ Term^szetrajzi Fiizetek, IX, p. 74, Feb. 20, 1885.

Description of Vireo forreri.
1887. RiDGWAY, Robert. <^ Manual of North American Birds, 1887. Descriptions

of laclie lawrencei (ex Berlepsch MS.), p. 320; Platypsaris insularis, p. 325;

Pirawja Jlaminea, p. 457.
1887. RiDGWAY, Robert. A Review of the Genus Fsittacula of Brisson. <[Proc.

U. S. Nat. Museum, X, 541, 1887. Description of Psif taenia insulai'is.
1891. Bryant, Walter E. Andrew Jackson Grayson. <^ Zoe, II, pp. 34-68, 1891.

A short account of Grayson's life, with extracts from his journals on the

habits of certain birds of the Tres Marias.
1898. Evermann, Barton Warren. Notes on Fishes Collected by E. W. Nelson on

the Tres Marias Islands and in Siualoa and Jalisco, Mexico. <^ Proc. Biol.

Soc, Washington, XII, pp. 1-3, 1898. Records Agonostomus nasuftts Giinth.

on Maria Magdalena and Maria Cleofa.
1898. Merriam, C. Hart. Mammals of Tres Marias Islands ott" Western Mexico.

<Proc. Biol. Soc. Washington, XII, pp. 13-19, 1898. Descriptions of

Marmosa insularis, Ory^omys nelsoni, Peromyscus madrcnsis, Procyon loior

insularis, Glossophaga mutica, with notes on other species of mammals

occurring on the islands.
1898. Nelson, Edward William. Descriptions of New Birds from the Tres Marias

Islands, Western Mexico. <Proc. Biol. Soc. Washington, XII, pp. 5-11, 1898.

Descriptions of the following new species and subspecies: Columha ftariros-

tris tnadrensis, Leptotila capitalis, Butco borealla fumosus, Polyborus cheriiniy

pallidas, Trogon antbignus goldmani, Nyctidromus albicollis insularis, Myio-

pagis placens minimus, Cardinalis cardinalis maria', Vireo hypochryseus sordidus,

Melanotis cwrulescens longirostris, Thr'yothorus lawrencii magdalenw.
1898. Greenman, Jesse M. Diagnoses of New and Critical Mexican Phanerogams.

<Proc. Am. Acad. Arts and Sci., XXXIII, No. 25, pp. 471-489, June, 1898.

-^Descriptions of Jigiphila pacifica, Beloperone nelsoni, Buxua pubescens, and

Cordia insularis.
1898. MiLLSPAUGH, Charles F. Notes and New Species of the Genus Euphorbi.').

<^Botanical Gazette, XXVI, pp. 265-270, Oct., 1898. Descriptions of Eupliurbia

nelsoni and Eupltorbia subcccrulea tresmariw.

I]^DEX.

[Names of now species in black-facii type.]

Abutilon reventum, 79.
Acacia, S''.
Acalyiiha, 90.
Actilis iiiacularia, 34.
A<liaiituiii tcnernm, 91.
32f:ialiti3 Sfiiiiiiahnata, 34.
j:>liphila pacitica, 13, 78, 87-88.
Agave, 9, 12, 90-91.
Agkistrodon biliiieatus, 71.
Ajr<iiii>stuiiius uasutus, 11.
Albi/.zia oicitlentalis. 83.
Amazilia ciiiuanioiuea, 45-46.

gray.soni, 12, 22, 45, 40.
Amazoiia finschi, 41.

oratrix, 39-41.
Aniyris, 80.
Auoli.s iieVmlo.sus, 65.
Auous stolidiis, 26.

stoliduM lidgwayi, 26-27.
Antigoiiou lc])t(i]>us, 88.
Aretuccplialus towiiiiendi, 18.
ArdL-a laiididissima, 33.

egret t a, 33.

liei'odiaa, 33.
Argeinono oeliroleuca, 78.
Avistolochia ])ardina, 88.
Anindo donax, 91.
A.'iliidimii patens, 91.

trit'oliatnm, 91.
Astragalinu.s psaltria mexicauus, 52.
Ateleia, 82.
Awaous, 11.
Baccbaris glntinosa, 84.
Bascanion lineatuni, 70.
Bassovia ddnnell-siuitbii, 86.
Bat, Big-ea,red, 18.

^laximilian's Black, 15, 18.

Mexican Ked, 19.
Bati.s maritima, 88.
Banliinia. 83.
Beloperone como.sa, 86, 87.

nelsoni, 13, 78, 86-87.
Biguonia a'quinoctialis, 86.

saruiento.sa, 86.
Bitbynisjaniaiceusis, 74-75.

vollenliiivenii, 75.
Boa lni])erati»r, 63, 69.
Booby, I?]u(-t'ooted, 31-32.

J?rewster'.s, 29,30.

AVebsters, 29.
Bracbyrlianipus breviro.stris, 23.

bypoleucus, 23.
Budd'leia verticillata, 84.
Biirseia gunimifera, 80.
Buteo ))()rea!i3 ealurus, 37.

boiealis fnniosns, 12, 37-38.

bmealis nioutana, 37.

borealis socorroeusis, 37-38.
r.uxus ]puliescens, 13, 78, 90.
Caiiiix'iiliiliis, 23.
Caiiavalia gladiata, 82.
Cajiparis bieynia, 78.

cyiiapliallopbora, 78.
Capj-aria IdHoia, 86.
Caracara, Trea Marias, 38-39.
Cardinalia cardinalis mariw, 12, 22, 52.

virginiaiius, 52.

virgininnus igneus, 52.
Cardin;d, Tres Marias, 52.
Caidiosperuium coriudum, 81,

Ca.searia, 83.

corymbosa, 83.

.sylvestris, 83.
Cassia, 12.

atoinaria, 82.

bitlora, 82.

emarginata, 82.
Catbartes aura, 37.
Celtis monoica, 90.
Cereiia, 12,51.
Cerylo alcyon, 43.
Chloriistili)on insularis, 62.

puclicraiii, 62.
Cba'ionycteris niexicana, 19.
Chordeiles actijieunis texensis, 45.
Cbrysotis levaillantii, 39.
Ciccaba squamulata, 39.
Circe latirostris, 46.
Cissampelos pareira, 78.
Cissolopba, 23.

beecheyi, 50.
Cissus sicyoides, 80.
Citharexylum atiinis, 87.
Cnemidophorus gularis mexicanns, 63, 68.

niariarum, 12, 03, 67-68.
Coccololia leptoslachya, 88.
Coccyzus minor, 42.
Colubriua arborea, 80.
Columba llavirostris, 35.

flavirostris madrensis, 12, 22, 35.
Columbigallina passerina pallescens, 37.
Compsotblypis inornata, 55.

iusnlaris, 11, 12, 22, 55-56.

nigrilora, 55.

pitiayumi, 55.

pnlcbra, 55, 56.
Conocarpus erectus, 83.
Coutopiis ricbardsoni, 49.
Conurns, 23.
Conyza lyrata, 84.
Coriiia insularis, 13, 78, 85.

sonorie, 85.
Cormorant, 32.
Corvu.s niexicanus, 50.
Craticva tapia, 78.
Crocodile, 11.

Crocodylus americanus, 64.
Crotalus, 71.
Crotolaria lupuliua, 81.
Crotou ciliato-glandulosus, 90.
Crow, Mexican, 50.
Ctenosaura teres, 65-66.
Cuckoo, Mangrove, 42.
Cyanos])iza, 23.
Cypenis iucompletus, 91.

lig(ilaris,91.
Dactylocteniuni »gyi)tiacum, 91.
Datura discolor, 86.
Dendroica a-stiva rubiginosa, 56.

a'stiva morcomi, 56.

anduboni, 56.

townsendii, 56.
Desmodium, 81.
Diplotrojiis diplotroi)is, 63, 69.
Bove, Mexican Ground, 37.

Mourning, 36.

Tres Marias, 36.

Wbite fronted, 36.

Wbitewinged, 36-37.
Drymarclion corals melauurus, 70

9G

INDEX.

Dryniobiiis hoddji'rti, 69-70.
Dryobates scalaris, 43.
" scalaris bairdi, 43.
scalaris graysoni, 12, 2"i, 43-44.
scalari.s lucasauiis, 43.
scalaris sinaloensis, 43-44.
Egret, American, 33.
Elaiuea placena, ^0.
Eleunine jntlica, 91.
Enipidonax ditticilis, 49.
Erythrina lanata, 13, 78, 81
Eiipatoriiiin, 84.
coUinuni, 84.
Euphorbia, 12,88.8!).
nelsoni, 13, 78, 89-90.
subcoerulea t^e.>^ma^iEB, 13, 78, 88-80.
Falco albigiilaris, 38.
coliiiuljaiius, 38.
peregrinus anatuni, 38.
peregrinus iiigi'iceps, 38.
sparvcrius, 38.
Falcon, A\"hite-tliroated, 38.
f icus fasciculata, 90.

radulina, 90.
Elorisuga niellivora, 62.
Flycatcber, Arizona Crested, 48.
Beardless, 49.
Golden Crowned, 50.
Little Golden Crowned, 50.
Olivaceous. 48-49.
Western, 49.
Eregata aquila, 33.
Garcia nutans, 90.
Gecarcinus diguetl, 73-74.
Gilibertia in.sularis, 13, 78, 83-84.
Glandina turria, 11.
Glossopliaga niutica, 11, 18-19.
Glossophaga, Tres Marias, 18-19.
Goldtin<li, Mexican, 52.
Gouolobus, 84.
Crackle, Great-tailed, 52.
Granatellus francescas 12, 22, 56-57.

venustus, 57.
Grapsus grap.sus, 74.
Guaiacuui coulteri, 79.
Guarea, 80.

Guazuma iilniifolia, 79.
Gull, American herring, 23.

Heermann'.s, 23-24.
Gymnogrammo calouielanos, 91.
Hadrostomus aglaia- attiuis, 47.
Hiematopus frazari 34-35.
galapagensis, 34-35.
palliatus, 34-35.
Haliplana fiiliginosa crissalis, 24.
Hawk. Duck, 38.
Pigeon, 38.
Sparrow, 38.

Tres Marias Ked-tailed, 37, 38.
Heliotriipluni curas.savicum, 85.

indicum, 85.
Heron, Great Blue, 33.
Snowy, 33.

YeHow-crowned Night, 33, 34.
Hetei'oiitciys tloribunda, 79.
Hibiscus tiliaceus, 79.
Hippocratia, 80.
Hiruudo ei-ythrogastor, 54.
Humming IJird, Grayson's, 45, 46.

Lawrence's, 40. 47.
Hyloclchla ustul.ata, 60.
ustulata alma;, 60.
ustulata swain.soni, 60.
Hypotriorchis rufigularis, 38.
Hyptis albida, 88.
laclie latirostris, 46, 47.
lawrencei, 12, 22, 46.
Icterus graysoni, 12, 22, 50-51.

pustulatus, 51.
Iguana, Black, 65, 66.
Ipomcea, 12.

bona-nox, 85.
jieduncularis, 85.
Iresine interrupta, 88.
Jacquemontia violacea, 86.
Jacquiuia macrocarpa, 84.
Jay, Beechey's, 50.
Kingbird, Couch's, 48.
Kiuglisher, Belted, 43.

Kinostornon integrum, 64.

LaiiirlluNis, 11.

Laiiiproiicltis micropholis oligozona, 70.

Lautana liurrida, 87.

Larus argentatus smithsonianus, 23.

heermanni, 23-24.
Lasiurus borealis niexicana, 15, 19.
Leptopbis <li]>liitropis, 69.
LeptotilaiiU)ifn.ns,36.
capitalis, I'J, 22, 36.
fulviventris brachyptera, 36.
Lepus graysoni, 11, 16-17.
Lonchocar])us, 82.
Lovebird, Tres Marias, 41-42.
Man-o'war bird, 33.
Marraosa insularia, 11, 15-16.
Melanerpes, 23.
Melanotis cairulescens, 59.

Cicrulescens longirostris, 12, 22, 59.
hypoleucus, 59.
Melochia tomentosa, 79.
Melopelia leucoptera, 22, 36-37.
Merula llavirostria, 60, 61.
grayi, 62.

graysoni, 12, 22, 60-62.
tristis, 62.
Micropallas, 39.
Mikania cordifolia, 84.
Mimus polyglottos, 57.
Mockingbird, 57.
Momotus, 23.
Mouse, Tres Marias, 16.
Murrelet, Short-billed, 23.
Mus rattus, 10.
Myadestes obscurus, 59.

obscurus inaularis, 12, 22, 59-60.
obscurus occidentalis, 60.
Myiarchus lawrencei olivascens, 48-49.

mexicanus magiater, 48.
Myioi)agis placens, 50.

placena minimus, 12, 22, 50.
Myiozetetes, 23.
Myotis nigricans, 15, 18.
Myriocarpa longipes, 90.
Nicotiana trigonophylla, 86.
Nighthawk, Texas, 45.
Nycticorax vicdaceus, 33-34.
Nyctidromus albicollia, 44.

albicollis insnlaris, 12, 22, 44.
albicollis merrilli, 44.
Oceanodroma melania, 28.
Ochna, 80.
Ocypodo, 74.
Ocypode kuhlii, 74.
' occidentalis, 74.
Opeas subula, 11.

Opossum, Tres Marias Pigmy, 11, 15-16.
Opuntia, 12, 83.
Oriole, Grayson's, 50.
Ornitliion iraberbe, 22, 49.
iniberbe ridgwayi, 49.
Orthalicus undatus, 11.

undatus melauocheilus, 11.
Oryzomya nelsoni, 11, 16.
Osprey, 39.

Otopterus mexicanus, 18.
Owl, Barn, 39.

Burrowing, 39.
Oxybelis acuminatus, 69.
Oyster-catcher, American, 34-35.
Piindion halia^tus carolineusia, 39.
Panicum brevifolium,91.
Parauque, Tres Marias, 44.
Parrot, Double yellow-bead, 39-41.
Partheniuin hysterophorus, 84.
Parula, insnlaris, 55.
Tres Marias, 55.
Passillora, 12, 83.
Paullinia seasiliflora, 81.
Pclecanus californicus, 32-33.
Pelican, California Brown, 32-33.
Petasophora thalassina, 62.
Perityle microglossa, 84.
Peromyscus madrensls, 11, 16.
Petrel,' Black, 28.
Plucthon athereus, 28-29.
Phalacrocorax, 32.
Phaseolu?, 82.
Phocicua commuuis, 19.

INDEX.

97

PhyllmLactylus tiiberculosus, 63, 64-65.
I'liysalia jiubescens, 86.
Phytolacca octaudra, 88.
Piaya, 23.
Picramnia, 80.

eiliata, 80.
Pious scalaria graysoni, 43.
Pigeon, Tivs Marias, 35.
Pilocarpus insularis, 13, 78, feO.
Piper aduucuiu, 88.
Pipilo,23.
Pirauga bidentata, 53, 54.

bitlentata flammea, 12, 22, 52, 53, 54.

ludoviciana, 23, 52.
Pithecolobium, 12.

dvilce, 39, 77, 83.

ligustrinuEi, 83.
Platypsaris aglaia;, 47-48.

aglaiffi albiventris, 47, 48.

aglaiffi insularis, 12, 22,47-4.'^.

aglaiiB sumiclirasti, 47-48.
Plover, Semipalmated, 34.
Pluchea odorata, 84.
Polyliorns audubonii, 38.

audubonii insularis, 22.

cheriway pallidus, 12,22,38-39.
Polygyra ventrosula, 11.
Porophyllum nuninmlariuni, 84.
Porpoise, Common, 10.

Long-no.sed, 19.
Portlaudia pterosperma, 84.
Procyon lotor insularis, 11, 17.
Prodelphinvis longirostris. 19. •
Psidiuni,35,83.
Psittacula cyanopyga, 41-42.

insularis, 12, 22, 41-42.
Pteris longifolia, 91.
Puffinus cuneatus, 27-28.

knudseni, 27.
Pyrgisoma, 23.
Pyrrhophiiena graysoni, 45.
Kaccoon, Tres Slarias, 17.
Rat, Black, 16.

Nelson's Rice, 16.
Ehogeessa parvula, 11, 18.
Rhogeessa, Tres Marias, 18.
Ehynchosia minima, 82.

precatoria, 82.
Russelia sarmeiitosa, 86.
Quiscalus macrourus, 52.
Saltator. 23.
Salvia aliena, 88.
Sandpiper, Spotted, 34.
SceloiJorus boulengeri, 67.

crarkii,67.

horridiis, 67.

olijioporus, 67.
Schcppna schreberi, 80.
.Sea Lion, 15, 17-18.
Serjauia mexicana, 80.
.Shearwater, Wedge-tailed, 27-28.
Solanuui, 12.

callicarpsefolium, 86.

lauceffifolium, 86.

nigrum. 86.

torvum, 86.

verbasciiolium, 86.
Solitaire. Tres Marias, 59-60.
Spanish cedar, 7, 8, 9, 12.
Speotyto cunicularia hypogiva, 39.
Stacbj's coccinea, 88.
Stegnosperma halimifolia, 88.
Sterna elegans, 24.

fuligiuosa crissalis, 24-26.

galerieulata, 24.

maxima, 24.
Stris pratincola, 39.
Sula ba.ssana, 29.

brewsteri, 29-30.

gossi, 31,32.

Sula nebouxii, 31-32.

piscator, 31.

websteri, 29.
Swallow, Barn, 54.
Tanager, Louisiana, 52.

Tres Marias, 52, 53.
Tephrosia, 81.
Tern, Elegant, 24.

Pacific iSToddy, 26-27.

Pacific Sootv' 24-26.

Royal. 24.
Ternostrcemia maltbya, 13,78.
Thalurauia glaucopis, 62.

lucia-, 02.
Thrush, 01i.ve backed, 60.

Russet-backed, 60.
Thryothorus felix, 22. 57, 58.

lawrencii, 12, 22, 57, 58.

lawrencii magdalena-, 12, 22, 58.
Tournefortia Candida, 85.

cymosa, 85.

velutina, 85.
Totanus tlavipes, 34.
Tres Marias, Blue Mockingbird, 59.

Caracara, 3S-39.

Cardinal. 52.

Cbat-AVarbler. 56-57.

Cottontail. 16-17.

Dove, 36.

Glossopbaga. 18-19.

Lovebird, 41-42.

Mouse, 10.

Parauque, 44.

Parula, 55-56.

Pigeon. 35.

Pigmy Opossum. 11, 15-16.

Raccoon, 17.

Red-tailed Hawk, 37-38.

Rbogeessa, 18.

Robin, 60-61.

Solitaire, 59-60.

Tanager, 52.

Vireo, 54-55.
Trichilia spondioides, 80.
Trixis frutescens. 84.
Trogon ambignus, 42.

ambiguus goldmani, 12, 22, 42-43.
Troiron, Goldman's, 42-43.
Tropic Bird, Red-billed, 28-29.
Turdus flavirostris, 60.
Turtle, Mud, 11.

Tyrannus melancholicus couclii. 22, 48.
Uranomitra guatemalensis, 62.
Urvillea ulmacea, 81.
Uta lateralis, 66-67.
Vireo flavoviridis forreri, 12, 21, 22, 54, 55.

hypochryseus, 54.

hypochryseus sordidns, 12, 22, 54-55.
Vireo, Forrei-'s, 54.

Tres Marias, 54-55.
Vulture, Turkey, 37.
Warbler, Alaskan Yellow, 56.

Audubon's, 56.

Pileolated, 57.

Townseud's, 56.

Tres Marias Chat-, 56, 57.

"Western Yellow, 56.
Wilsouia pusilla pileolata, 57.
Wissadula hirsutitlora, 79.
Wood Pewee, Western, 49.
Wren, Magdalena, 58.

Maria Madre, 57-58.
Ximenia americana, 80.
Yellow-legs, 34.
Zalophus californianus, 17-18.
Zamia loddigesii, 91.
Zanthoxylum insularis, 13, 78, 79.

nelsoni, 13, 78, 79.
Zenaidura macroura, 36.

139r)(>— No. 14-

U. S. DEPARTMENT OF AGRICULTURE
DlVISIOlN OF BIOLOGICAL SURVEY

NORTH AMERICAN FAUNA

ISTo. 15

[Actual date of iniblication, August 8, 1899]

REVISION OF THE JUMPING MICE OE THE GENOS ZAPOS

BY

EDWARD A. PREBLE
ASSISTANT IN BIOLOGICAL SURVEY

Pifjiared iiuder the direction of

Dr. C. HART MERRIAM
CHIEF OF DIVISION OF BIOLOGICAL SURVEY

- c-^^-^sx-'

WASHINGTON

GOVERNMENT PRINT INGr OFFICE
1809

':J^^ ':t" cAvrt^^^-

A>t^ «

LETTER OF TRANSMITTAL.

U. S. Depaetment of Agriculture,

Washington, T). C, June 15, 1899.
Sir : I liave the l)onor to traDsmit herewith for pubHcation, as Ko. 15
of ]S^orth American Fauna, a 'Eevision of the Jumping Mice of tlie
Genus Zapus.'' by Edward A. Preble, assistant in the Biological Survey.
Eespectfully,

T. S. Palmer,
Acting Chief, Biological ISicrrey.
Hon. James Wilson,

Secretary of Agriculture.
2

CONTENTS.

lutrodiictiou 5

History and material 5

Distiibutiou 6

External characters 7

Habits "

Nomenclature 9

Generic names ^

Specific and suLspecific names 10

Genus Zapus Cones 13

Key to subgenera 13

List of species and subspecies, with type localities 13

Subgenus Zapus Cones 1^

Key to species 1^

Xapa'o^apus subgenus novum 33

Key to species 33

Eozapus subgenus novum 37

ILLUSTRATIONS.

PLATE.

Page.

Plate I. Skulls of Zapus insignis, Z. setchhanus, Z. hudsonius, Z. orariiis,

Z. irinotattis, and Z. princeps -12

TEXT FIGURES.

Fig. 1. Molar teeth of Zapus h udsonius li

2. Molar teeth of Zapus insignis 33

3. Molar teeth of Zapus setcliuauus.i >"

4. Skull of Zapus setchuaitus 38

3

No. 15. NORTH AMERICAN FAUNA. August, 1899.

REVISION OF THE JUllPING MICE OF THE (lENOS ZAPllS.

By Edward A. Preble.
INTRODUCTION.

History and materiah — Tlie^Jumpiug Mice of Kortli America early-
attracted the attention of naturalists, but their true status and rela-
tionships long remained in obscurity. The first reference to any of the
species seems to have been made by Thomas Pennant, in the latter part
of the last century-, who mentioned the animal under the name of the
'Long-legged Mouse of Hudson's Bay.' In 1780 Zimmermann, basing
his description on that of Pennant, and supposing the animal to be con-
generic with the jerboas of the Old World, named it Blpus Imdmnius.
Subsequent authors, recognizing the inapplicability of Dipus, referred
the species successively to Gerbillus, Meriones, and Jaculus, until finally
Dr. Elliott Cones, in 1875, after showing that all the generic names
previously used were untenable, proposed Zapus.

Previous to 1857 a number of specific names were used for members
of this genus (see pp. 10-13), but were applied mainly to specimens from
the eastern United States and Canada. Suffice it to saj^ that Baird,
in 1857, and Cones, in 1877, recognized but one species, having a range
nearly coincident with the i)resent geographic distribution of the genus
in North America. This resulted mainly from two causes — poor qual-
ity of existing material and the prevailing teudenc}^ of the "times to
lump distinct species having a superficial resemblance to one another.
The material consisted mainly of poorly preserved alcoholic speci-
mens, or skins badly made (in most cases with the skulls inside),
and without satisfactory measurements. The characters distinguishing
the species, therefore, were not apparent or were considered as being-
due merely to individual variation. In failing to examine skulls, both
Baird and Cones referred specimens of the animal since described as

5

6 NORTH AMERICAN FAUNA.

Zapiis insi(inis to Z. hudsonius, though. iHsii/nis has a different number of
teeth and is now considered subgenerieally distinct from the hitter.
Under tlie improved methods of collecting and preserving mammals
now in vogue large series of well-prepared specimens have been brought
together during the ])ast few years, and several new forms have been
desciibed, mainly from the west and northwest.

The following revision of the genus Zapus is based on a study of more
than 900 specimens, mainly in the collections of the Biological Survey of
the U. S. Department of Agriculture, the V. S. Xational Museum, and
several x^rivate individuals.^ To the custodians and owners of these
collections my thanks are cordially extended. For kind assistance and
advice my thanks are also especially due to Dr. 0. Hart Merriam, Dr.
T. S. Palmer, and ]Mr. Gerrit S.Miller, jr. Through the courtesy of
Mr. Miller I have had an opportunity of examining the types of Zapus
setchuanus Pousargues, temporarily placed in his hands for comparison,
and am thus able to present a complete account of the genus, includ-
ing a description of this very interesting Asiatic species, the only oue
known from the Old World.

The Xational Museum collection of Zapus, though not extensive,
contains many si^ecimens of unusual interest from widely separated
localities; that of the Biological Survey is especially rich in westeru
specimens; and those of Miller and Bangs contain good series of east-
ern species. These collections, comprising many large suites of well-
prepared skins, nearly all accompanied by perfect skulls and by meas-
urements taken in the flesh, have afforded facilities for comparison never
before enjoyed. Much additional material is needed, however, before
certain questions concerning relationship and seasonal variation can
be satisfactorily settled.

Disfrihufion. — Zapus is a Boreal genus, and, with the exception of a
single species, is confined to Xortli America. The exact northern lim-
its of its range are not well known, but are indicated by specimens
from Hamilton Inlet, Labrador; Fort Churchill, on Hudson Bay; Fort
Eae, on Great Slave Lake; and the Yukon River (probabl}' Fort Yukon,
just below the Arctic Circle). Southward it extends as far as Ealeigli,
K C; Wheeling, W. Va.; Jarksou County, Mo.: Santa Fe, X. Mex.;
and Kern Eiver, California. Within the region thus roughly outlined
there are probably no large areas where at least one member of the
genus is not found. In regions where there is little or no woodland its
distribution is more or less local, and is confined almost exclusively to
the borders of streams. Of course, the arid deserts of the West do
not offer a conoenial environment.

•These specimens are as follows: Biological Survey, 378; U. S. National Museum,
85; Merriam Collection, 60; Bangs Collection, 132; Miller Coxlection, 200; and a few
from each of the following collections, American 3Iuseum of Natural History, New
York; Academy of Natural Sciences, Philadelphia; Carnegie Museum, Pittsburg,
Pa.; Field Columbian Museum, Chicago; and the collections of Samuel N. Rhoads
and Charles F. Batchelder.

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. (

Aery little is kiiowu of the geological history of Zapus, except that
the genus dates back to the Postpliocene. In 1871 Prof. E. D. Cope
recorded a mandibular ramus with incisor and second molar of a Jumj)-
ing Mouse from the Port Kennedy Bone Cave in Pennsylvania.' On
account of lack of recent specimens for direct comparison he was some-
what in doubt as to the specihc identity of the remains, but referred
tbem to Jaculus (= Zapus) hndsonius.'

External characters. — The external and cranial characters of Zapus
have been so exhaustively treated by Baird, Coues, and others that
a brief reference to them here will be sufficient. The body is consider-
ably enlarged posteriorly, the hind legs and tail greatl}' developed, the
forelegs rather short, and the ears somewhat longer than the surround-
ing fur. All the sjiecies agree closely in color and markings. A broad
dorsal stripe of some shade of yellowish brown, quite thickly flecked
with black-tipped hairs, is bordered by a slightly narrower lateral stripe
of a lighter color and usually with less black. The fur of the entire
upper parts is uniform plumbeous at base, only the tips of the hairs
att'ecting the external appearance of the animal. Between the darker
color of the sides and the white of the lower parts is a narrow stripe
of clear yellowish orange, unmixed with black, and with the base of
the hairs Avhite. The lower parts are white, sometimes suffused with
the color of the sides, and one species, Z. setchuanus, from China, has
a well-defined brown ventral stripe. The tail is very long, usually about
60 per cent of the total length, and more or less distinctly bicolor,
brownish above and whitish below. It is thinlj" covered with short
hairs, which are longer on the terminal part and form a short pencil.
In two of the three subgenera the tail is consi)icuously tipped with
white. The spring and early fall jielages differ noticeably, except in
members of the subgenus Xapaozapui'. the bright color of the spring
and early summer pelage becoming duller in the fall, and the dark dorsal
area partially obliterated. In some species, however, the dorsal area
is darker and more sharply defined in late summer and early fall than
in spring and early summer.

The teats are normally 8, and arranged in pairs at regular intervals.
The pectoral pair is situated just back of the forelegs, the inguinal pair
far back at the base of the thighs, and the remaining pairs between.
Sometimes the anterior or posterior pair remains undeveloped.

Habits. — In general but one species occurs in a given locality, but
Zapus Imdsonins, wLich has the widest range of all, seems to be always
present throughout the range of the subgenus Xapa'ozapus, the members
of which are almost invariably found in deep woods near streams. All
the other species delight in meadows, shrubby fields, and thickets along

' Preliminary report on the Vortebrata discovered in the Port Kennedy Bone Cave
<Proc. Am. Philos. Soc, XII, p. ^Q. 1871.

Vertebrate Eemains from Port Kennedy Bone Deposit <Joiirn. Acad. Nat. Sci.
Phila., 2d ser., XI, p. 200, 1^99.

■^Spelled huiUonianus by Zittel, Haudbiich der PaLeoutohjgie, IV, p. 527, 1893.

8 NORTH AMERICAN FAUNA.

the edges of woods, and show a marked preference for moist locations.-
During' the summer Jumping Mice are often seen in meadows and fiekls,
and are more readih' detected during the haying season than at other
times, when they are driven from their hiding places while the grass is
being cut. When disturbed tliey move oft' by a series of frog like leaps,
and often remain motionless after a jump or two, especially when
frightened from a nest. In leaping they are greatly assisted by their
long tails, which aid in preserving their balance as in the case of otber
animals similarly endowed. If, by any accident, a portion of the tail is
lost, the power of balancing is greatly impaired, and the animal, if
startled, seems unable to pursue a direct course because of failure to
land properly on its feet. Some of the earlier writers ascribe to the
animal the power of leaping 4 or 5 yards. Such statements are probably
exaggerated,- but it is certain that Zapus does possess remarkable leap-
ing powers, and when disturbed will jump 0 or 8 feet, and under some
circumstances perhaps much farther. In their chosen haunts these
mice do not follow beaten paths or runways like many small mammals,
notably meadow mice, b*ut seem to wander rather indiscriminately,
availing themselves to some extent of natural pathways or open places.

The birth of the young (five or six in number) which occurs in a nest,
ordinarily underground but sometimes placed in a hollow tree, usually
takes place in May or June. Sometimes, however, it occurs as late as
September; from which it seems probable that a second litter is raised,
or that the breeding season continues throughout the summer. Three
specimens obtained by my brother at Wilmington, Mass., September 25,
1897, which with their parents were turned up by a plow, were so young
that the posterior upper molars had just appeared.

During the latter part of summer Zapus makes a globular nest of
grass about 4 inches in diameter, with a small entrance at the side.
All that I have seen were on the ground in meadows among thick grass
or small bushes. One that I examined was made entirely of the straight
narrow leaves of grass, and was a very beautiful little home; but though
seemingly compact, it was so frail that it was impossible to preserve it.
These nests are usually occupied by two individuals, presumably a pair,
and seem to be used only at the close of the breeding season.

The hibernation of Zapus has been frequently noticed and several
interesting articles have appeared mainly on this habit.' All the species,
so far as known, hibernate during winter, though in one, at least, hiberna-
tion is not always complete — that is, individuals may awake and come
out during warm intervals, after the manner of the common skunk. Dr.
Merriam mentions having seen Jumping Mice abroad in northern Kew
York on several occasions during the unusually mild winter of 1881-82.
Messrs. H. H. and C. S. Brimley report that they have no positive evi-
dence of the hibernation of Zapus hudsonius amerieanus at Ealeigh,
N. C, although they add that the animal is too rare to enable them to

'Davies, Traus. Liun. Soc. London, IV, p. 155, 1798; Barton, Trans. Am. Pliilos.
Soc, VI, p. 143, 1804 : Tenney, Am. Nat., VI, p. 330, 1872.

REVISION OF THE JUMPTXG MICE OF THE GENUS ZAPUS. 9

find out much about it. Hiberuation varies with the locality, but usu-
ally begins about the time of the first heavy frosts and lasts until
spring. The fall pelage is usually assumed and the animals become
ex(>eedingly fat before entering winter quarters. Although they often
lay up stores of food in nests or burrows during summer, it is not known
that they use this food during winter. The animals are generally found
singly (sometimes in pairs) in nests in holes at a depth varying from
a few inches to 2 or 3 feet below the surface. They lie rolled up like
a ball with the feet close together and tail curled about them. If
removed from the nest and subjected to a moderate degree of heat they
revive and in the course of a few hours move about freely, but gen-
erally resume their lethargic state if again exposed to cold. The pulse
and respiration are very slow.

Hibernation sometimes takes place in a nest above ground. Dr.
A, K. Fisher tells me that some years ago a Zajms was brought to him
at Lake George, Xew York, which some men at work in the woods had
found hibernating in sm elaborate nest of grass and moss, exposed by
moving a log. This nest was destroyed, but when the animal was
given material it constructed another, in which it resumed its inter-
rupted sleep. It ^as occasionally awakened, and observations were
taken on its weight, respiration, and heart beats, after which it would
relapse into its state of lethargy.

NOMENCLATUEE.

Five generic and 30 specific and subspecific names have been applied
to various forms of Jumping- Mice now included in the genus Zapus.
In order to discuss the ditferent species and subspecies intelligently it
becomes necessary to separately consider each of these names, which
may be taken up alphabetically.

GENEKIC NAMES.

Dipus Zimmermann, Geog, Geschichte des Menschen u. vierfiissigen
Thiere, II, p. 358. 1780. The original description of the genus Dipus
included six species: D. jaciiJiis, D. sagitta, Yerhua capensis { = Mus
cafer), I), longipes, D. tamaricinus and I), hudsonius. The name was
subsequently restricted to the Old World jerboas.

Gerbillus -Desmarest, Nou v. Diet. d'Hist. is^at., XXIY. p. 222. 1804. A
genus of Old World rodents allied to Meriones. Four nominal species
of American Jumping Mice were referred to this genus by IJafinesque
in 'Precis des Decouv. somiologiques ' (p. II, 1814), and Am. Monthlj'
Magazine (III, p. 440, Oct.. 1818).

Meriones Illiger, Prod. Syst. Mamm. et Avium, p. 82, 1811. The name
was originally proposed by Illiger for a group including Dipus tamari-
cinvs and D. meridianns. Cuvier apparently proposed it independently,
twelve years later, for the American Jumping Mouse (Dents des Mamm,,
pp. 187, 250, 1823j.

10 NORTH AMERICAN FAUNA.

Jaculus Erxleben, Syst. Xat., p. 404, 1777. A geims of Old World
jerboas. The name was first used for jSfortli American species by
Wagler (Nat. Syst. Ampbibien, p. 23, 1830).

Zapus Coues, Bull. U. S. Geol. Surv. Terr., I, 2d ser., No. 5, p. 253,
1875. The first tenable generic name based on a North American
Jumping Mouse.

SPECIFIC AND SUBSPECIFIC NAMES.

Acadicus (Meriones) Dawson, Edinburgh New Philos. Journ., new
series, III, p. 2, 1856. Dawson, after unmistakably referring the animal
since described as Zapus insignis to the Meriones lahradorms of Richard-
son's Fauna Boreali- Aniericana (i. e., Z. hndsonins), goes on to describe
Z. hudsonius from Nova Scotia, calling it provisionally Meriones acadicus.

Alascensis (Zapus) Merriam, Proc. Biol. Soc. Wash., XI, p. 223, July
15, 1897. Dr. 0. Hart Merriam has described a subspecies of Z. hud-
sonius under this name, from specimens collected at Yakutat Bay,
Alaska.

AUeni (Zapus) Elliot, Field Columbian Mus., Pub. 27, Zool. Ser., I,
No. 10, pp. 212-213, March, 1898. Under this name Mr. D. G. Elliot
described the form of Zapus inhabiting the Sierra Nevada in California.

Americanus (Dipus) Bartosi, Trans. Am. Philos. Soc, IV, No. XII,
p. 115, 1799. Under this name Benjamin Smith Barton described
specimens from the vicinity of Philadelphia, Pa. Later (ibid, VI,
1804) he discussed the habit of hibernation and presented a figure of
the animal. Mr. C. F. Batchelder has recently revived americanns for
the form of Z. hudsonius inhabiting the Upper Austral zone in the
Eastern States. (Proc. New Eug. Zool. Club, I, p. 6, 1899.)

Brachynrus (Gerbillus) ' Rafiuesque.' A name referred to by Fischer
(Syu. Mamm., p. 340, 1820) without reference, and included in the sj^n-
onomy of Zapus by Cones (Mon. N. Am. Rodentia, p. 4G8, 1877). I have
been unable to find the original reference, but the inapplicability of the
name is self-evident.

Canadensis (Dipus) Davies, Trans. Linn. Soc. London, IV, p. 157,
1798. Maj. Gen. Thomas Davies described specimens which he had
procured in the vicinity of Quebec, Canada. He undoubtedly referred
to an animal of the hudsonius type, and as specimens from that region
do not deserve separation fiom the typical form, the name becomes a
synonym of hudsonius.

Daviesii (Gerbillus) Rafinesque, 'Precis des Decouv. somiologiques,
p. 14, 1814.' Rafinesque is said to have renamed Dijjus canadensis after
its describer.

Hardyi (Zapus) Batchelder, Proc. New. Eng. Zool. Club, I, p. 5,
February 8, 1899. Mr. C. F. Batchelder has recently applied this name
to specimens from Mount Desert Island, Maine. From a careful study
of the material at present accessible, I can find no characters of impor-
ta.Qce to separate this form irom hudsonius as here restricted.

REVISIOX OF THE JUMPING MICE OF THE GENUS ZAPUS. 11

Hudsonius (Dipus) Ziinmerinanu, Geog. Gescliiclite des Meuscheii n.
vierfiissigen Thiere, II, p. 3o>>^ 1780. This is tbe first specific name pro-
posed for any species of Jiiinpiug Mouse iu America, and imdouMedly
refers to the common eastern species which has so long borne the name.
It is tlie only species thus far known to occur about Hudson Bay — the
locality of Pennant's 'long-legged mouse,' to which Zimmermanu
referred,

Imperator (Zapus) Elliot, Field Columbian Mus., Pub. 30, Zool, Ser., I,
No. 11, p. 228, February 1, 1893. Mr. 1). G. Elliot has recently described
a Zapus from the Olympic Mountains of Washington, but specimens
from that region not being considered separable from typical Z. trinofa-
tiis, the name becomes a synonym.

Iiisignis (Zapus) Miller, Am. Naturalist, XXV, p. 472, August, 1891.
Mr. Gerrit S. Miller, jr., described the species erroneously referred to
.1/. Uihradorius {Z. hudsonius) by Dawson, from specimens collected on
the Pestigouche River, New Brunswick.

Labradorius (Dipus) Kerr, Animal Kingdom, p. 27G, 1792. Based on
the 'Labradore Jerboid Rat' of Pennant's History of Quadrupeds, 1781.
Kerr says, " Inhabits Hudson's Bay and Labradore. This species was
sent over from Hudson's Bay by 31r. Graham." Although Labrador is
mentioned in the habitat of the species, the specimen came from Hud-
son Bay, and consequently the name shouhl be restricted to the animal
from that region, thus becoming a synonym o? hudsonius.

Ladas (Zapus) Bangs, Proc. New Eng. Zool. Club, I, p. 10, February
28, 1899. Under this name Mr. Outram Bangs has described, as a
subsi^ecies, the form of hudsonius inhabiting the Labrador Peninsula.

Leonurus (Gerbillus) Rafinesfjue, Am. Monthly Mag., HI, p. 446, Octo-
ber, 1818. From the description — "body fallow, ears very long, white
inside, tail as long as body, black with a fallow tuft at end, length G
inches, body 3" — the animal is plainly not referable to Zapus.

Longipes (Mus) Zimmermaun, in Pennant's Arkt. Zool. I, p. 131, 1787.*
Zimmermanu erroneously identified a Jumping Mouse in spirits, sent
from Hudson Bay by Mr. Graham, as Mr(s longipes of Pallas, an animal
from the deserts about the Caspian Sea.

Macrourus (Gerbillus) 'Rafinesque,' a name referred to by Fischer (Syn.
Mamm., p. 340, 1829), without reference, and included in the synonomy
of Zapus by Dr. Coues (Mon. N. Am. llodentia, p. 408, 1877). I have
been unable to find the original reference.

Megalops (Gerbillus) Rafinesque, Am. Monthly Mag., Ill, p. 446, Octo-
ber, 1818. This animal he describes as follows: "Body gray, belly
white." * * * "Total length 5 inches, body only 2. From the
barrens of Kentucky," This can not apply to any species of Jumping
Mouse.

Mellivorus (Dipus) Barton, Trans. Am. Philos. Soc, VI, p. 143, 1804.

*Fi(le Coues, Mon. N. Am. Eodeiitia, p. 407, 1877. I have not seen this edition of
Pennant.

12 NORTH AMERICAN FAUNA.

This name occurs incidentally iu the text, being applied to a supposed
new species, with remarks on its reputed habit of passing the winter in
beehives and living on the honey.

Microcephalus (Meriones) Harlan, Proc. Zool. Soc. Loudon, p. 1, 1839.
Under this name Dr. Harlan described specimens from the vicinity of
Philadelphia, Pa. The name, being antedated by anierieamis Barton,
becomes a synonym.

Montanus (Zapus) Merriam, Proc. Biol. Soc. Wash., XI, p. 104, April
20, 1897. Br. C. Hart Merriam described the form inhabiting the Cas-
cade Range in Oregon, under the name Zapus trhiofatKS montanus, but
the characters seem to warrant its recognition as a full species.

Nemoralis (Meriones) Geoft'roy St. Hilaire, Diet. Classique d'Hist. I^at.,
VII, p. 323, Fev., 1825. Under this name Geoft'roy published a careful
description of two specimens (without locality), iu the Museum d'His-
toire ISTaturelle of Paris, which were figured by F. Ouvier and served as
the basis of Cuvier's genus Meriones. There is nothing iu the descrip-
tion of ncmoralis that gives any clue to the specific identity of the animal.

Niger (Gerbillus) Eafinesque, Am. Monthly Mag., IV, p. 100, December,
1818. A nomennudum used by Eafinesque in an article entitled 'General
Account of the Discoveries made in the Zoology of the Western States.'

Pacificus (Zapus) Merriam, Proc. Biol. Soc. Wash., XI, p. 104, April
25, 1897. Under this name Dr. C. Hart Merriam described a form
represented in the Biological Survey collection by specimens collected
in upper Rogue River Valley, Oregon.

Princeps (Zapus) Allen., Bull. Am. Mus. Nat. Hist., V, pp. 71-73, April
28, 1893. Dr. J. A. Allen described under this name a large species
from Florida, La Plata Couuty, Colo., which proves to be a very dis-
tinct form inhabiting the Rocky Mountain region of the United States.

Saltator (Zapus) Allen, Bull. Am. Mus. Xat. Hist., N. Y., XII, p. 3,
March 4, 1899. Dr. J. A. Alleu has recently described under this name
a well-marked form inhabiting northern British Columbia and the
adjacent region.

Setchuanus (Zapus) Pousargues, Ann. Sci. Nat., 8e ser., I, Xo. 4, p. 220,
Avril, 1896. The only name based on the single Old World species of
Zapus.

Soricinus (Gerbillus) Rafinesque, 'Precis des D;'Couv. somiologiques
p. 14, 1814.' I have not seen the original reference.

Sylvaticus (Gerbillus) Rafinesque, Am. Monthly Mag., Ill, p. 354, Sep-
tember, 1818. A nomen nudum used by Rafinesque in a letter addressed
to "Samuel L. Mitchill, president, and the other members of the
Lyceum of Xatural History, * * * dated at Louisville, Falls of
Ohio, 20 July, 1818.''

Tenellus (Zapus) Merriam, Proc. Biol. Soc. Wash., XI, p. 103, April 26,
1897. Under this name Dr. C. Hart Merriam has described a form
from Kamloops, British Columbia.

Trinotatus (Zapus) Rhoads, Proc. Acad. Xat. Sci., Phila., 1849, p. 421
(January 15, 1895). Under this name IMr. S. X. Rhoads described, from

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 13

specimens taken on Lulu Island, near the mouth of Fraser River, a
well marked form inhabiting- the Pacific coast region from Fraser
Kiver, British Columbia, to Humboldt Bay, Calif.

Genus ZAPUS Coues.

Dipus Ziminermauti, Oeog. Geschithte d. Meiischeu n. vierfiissigen Thiere, II, ji. 358,

1780.
Gerbillus Eafiuesqiie, 'Precis des Decouv. somiologiques, p. 14, 1814'. (Not Gerbillus

Desmarest, 1804.)
Meriones F. Cuvier, Dents des Maairu., pp. 187, 256, 1823. (Not Meriones lUiger, 1811.)
Jacnlus Wagler, Nat. Syst. Amphibien, p. 23, 1830. (Not Jacuhis Erxlebeu, 1777.)
Zapns Coues, Bull. U. S. Gaol. Surv. Terr., I, 2d ser., No. 5, p. 2.53, 1875.

Characters. — Skull not massive; braiu case rather high and rounded;
antorbital foramen large and oval; zygomata not widely spreading,
broadly expanded anteriorly where the malar extends upward to the
lachrymal. Teeth 16 or 18 (upper premolar i^reseut or wanting);
enamel much folded ; upper incisors compressed, much curved, deeply
salcate, and deeji orange.

Body enlarged posteriorly; hind legs and tail greatly developed, the
latter much exceeding length of head and body. Tail slender, uni-
formly tapering. Toes of hind foot five, each with separate metatarsal.
Upper parts ochraceous; median dorsal band dark. Under parts
white (often suffused with ochraceous) ; one species with brown ventral
stripe.

KEY TO THE SUBGENERA OP ZAPUS.

Premolar present:

Enamel folds closely crowded; tail not tipped with white; ears rather Jong;

lower parts white or ochraceous Zapus

Enamel folds not crowded; tail tipped with white; ears rather short; lower

parts white with brown ventral stripe Eo-apus

Premolar absent :

Enamel folds closely crowded ; tail tipped with white; ears rather long; lower
parts white Napwozapus

LIST OK SPECIES AND SUBSPECIES, WITH TYPE LOCALITIES.

Subgenus Zapus.

Zapus alascensis. Yakutat, Alaska.

aUeni. Pyramid Peak, Lake Tahoe, California.

americanus. Philadelphia, Pennsylvania.

campestris nob. Bear Lodge Mountains, Wyoming.

hudsonius. Huds(m Bay. *

laclas. Kigoulette, Hamilton Inlet, Labrador.

major nob. Warner Mountains, Oregon.

minor nob. Wingard, Saskatchewan.

monfanus. Crater Lake, Mount Mazama, Oregon.

ncvadcnsis nob. Euby Mountains, Nevada.

orarins nob. Point Reyes, California.

oregonus nob. Elgin, Oregon.

pacificus. Prospect, Rogue River Valley, Oregon.

princeps. Florida, La Plata County, Colorado.

saltator. Telegraph Creek, Northwest Territory.

tenellus. Kamloops, British Columbia.

trinotatus. Lulu Island (mouth Fraser River), British Columbia.

14 NORTH AMERICAN FAUNA.

. Siibgeuus Xcqjwocapita.

(tbietorum uob. Peninsula Harbor, Ontario.

insignis. Restigouche River, New Brunswick.

roanensia uob. Magnetic City, Roan Mountain, North Carolina.

Subgenus Eozapus.
setchnaniis. Ta-tsien-lou, Szecbuen, China.

Subgenus ZAPUS Coues.

Tyjye Zapiis hitd.soiihts (Zimmermanu), from Hudson Bay.

Subgeneric characters. — Teeth : i. j^^, ^j. J^^, m. ^~l = 18. Upper pre-
molar very small, nearly cylindrical ; anterior upper molar large, broadly
oval; m 2 similar, but noticeably smaller; m 3 con-
^l^^^l^^^j© siderably smaller and nearly circular. Lower
^"'^^^^^^^"^ molars similar in shape and relative size (fig. 1).
/^^)^^^\f^^^h, Enamel folds closely crowded. Interorbital con-
^'^^^^'^^mimj^^^^'ii^^^^i^^ strictiou or medium width; irontal region moder-
FiG. 1— Molar teeth of atcly swollcu; brain case rather high; rostrum
zapus hud807iius from gieuder and somewhat attenuated.

Tower, Minnesota (No. ...

72695, u. s. xat. Mus.) Uppei parts as in entire genus; lower jjarts white,
X 6- sometimes suffused with color of sides; tail more or

less distinctly bicolor, normally without white tip.

KEY TO SPECIKS OF THE SUBGENUS ZAPUS.

Lower parts mucb suffused with color of sides orariiis.

Lower parts not much suffused with color of sides.
Size, large; hind foot usually more than 32 mm.
Ears never conspicuously bordered with whitish.

Larger; pencil of tail never normally white; lower parts often suffused with

ochraceous trinotatns.

Smaller; pencil of tail usually white; lower parts never suffused with

ochraceous alhni.

Ears usually conspicuously bordered with whitish.

Sides light ochraceous buff nevadensis.

Sides not light ochraceous buff.

Skull narrow; incisive foramina very large mtUuior.

Skull broad; incisive foramina not very large.

Hind foot about 35 mm major.

Hind foot about 32 to 34 mm.

Incisive foramina broad posteriorly oregonus.

Incisive foramina narrow posteriorly prbiceps.

Size medium or small; hind foot usually below 32 mm.

Size medium; hind foot 30 to 32 mm. ; total length of adults more than 215
mm.
Skull rather heavy ; brain case broad; incisive foramina rather large.

Dorsal area not very distinct patlficua.

Dorsal area distinct.

Rostrum considerably deflected minor.

Rostrum not much deflected montanus.

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 15

Skull rather light: lirain case uarrt)W ; incisive I'nrauiiiia small.
Coloratiou dark and dull.

Larger; basilar length of skull usually more than 18 mm.

alasceiisis.
Smaller; basilar length of skull usually under 17.5 mm. .ieneUns.
Coloration not dark and dull.

Dorsal area very much flecked with color of sides; fall pelage

with dorsal area nearly black campestris.

Dorsal area not much flecked Avith color of sides; fall pelage
with dorsal area not very dark.
Larger; hind foot about 32 mm.; tail usually more than

140 mm ladas.

Smaller; hind foot usually 30 or 31 mm. ; tail usually below

135 mm hudsonins.

Size small; bind foot usually below 30 mm.; total length of adults usually
under 215 mm americanus.

ZAPUS HUDSONIUS (Zimmcrmaun). Hudson Bay Jumping Mouse.
(PI. I, fi^s. 3, 3a.)

1780. Dipus hudsonins Zimmermann, Geog. Geschichte d. Menscheu u. vierfiissigen

Thiere, II, p 358.
1792. Dipus lahradorius Kerr, Animal Kingdom, i>. 276 (liased on the " Labradore

Jerboid Eat of Pennant ' ). '
1798. Dipus canadensis Davies, Trans. Linn. Soc. London, IV, ji. 157.
1822. Gerhillus canadensis Desmarest, Mammalogie, II, p. 321.
1825. Gerbillus luhradorius Harlan, Fauna Am., p. 157 (in part).

1829. Meriones iahradorius Richardson, Fauna Boreali-Americaua, I, p. 141. (Not M.
lahradurius Dawson, 1856.)

1830. Jacnius americanus AVagler, Nat. Syst. Amphibieu, p. 23.

1839. Meriones microccphalns Harlan, Proc. Zool. Sec. London, 1839, p. 1.
1843. Jaculus luhradorius AVaguer, Suppl. Schrebcr's Saugthiere, III, p. 294.

1856. Meriones acadicus Dawson, Edinburgh New Philos. Journ., new ser.. Ill, p. 2.

1857. Jaculus hndsonius Baird, Mamm. N. Am., p. 430 (in jiart).

1875. Zapus liuusonius Cones, Bull. U. S. Geol. Surv. Terr., 2d Ser., No. 5, p. 253; Mou. N.

Am. Rodentia, p. 467, 1877 (in part), and recent authors (in part).
1899. Zapus hudsonins canadensis Batchelder, Proc. New Eug. Zool. Club, I, p. 4.
1899. Zapus hudsonins hardiji Batchelder, Proc. New Eng. Zool. Club, I, p. 5. (Mount

Desert Island, Maine.)

Type locality. — Hudson Bay.

Geographic distribution. — As restricted iu the present i^aper, tliis spe-
cies is found from the soutberu shores of Hudsou Bay south to Xew
Jersey, and in the mountains to Xorth Carolina, west to Iowa and Mis-
souri, and northwest to Alaska. It intergrades Avith Z. americanus in
the upper i)art of the Carolinian zone in the Eastern States; with cam-
pestris along the edge of the Great Plains, and probably with alascensis
somewhere iu northern British America.

General characters. — Size medium, smaller than Z. insignis, Z. trinoia-
tus, or Z. princeps. Skull less massive, with small molars and incisive
foramina.

' "Inhabits Hudson's Bay and Labradore. This species was sent over from Hud-
son's Bay by Mr. Graham." — Kerr.

16 NORTH AMERICAN PWUNA.

Color. — Summer pelage : Fur of entire upper parts i)lumbeous at base,
with tips of hairs ranging- from yellowish fawn to rather dark oehra-
ceous, rather thickly flecked with black-tipped hairs; broad dorsal
sti'ipe reaching- from upper part of face to base of tail much darker than
remainder of upper parts, the black-tipped hairs predominating; tail
rather sharply bicolor, grayish brown above and whitish beneath ; lower
parts white, sometimes suffused with color of sides; feet white. Fall
pelage: Ochraceous of sides duller and more yellowish, encroaching
more on the dark dorsal area, which is sometimes almost obliteiated.
Young of the year with more ochraceous in fall pelage than the adults,
and a greater tendency to fulvous wash on lower parts.

Cranial characters. — Skull rather lightly built, with rather narrow
brain case, small incisive foramina, and small molars; zygomata not
widely spreading; rostrum not appreciably deflected.

Measurements. — Two specimens from James Bay, Canada, measured
from alcohol, average: Total length, 209; tail vertebra^, 124; hind foot,
30.5. Eleven from Tower, Minn., average: Total length, 218; tail ver-
tebne, 133 ; hind foot, 30.2. Thirty-four adults from Keene Valley, Essex
County, N.Y., average:^ Total length, 21G.G; tail vertebrae, 129.25; hind
foot, 31.25. Skull: An adult skull from James Bay measures: Basilar
length, 18.5; zygomatic breadth, 11; mastoid breadth, 10; interorbital
constriction, 4.2; incisor to posti)alatal notch, 8.5; foramen magnum to
postpalatal notch, 8; fronto-palatal depth at middle of molar series, 6.5.
An adult ? from Ossipee, N. H., measures: Basilar length, 18.8; zygo-
matic breadth, 11. G; mastoid breadth, 10; interorbital constriction, 4,3;
incisor to postiialatal notch, S.6- foramen magnum to postpalatal notch,
8.2; fronto-palatal depth at middle of molar series, G.2.

General remarJiS. — Zapus hudsonius^ the type of the genus, was until
recently the only well established species. The foregoing list of
synonyms indicates that it attracted the attention of many naturalists
during the early part of the century, all the names jDublished previous
to 1856 referring almost exclusively to it. Jaculus hudsonius of Baird
[= Zapus hudsonius Coues) is a composite species including nearly a
dozen forms now recognized as distinct. The distribution of hud-
sonius, including its four well-marked subspecies, is by far the most
extensive in the genus, few North American rodents ecpialing it in
this respect.

The only specimens of Zapus from Hudson Bay at present available
are four alcoholic examples, in rather poor condition, from James Bay,
and a part of a skin from Fort Churchill, Northwest Territory. This
skin has evidently been in alcohol, and hence its color can not be
depended upon. I have removed several skulls from the James Bay
specimens, including one perfect adult. Since this skull agrees essen-
tially with skulls from Tower, Minn., from which place the Biological
Survey has a large series, these specimens have been assumed to be

' Fide Batclielder.

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 17

fairly typical and have been used for comparison in most instances. It
is qnite possible, liowever, that a series of skins from Hndson Bay
would measure slightly more than the ^Minnesota specimens, and would
perhaps differ in color. While this material, taken in connection with
what is available from the type locality is sufficient for comparison with
the subspecies here recognized, it will not permit the recognition of
other nominal forms which may eventually prove to be well founded,
A skull from Saint Catharines, Ontario, however, which may be con-
sidered a topotype of canadensis, is identical in appearance with one
from James Bay.
Specimens examined. — Total number, 319, from the following localities :

Northwest Territory: James Bay, 4; Fort Churchill, 1; Fort Eae, 1; Fort
Ke.solutioD, 1.

Ontario: Xepigon, 7; North Bay, 6; Mount Forest, 3; Michipicoten, 1 ; Saint
Catharines, 6.

Ne'w Brunswick: Oak Bay, 30.

Nova Scotia: Halifax, 6; James River, 2; Digby, 4.

Maine: Walker Pond, 4; Mount Desert Island, 4.

New Hampshire : Ossipee, 1 ; Fabyans, 1.

Vermont: Brandon, 1; Mount Mansfield, 1; St, Johnsbury, 2.

Massachusetts: War eh am, 70; North Truro, 6; Middleboro, 2; Wilmington,
3; Lunenburg, 2; Belmont, 2; Ipswich, 1; Concord, 4; Lexington, 1;
Dightou, 1; AVellesley, 1; Barnstal)]e Neck, 5.

Connecticut: East Hartford, 1 (not typical) ; Wetherstield, 2 (not typical).

New York: Peterboro, 39; Catskills, 8; Elizabethtown, 2; Waterville, 1;
Patten Mills, 1; Locust Grove, 7; Lake George, 5.

New Jersey : Lake Hopatcong, 5.

Pennsylvania: Leasuresville, 3.

Maryland: Cumberland, 1; Finzel, Garrett County (6 miles north of Frost-
burg), 1.

West Virginia : Wheeling, 1.

North Carolina: Roan Mountain, 1 ; Magnetic City, 1.

Ohio : Portland Station, 1.

Indiana : Terre Haute, 1.

Wisconsin : Racine, 1.

Michigan : Seney, 1.

Minnesota: Lac qui Parle, 1; Elk River, 25; Tovrer, 26.

ZAPUS HUDSONIUS LAD AS Bangs. Labrador Jumping Mouse,
Zapus hudsonius ladas Bangs, Proc. New Eug. Zool. Club, I, p. 10, Feb. 28, 1899.

Type locality. — Rigoulette, Hamilton Inlet, Labrador.

Geographic distribution. — Eastern Quebec north to Hamilton Inlet,
Labrador; limits of range unknown.

General characters. — Larger than typical Zapus hudsonius with longer
tail and hind foot; coloration darker; differing also in cranial char-
acters.

• Color. — Summer pelage: Sides rather dark, bright ochraceous buff",
model ately lined with blackish-tipped hairs; dorsal area usually less
distinct than in typical hudsonius, much flecked with ochraceous.
Lower parts white, usually more or less suffused with color of sides.
22095— is^o. 15 2

18 NORTH AMERICAN FAUNA.

Tail quite sliarply bicolored, dusky above and wbitisli beueatb; ears
usually edged witb ocbraceous (wliitisli iu liudsonius). Feet wbite.
Fallpclaf/e: Dorsal area usually eveu less distinct tban iu summer, with
tbe general tinge of entire upper parts yellowisb instead of ocbraceous.

Cranial characters. — Skull longer tban tbat of typical hudsoniiis;
zygomata considerably longer; zygomatic breadtb about tbe same as
in largest skulls of hudsonius; interorbital constriction sligbtly wider;
brain case bigber and averaging sligbtly longer; rostrum and nasals
longer; bulbie considerably larger; molar series averaging sligbtly
longer.

Measurements. — Type iso. 41G9 9 ad.j collection of E. A. and O.
Bangs: Total lengtb, 238; tail vertebne, 153; bind foot, 32. Average
o{ five adults from Black Bay, Labrador: Total lengtb, 228; tail verte-
brie, 142; hind foot, 32. ^SkMU : Average of four adults from Black
Bay, Labrador : Basilar lengtb, 18.8; zygomatic breadtb, 11.4; mastoid
breadtb, 10,2; interorbital constriction, 4.5; incisor to postpalatal
notch, 8.9; foramen magnum to postpalatal notch, 8.5; fronto-palatal
depth at middle of molar series, G.3.

General remarks. — Zapus li. ladas is a fairly well-marked subspecies
represented by 23 specimens in tbe collection of E, A. and O. Bangs,
and 5 in tbe Biological Survey collection. Tbe best characters separat-
ing this form from typical hndso7iins, as here restricted, are tbe longer
tail, larger hind foot, and longer skull, with larger audital bulla'.
Tbis latter character is very evident, eveu in young skulls.

Spechnens examined. — Total number, 28, from the following localities:

Labrador: Hamilton Inlet (type locality), 3; Black Bay, 20.
Quebec: Godbout, 5.

ZAPUS HUDSONIUS ALASCENSIS Merriam. Alaska Jumping Mouse.
Zapns hudsonius alascensis Merriam, Proc. Biol. Soc. "Wash., XI, p. 223, 1897.

Type locality. — Yakutat, Alaska.

Geographic distribution. — Yakutat Bay, north to Y^ukon Eiver; limits
of range unknown.

General characters. — Similar to Zapns hudsonius, hut slightly larger
and darker.

Color. — Sides dull, dark ocbraceous, with many black-tipped bairs;
dorsal area quite distinct, but tbickly flecked witb brownish; beueatb
pure wbite; tail rather sharply bicolor.

Cranial characters. — Skull similar to tbat of typical hudsonius, but
slightly longer and narrower; zygomata less bowed outward and
slightly longer; brain case narrower; molar series longer; crown of
last lower molar longer.

Measurements. — Average of four specimens from type locality: Total
lengtb, 218; tail vertebra', 132; bind foot, 31.5. Skull: Average of
two adults from type locality: Basilar lengtb, 18.2; zygomatic breadth,

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 19

10.7; mastoid breadth, 10.2; iuterorbital constriction, 4.2; incisor to
])OStpalatal notch, 8.5; foramen magnum ,to postpalatal notch, 8;
frouto-palatal depth at middle of molar series, C.2.

General remarks. — Zapus h. alascensis is a fairly Tvell-marked subspe-
cies based on four specimens collected at Yakutat Bay, Alaska. Four
additional specimens in the National Museum from Alaska have been
examined, but are in such poor condition as to furnish no additional
characters. This form undoubtedly iutergrades with true hudsonius
somewhere in northern British America.

Specimens examined. — Total number, 8, from the following localities:

Alaska: Nusliagak, 2; Yakutat Bay, 4; Yukon River, 2.

ZAPUS HUDSONIUS AMERICANUS (Barton). Carolinian Jumping Mouse.

Dipiis ameriraniis Barton, Trans. Am. Philos. Soc, IV, No. XII, p. 115, 1799.
Zapus hudsonius americanus Batchelder, Proc. New Eug. Zool. Club, I, p. 6, 1899.

Type locality. — Philadeli)hia, Pa.

Geographic distribution. — From vicinity of Ealeigh, N. C, north
through Upper Austral zone along coastal i)laiu to southern Connec-
ticut and lower Hudson Valley, intergrading in upper edge of its range
with Z. hudsonius.

General characters. — Similar to Zap us hudsonius, but smaller; dorsal
area less distinct; skull and teeth smaller.

Color. — Description of specimens in summer pelage from Ealeigh,
N. C. : Entire upper parts deep dull ochraceous, the dorsal area very
indistinct and but slightly mixed with black-tipped hairs; sides of head
much lighter than remainder of upper parts; ears very dark, nearly
black; under parts whitish, suffused with ochraceous. In the fall the
dorsal area is indistinct and very thickly flecked with j'ellowish-tipped
hairs; sides dull yellowish.

Cranial characters. — Skull considerably smaller than in Z. hudsonius.,
with zygomata more arched and brain case shorter; incisive foramina
shorter and relatively broader; interorbitiil breadth about the same;
rostrum relatively deeper and shorter; molar series shorter.

Measurements. — Average of 7 adults from Ealeigh, N. C: Total
length, 191.3; tail vertebrie, 115.4; hind foot, 28.3.i Two Trom Wash-
ington, D. C, average: Total length, 198.5; tail vertebrfe, 117.5; hind
foot, 28. Skull: Average of two from Ealeigh, IST. 0.: Basilar length,
17; zygomatic breadth, 10.3; mastoid breadth, 9.2; iuterorbital con-
striction, 3.9; incisor to postpalatal notch, 8,2; foramen magnum to
postpalatal notch, 7.5; fron to-palatal depth at middle of molar series, 6.

General remarks. — It is unfortunate that the name americanus must
be adopted for this subspecies, since it was based on specimens clearly
intermediate between the extremes of this form and true hudsonius.

' Fide Batchelder.

20 NORTH AMERICAN FAUNA.

Specimens examined. — Total uumber, 54:, from the followiug localities:

New York : Long Islaml, 3 ; Montauk Point, 8; Raynor Beach, 1; Roslyn, 3;
Sing Sing (not typical), 1.

New Jersey : Chatham, 1; Englewood, 1; Mount Holly, 2; 'New Jersey', 1;
Phillipsburg, 1.

Pennsylvania : Chester County, 1 ; Culver Lake, 2; Delaware County, 2; Mar-
pie (Delaware County), 1; Schuylkill,!; Tinicum (Delaware County), 1;
Upper Darby, 1.

Maryland: Kensington, 1; Laurel, 4; Prince George County, 5; Sandy
Spring, 1.

District of Columbia : Washington, 2.

Virginia: Accotink, 1; Falls Church, 1.

North Carolina : Raleigh, 7; Weaverville, 1.

ZAPUS HUDSONIUS CAMPESTRIS subsp. uov. Prairie Jumping Mouse.

Type from Bear Lodge Mountains, Wyoming, No. 65872, $ ad., U. S. Nat. Museum,
Biological Survey Coll. Collected June 12, 1894, by B. H. Dutcher. Original
No. 600.

Geographic distribution. — Great Plains from Manitoba southward to
Nebraska and westward to Colorado and Wyoming.

General characters. — Similar to Z. hudsonius, but slightly larger and
brighter in color; differing also in cranial characters.

Color. — Summer pelage: Sides bright ochraceous-buflf, similar to
brightest specimensof //Mrfsow/?fs from Eastern States, moderately lined
with black-tipped hairs; dorsal area distinct, strongly suffused with
color of sides; tail yellowish white below, dark grayish above. Fall
pelage: Sides dull yellowish instead of ochraceous; dorsal area nearly
black, very slightly flecked with yellowish-tipped hairs; dorsal surface
of tail much darker than in summer; ears darker, conspicuously
bordered with whitish.

Crania I characters. — Skull slightly larger than in Z. hudsonius. Skulls
from the type locality, compared with those of Z. hudsonius from Tower,
Minn., agree in interorbital breadth; brain case about as wide but
higher; incisive foramina slightly larger; process on lower border of
maxillary portion of zygoma more pronounced; viewed in profile, the
cranium is more evenly curved.

Measurements. — Average of 4 adults from type locality : Total length,
222; tail vertebric, 135; hind foot, 30.5. ShuU: Average of 4 adult
skulls from type locality : Basilar length, 18.7; zygomatic breadth, 11.4;
mastoid breadth, 10.4; interorbital constriction, 4.2; incisor to post-
palatal notch, 8.9; foramen magnum to postpalatal notch, 7.9; fron to-
palatal depth at middle of molar series, 6.3.

General remarls. — The present well-marked form replaces Z. hudsonius
over the northern and more elevated portions of the plains west of
Minnesota and Iowa to the base of the Eocky Mountains. The char-
acters of this form remain remarkably constant throughout its known
range. While no good intergrades have been examined, it doubtless

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 21

does iutergrade with true hudsonius iu western Minuesota. The dif-
erence between summer and early fall pelage is very great, apparently
greater than in any other species.
Specimens examined. — Total number, 35, from the following localities:

Manitoba: Carberry, 2; Portage la Prairie, 1; Red River settlement, 4; Sel-
kirk settlement, 1.

North Dakota : Devils Lake, 1 ; Fort Sisseton, 1 ; Fori Wadsworth, 1 ; Pem-
bina, 2.

Montana : Little Big Horn River, 1.

South Dakota : Cnster, 4.

Wyoming: Bear Lodge Menu tains (type locality), 6; Cheyenne, 1; Devils
Tower, 1 ; Sherman, 1 ; Sundance, 1

Nebraska : Columbus, 1.

Colorado : Loveland, 5.

Missouri : Jackson County, 1.

ZAPUS TENELLUS Merriara. Kamloops Jumping Mouse.

Zainis leneUiis Merriani, Proc. Biol. Soc. Wash., XI, p. 103, April 26, 1897.

Type locality. — Kamloops, British Columbia.

Geographic distrihution. — Known only from vicinity of type locality.

General characters. — Size rather small; ears rather large; coloration
dark.

Color. — Early faU pelage : Somewhat similar to Zap us hudsonius in
corresponding pelage but much darker; sides olive yellowish, heavily
intermixed with black-tipped hairs; dorsal area somewhat as in hud-
sonius, but not well defined, contrasting less with sides; tail sharply
bicolor, darker above and at tip than in hudsonius ; nose, ears, and
outer sides of thigh and forearm dusky, much darker than in hudsonius;
fore and hind feet soiled whitish.

Cranial characters. — Skull similar to that of Zapus hudsonius, but
less swollen in frontal region; border of ascending portion of jugal
straigliter; cranium more arched.

Measurements. — Average of four specimens from type locality: Total
length, 208; tail vertebrt^, 128; hind foot, 30..J. Skull: Average of 3
skulls from type locality : Basilar leugtb, 1G.5; zygomatic breadth, 10.6;
mastoid breadth, 9.0; interorbital constriction, 4.2; incisor to post-
palatal notch, 7.0; foramen magnum to postpalatal notch, 6.9; fronto-
palatal depth at middle of molar series, 5.7.

General remarks. — Zapus tenellus is a well-marked form requiring
comparison only with Z. hudsonius and its subspecies alascensis. In
color it resembles alascensis, but in cranial characters it departs from
hudso7iius iu anothev direction. There is a possibility that it may be
found to intergrade with one of them to the northward, but until such
intergradation is proved, it seems best to allow it full specific rank.

Specimens examined. — Total number, 7, from the following localities:

British Columbia : Ducks, 2 ; Kamloops, 5.

22 NORTH AMERICAN FAUNA.

ZAPUS TRINCEPS Allen. Rocky Mountain Jumping Mouse.

(PI. I, figs. 6, 60.) -

Zapus princeps .J. A. Allen, Bull. Am. Mus. Nat. Hist., N. Y., V, p. 71, April 28, 1893.

Type locality. — Florida, La Plata County, Colo.

Geographic distribution. — Rocky Mouutaiu region from northern New
Mexico northward to Henry House, Alberta.

General characters. — Size large; skull broad and heavy, much as in
Z. hndsonius, but much larger.

Color. — Slimmer pelage: "Above with the middle of the dorsal region
pale yellowish brown, i^rofusely inixi d with blackish, so that sometimes
the blackish color, sometimes the pale yellowish brown, predominates;
sides of the body, forming a baud on either side about equal to the
dark dorsal area, yellowish brown, slightly mixed with blackish, except
over a narrow lateral line, adjoining the white of lower parts, which is
a clear, strong yellowish brown ; lower ])iiTts white to the base of the
hairs, varjdug in some specimens to strong ochraceous; tail indistinctly
bicolor — grayish white below and pale brown above, and very thinly
haired; hind feet grayish white above like the lower surface of the
tail; ears narrowly edged with yellowish white."' In fall pelage stdnlts
have the dorsal area with more black and sides more yellowish than in
summer; under i^arts pure white; immature in same pelage similar,
but with dorsal area more suffused with yellowish; under parts some
what suffused with color of sides.

Cranial characters. — Skull large and heavy, with rather heavy denti-
tion. In some specimens the premolar is very small, occasionally being
flattened against the adjacent tooth and scarcely functional. Imma-
ture skull short and broad, with large brain case. The incisive foramina
in this and closely related species much larger than in those more nearly
lelnted to Z. hudsonius.

Measurements. — Average of 7 specimens from Fort Garland, Colo. :
Total length, 245; tail vertebra^, 147; hind foot, 32. Average of 4 from
La Barge Creek, Wyo. : Total length, 239; tail vertebrae, 150; hind
foot, 32. Average of 6 from St. Mary Lake, Mont. : Total length, 232;
tail vertebrae, 139; hind foot, 32. Slull: Average of 2 adult skulls
from Fort Garland, Colo. : Basilar length, 20.7 ; zygomatic breadth, 12.5 ;
mastoid breadth, 11; interorbital constriction, 4 5; incisor to post-
palatal notch, 9.8; foramen magnum to postpalatal notch, 8.8; fronto-
palatal depth at middle of molar series, 0.0. One irom Preuss Moun-
tains, Idaho: Basilar length, 20.5; zygomatic breadth, 13.2; mastoid
breadth, 11; interorbital constriction, 4.8; incisor to postpalatal notch,
9.8; foramen magnum to postpalatal notch, 8.8; frontopalatal depth
at middle of molar series, 0.5.

General remarls. — Zapus princeps is one of the largest species of
the genus and has the widest range of any except Z. hudsonius. It
is mostly confined to mountains, though northward its range extends

'From original description in Bull. Am. Mus. Nat. Hist., N. Y., V, p. 71, 189.3.

REVIH.ION OF THE JUMPING MICE OF THE GENUS ZAPUS. 23

over the plaius of Assiniboia and Saskatchewan, where it grades into
the form here described as subspecies minor.
Specimens examined. — Total number, 101, from the following localities:

Alberta: Crow Nest Pass (Rocky Monntaius near forty-ninth parallel), 1;

Henry House, 2; 15 miles south of Henry House, 3.
British Columbia: Field, 3; Glaeier, 1.
Montana: Bear Pan' Mountains, 6; Big Snowy Mountains, 12; Blackfoot

Agency, 2; Fort Ellis, 1; Paola, 1; Pryor Mountains, 1; St. ^larys Lake,

13; Suuiniit, 5; Upper Stillwater Lake, 2.
Idaho: Preuss Mountains, 8; Salmon River ]\Iountaius, 5.
Wyoming: Clark Fork, 1; La Barge Creek, 11; Yellowstone Lake, 1.
Colorado: Co( hetope Pass, 1; Florida (type locality), 5; Fort Garland, 10;

Gold Hill, 2; Rocky Mountains (39-), 1.
Nevr Mexico: Camp Burgwyn, 2; Santa Fe, 1.

ZAPUS PRINCEPS MINOR subsp. nov., Saskatchewan Jumping Mouse.

Tij2)e from Wingard, near Carlton House, Saskatchewan, No. 73673, 9 ad., U. S. Nat.
Museum, Biological Survey Coll. Collected July 2.3, 1895, hy J. Alden Loring.
Original No. 3123.

Geograpliie distribution. — Plains of Saskatchewan; limits of range
unknown.

General characters. — Similar to Zapiis princeps, but smaller and difter-
iug in color and cranial characters.

Color. — Summer pelage: Similar to Z.2)rinceps, but dorsal area darker;
lower iiarts suffused with salmon. Fall pelage with dorsal area thickly
tiecked with yellowish and sides yellowish olive; lower parts much as
in summer.

Cranial characters. — Skulls of the present form from Osier and AVin-
gard, Saskatchewan, compared witU those of Z. prinreps from Colorado,
differ as follows: Smaller; rostrum shorter and more deflected; nasals
much narrower anteriorly ; brain case relatively higher; interpterygoid
fossa shallower.

^[easurements. — Average of 10 specimens from Osier, Saskatchewan:
Total length, 219; tail vertebra^, 131; hind foot, 29.0. Average of 2
from tyi)e locality: Total length, 220; tail vertebni^, 131,5; hind foot,
28. Skull: Average of 3 skulls from Osier, Saskatchewan: Basilar
length, 18.7; zygomatic breadth, J 2.2; mastoid breadth, 10.7; interor-
bital constriction, 4.1; incisor to postpalatal notch, 9; foramen mag-
num to postpalatal notch, 8.3; fronto-palatal depth at middle of molar
series, C.7.

General remarks. — Z. minor is a well-marked subspecies requiring
comparison only with Z. princeps. It is evidently the northern plains
representative of that species, probably intergrading with it in Assini-
boia and eastern Alberta near the United States boundary and along
the eastern base of the Canadian Rockies. Lack of specimens from
this region makes it impossible to conclusively sel;tle this point.

Specimens examined. — Total number, 21, from the following localities:

Saskatchewan: Osier, 15; Wingard (type locality), 3.
Assiniboia: Indian Head, 3.

24 NORTH AMERICAN FAUNA.

ZAPUS PRINCEPS OREGONUS snbsp. iiov. Blue Mountaius Juiiipiu<v Mouse.

Type from Elgin, Blue Mountaius, Oregon. No. 78156, i ad., U. S. Nat. Museum,
Biological Survey Coll. Collected May 29, 1896. by Edward A. Preble. Original
No. 959.

Geographic distribution. — Blue Mountains of Oregon.

General characters. — Similar to Zapus 2Jrinc€ps, but differing in color
and in cranial cliaracters.

Color. — Early summer pelage : Very similar to Z.princeps^ but slightly
lighter on sides, especially anteriorly; dorsal area and head more finely
tiecked with color of sides, the head especially presenting a grayish
appearance ; beneath j)ure white ; whitish edging of ears very indistinct.

Cranial characters. — Compared with Z. princeps from Colorado, the
skull of the present form differs as follows: Brain case more rounded,
especially shorter and more rounded behind; zygomata shorter; inci-
sive foramina larger and very much broader behind; audital bulhe
slightly smaller.

Meastirements. — Type: Total length, 250; tail vertebrae, 154; hind
foot, 33. Average of 3 adults from Strawberry Butte, Oregon: Total
length, 243; tail vertebra^ 140; hind foot, 32. Slcull: Average of 3
adult skulls from type locality : Basilar length, 20,1 ; zygomatic breadth,
12.9; mastoid breadth, 11; interorbital constriction, 4.4; incisor to post-
palatal notch, 0.7; foramen magnum to postpalatal notch, 8.5; fronto-
palatal dei)th at middle of molar series, 0.5.

General remarks. — The present form seems to replace Z. princeps
throughout the Blue Mountains region of northeastern Oregon. Speci-
mens from this region agree very constantly with each other and differ
from typical Z. princeps as above indicated. It is much larger than and
not at all closely related to Z. montanus, its relative on the west. A
specimen from Mountain City, Nev.. in the Brunneau Mountains, though
differing in many details, is doubtfully referred to this form. Exter-
nally it differs considerably from Z. oregoniis and agrees with Z. neva-
decisis, from the Euby Mountains, Nev., though the skull shows that it
is not closely related to that form. The skull of the Mountain City
specimen also differs somewhat from that of typical Z. oregouus, and addi-
tional material will doubtless show it to represent a form which merits
separation.

Specimens examined. — Total number, 9, from the following localities:

Oregon: Elgin, 3; Harney (10 miles north), 2; Strawberry Butte, 3.
Nevada: ^Mountain City (not typical), 1.

ZAPUS MAJOR sp. nov. Warner Mountain Jumping Mouse.

I'Hjje from Warner Mountains, Oregon. No. 79983, 9 ad., U. S. Nat. Museum, Biolog-
ical Survey Coll. Ceik-cted August 4, 1896, by C. Hart Merriam and Vernon
Bailey. Original No. 5720.

Geographic distribution. — Known only from the type locality.
General characters. — Similar externally to Zapus princeps^ but larger;
skull closelj" resembling that of Z. trinotatus.

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 25

Color. — Type in summer pelage: Sides ocbraceous-buff, moderately
lined with black-tipped hairs; back slightly darker, thickly flecked
with black; tail not sharply bicolored; beneath, dull white; feet soiled
white.

Cranial characters. — Skull massive; brain case high and broad;
zygomata rather short; palate broad and long; interpterygoid fossa
broad and shallow, with bordering edge of palate much excavated.
IMaxillary portion of zygomata heavy and nearly jierpendicular to main
axis of skull; incisive foramina large and elliptical. The skull of Z.
major difiers from those of Z. trinotatus taken near type locality and from
northern Washington, as follows: Rostrum larger and nasals broader;
brain case higher; ascending portion of jugal steeper.

Measurements. — Type: Total length, 255; tail vertebrie, 155; hind
foot, 35. ISlxuU (type): Basilar length, 20.4; zygomatic breadth, 13;
mastoid breadth, 11.2; iuterorbital constriction, 4.7; incisor to post-
palatal notch, 10; foramen magnum to postpalatal notch, 8.0; fronto-
palatal depth at middle of molar series, 0.5.

General remarls. — The ])resent form is based on a single adult speci-
men from the Warner Mountains, in southern Oregon. The skull bears
some resemblance, especially in the form of the brain ease, to those ot
Z. trinotatus and its subspecies all en i. It is apparently shut off by
natural barriers from all of the forms surrounding it, with the possible
exception of Z. montamis, and has no close afiinities with them. From
Z. montamis it differs to such an extent that intergradation is out of the
question.

Specimens examined. — The type.

ZAPUS NEVADEXSIS sp. uov. Nevada Jumping Mouse.

Type fvom Ruby Mountains, Nevada, No. 94185, 9a<^I-, U- S- Nat. Museum, Biological
Survey Coll. Collected Juue21, 18!)8, by Vernon Bailey. Original No. 6581.

Geo(iraphic distribution. — Known only from type locality.

General characters. — Size rather large; color light; molar series long
and rather nariow.

Color. — Dorsal area about as in Z. princeps^ pale yellowish-brown,
profusely mixed with black-tipped hairs; sides light ochraceous-buff,
becoming almost white on cheeks, moderately lined with black-tipped
hairs, the basal i)ortion of fur cinereous, noticeably lighter in color than
ill Z. princeps, with a few white hairs intermixed; beneath, pure white.

Cranial characters. — The skull of the type and only known specimen
is large, but rather lightly built. Compared with skulls of Z. princeps
from Colorado it is smaller and flatter; brain case shorter and more
rounded; incisive foramina small and elliptical; bulla? smaller; molars
rather narrow, but molar series long; zygomata short and not broadly
spreading. Compared with skulls of Z. trinotatus allcni fioni the Sierra
Nevada, Calif., it differs as follows: Smaller, but with molar series
longer; incisive foramina smaller and narrower posteriorly ; zygomata

2G NORTH AMERICAN FAUNA.

shorter and less spreading; brain case flatter; rostrum shorter. Com-
pared with the skull of the specimen from Mountain City, Nev., doubt-
fully referred to Z. oregomts, the type differs as follows : Slightly smaller
and younger; zygomata shorter and zygomatic breadth slightly less;
shape of brain case not strikingly different; molar series decidedly
longer; incisive foramina very much smaller and narrower posteriorly;
postpalatal notch indenting palate farther, nearly to middle of posterior
molars; bullaj slightly smaller. Mandible slighter, but lower molar
series longer.

Measurements. — Type: Total length, 242; tail vertebr.T, 150; hind
foot, 33, Skull (type): Basilar length, 19.4; zygomatic breadth, 12;
mastoid breadth, 11; interorbital constriction, 4.6; incisor to postpalatal
notch, 9.5; foramen magnum to postpalatal notch, 8; fronto-palatal
depth at middle of molar series, C.

General remarks. — The present form is based on a single adult speci-
men collected in the Ruby Mountains in east-central ;N"evada. It does
not seem to be closely related to any of the forms whose ranges partially
surround its habitat. While it is probably most nearly related to Z.
princeps, it seems to differ enough to warrant specific separation.

8x)eclmens examined. — The type.

ZAPUS TRINOTATUS Rhoads. Northwest Jumping Mouse.

(PI. I, tigs. 5, 5a.)

Zapus Irinotatus Rhoads, Proc. Acad. Nat. Sci. Phila., 1894, p. 421, Jau. 15, 1895.

Zapus imperator Elliot, Field ColuiubiaD Mus., Pub. 30, Zool. Ser., I, No. 11, -p. 228,

Feb. 1, 1899; ibid. No. 13, pp. 260-261, 1 fig. in text, March, 1899. (Type from

Sieg's ranch, Elwah River, Clallam County, Wash.)

Type locality, — Lulu Island, mouth of Fraser Eiver, British Columbia.

Geographic distribution. — Coast region of southern British Columbia,
Washington (including Cascades), Oregon (west of western base of
Cascades), and northern California, south to fJumboldt Bay.

General characters. — Size large; color bright; skull large and broad.

Color. — Summer pelage: Sides dark ochraceous-buff, rather heavily
lined with black-tipped hairs; dorsal area very distinct, moderately
flecked with color of sides; beneath white, frequently suffused or
blotched with fulvous, especially in young; outer surface of forearm
and legs dusky; tail quite strongly bicolored, in dried skins dusky
brown above and yellowish-white below. In early fall pelage the sides
are dull yellowish instead of ochraceous; dorsal area duller than in
summer, much more thickly flecked with yellowish hairs. In immature
individuals the dorsal area is especially indistinct.

Cranial characters. — Skull large and rather heavily built, about the
size of that of Z. xninceps. Brain case broader and more globular;
zygomatic breadth greater; palate shorter; molars more lightly built
than in that species.

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 27

Measurements. — Average of 4 adults from !N"eali Bay, Wash.: Total
length, 248; tail vertebrie, 153; hind foot, 33.5. Four specimens from
Lake Washington, Wash., average: Total length, 240; tail vertebme,
147.5; hind foot, 34.2. Ten specimens from Lake Cnshman, Wash.,
average: Total length, 239.5; tail vertebrti^, 144.5; hind foot, 32.5.
Sl'vU : Six adult skulls from five localities in Washington, average:
Basilar length, 20.2; zygomatic breadth, 13; mastoid breadth, 10.9;
interorbital constriction, 4.3; incisor to postpalatal notch, 9,0; foramen
magnum to postpalatal notch, 8.8; fronto-iDalatal depth at middle of
molar series, 6.7.

General remarls. — Zapus trinotatKs is one of the largest, and in early
summer pelage one of the most showy species in the genus. In the
absence of specimens from the exact type locality, sx^ecimens from Port
Moody, British Columbia, a few miles distant on the mainland, are
assumed to be typical.' The species is remarkably constant in char-
acters throughout its range, examples from near the type locality not
differing much from specimens from Yaquina Bay, Oregon. In the
Sierra Kevada of California this species is replaced by the closely
related subspecies aJleni. In a recent paper entitled ' Preliminary
descriptions of ]S^ew Eodents from the Olympic Mountains,' Mr. D. G.
Elliot has described the Zapus from that region under the name Zapus
imperator, but in making his comparisons he ignores Z. trinotatns, con-
trasting imperator only witli Z. prineeps. Through the courtesy of the
author I have been able to examine the type and a topotype of imper-
afor. They do not differ sufiSciently from Z. trinotatns^ from the con-
tiguous coast region and from the type locality, to admit of even sub-
specific separation.

Specimens examined. — Total number, 99, from the following localities:

British Columbia: Lulu Islaud (type locality), 2; Mount Baker Eange, 4;

Mouut Lebmau, 1 ; Port Moody, 3 ; Sumas, 3.
California: Crescent City, 1; Mad River (Carsou"s Carap), 1.
Oregon: Astoria,!; Beaverton, 3; Elk Head,!; Gieudale, 1; Lincoln County,

1; Marshfield, 1; Salem, 1; Yaquina Bay, 9.
Washington: Cascade River (head), 2; Easton, 3; Granville, 1; Fort Steila-

coom, 1; Kichelos Lake, 1; Lake Cusbniau, 15; Lake Washington, •!;

Lapush, 2; Mount Rainier, 12; Mount St. Helens, 3; Neah Bay, 5; Nis-

qually River, 4; Olympic Mountains (head Elwah River), 2-; Olympic

Mountains (head Soleduc River), 2; Pacific County, 5; Puget Sound, 3;

'Washington,' 1.

ZAPUS TRINOTATUS ALLENI Elliot. Allen's .lumping Mouse.

faptis alleiii PLlliot, Field Columbian Mus., Pub. 27, Zool. Ser., I, No. 10, pp. 212-213,
-March, 1898.

Type locality. — Pyramid Peak, Lake Tahoe, California.
Oeof/raphic distribution. — Mount Shasta and southward in the Sierra
Nevada to Mammoth and North Fork of Kern Eiver, California.

' Since the above was written the type and a topotype have been examined and
found to agree with the specimens from Port Moody.
''Type and topotype of Zcqjus imptrator.

2H NORTH AMERICAN FAUNA.

General characters. — Similar to Z. irinotatun; differing slightly in
external and cranial characters.

Color. — Early summer pelage: Sides rather pale ocbraceous-buff,
only moderately lined with black-tij^ped hairs; dark dorsal area well
defined, rather lighter than in Z. trinotatus; under parts pure white;
never marked with fulvous, except that rarely young specimens are
slightly sutiused with this color beneath; tail in dried skins rather
light grayish-brown above and yellowish- white below; tail lighter
toward tip, the pencil often white. Compared with Z. trinotatus, adults
of this species have dorsal area less flecked with color of sides and fewer
black-tipped hairs on sides. Upper surface of tail, ears, head, and
outer surface of thighs lighter than in Z. trinotatus. Immature indi-
viduals in late summer pelage have dorsal area very indistinct and
much mixed with ochraceous. Fall pehu/e : An adult taken at Cassel,
Calif., August 29 (very fat and doubtless about to hibernate), is in a
l^elage quite similar to that worn in earh' summer, but the sides are
lighter ochraceous, especially anteriorly, and the dorsal area is more
thickly flecked with color of sides. Z. alleni apparently does not assume
the dull yellowish fall pelage of Z. trinotatus.

Cranial characters. — Skulls similar to those of Z. trinotatus, but
smaller, with much smaller bulla'. From that of Z. montanus of the
Cascade Eange the skull of the present species differs so much as not to
require comparison.

Measurements. — Eight adults from Emerald Bay, Lake Tahoe, aver-
age: Total length, 234.6; tail vertebn^, 143.6; hind foot, 32.5. Two
adults from Mount Shasta, Calif., average: Total length, 249.5; tail
vertebrre, 152.5; hind foot, -32.5. Sl'ull : Eight skulls from Emerald
Bay, Lake Tahoe, average: Basilar length, 19.7; zygomatic breadth,
12.6; mastoid breadth, 10.5; interorbital constriction, 4.5; incisor to
postpalatal notch, 9.3; foramen magnum to postpalatal notch, 8.2;
fronto-palatal depth at middle of molar series, 6.~i.

General remarlcs. — Zapus t. alleni is a fairly well-marked subspecies
occupying the Sierra Nevada of California, from Mount Shasta south
to the region about the head of the North Fork of Kern River. It is
a very beautiful species and differs from all others of this subgenus in
often having the pencil, and occasionally more of the tail, white.

Specimens examined. — Total number, 61, from the following localities:

California: Big Trees, 1; Cassel, 1; East Fork Kaweah River, 2; Emerald
r)ay, Lake Tahoe, 19; Keru River (North Fork — 'Keru Lakes'), 1; Lassen
Peak, 14; Mammoth,!; Mount Shasta, 20; Summit, 1; Upper Bear Creek. 1.

ZAPUS MONTANUS (Merriam). Mountain Jumping Mouse.
Zapus Irinotatus montanus Merriam, Proc. Biol. Soc. Wash., XI, p. 104, April 26, 1897.

Type locality. — Crater Lake, Mount Mazama, Oregon.
Geographic distribution. — Cascade Eange in Oregon.
General characters. — Smaller and duller in color than Z. trinotatus^
also differing in cranial characters.

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 29

Color.— t<ummer pelage Sides rather dark, oebraceous-buff, heavily
lined with black-tipped hairs; dorsal area quite sharply defined, griz-
zled dusky and yellowish; outer side of legs dusky to heel; tail sharply
bicolor, dark gray above aud whitish beneath ; fore and hind feet soiled
white ; immature, suffused beneath with yellowish. Fall pelage : Dorsal
area darker than in summer, thickly flecked with light yellowish, pre-
seuting- a pepper-and salt appearance; sides yellowish-olive, heavily
lined with black-tipped hairs.

Cranial characters. — Skull smaller than those of Z. trlnotatus and
Z, princeps oregonus and much narrower throughout; incisive foramina
much narrower posteriorly; bulUe smaller.

Measurements.^-Eight specimens from tyj)e locality average: Total
length, 228; tail vertebrae, 135; hind foot, 31. SJcull: Three skulls
from type locality average: Basilar length, 19.5; zygomatic breadth,
12.1; mastoid breadth, 10.1; interorbital constriction, 4.5; incisor to
postpalatal notch, 9; foramen magnum to postpalatal notch, 8.3;
fronto-i)alatal depth at middle of molar series, C.l.

General remarks. — Zapus montanus is a very well-marked form inhabit-
ing the entire Cascade Range in Oregon. It is apparently totally dis-
tinct from all of the species whose ranges surround its habitat. The
type series was taken in August, 1896, near Crater Lake at the head
of a tributary of Anna Creek, on Mount Mazama, Oregon. The species
was quite abundant in the meadows and the shrubbery of the wet hill-
sides, and nearly a dozen were taken within a few days. Several of
their summer nests were found. A week or two later other specimens
were taken in the same valley a few miles below. These had begun to
assume the fall pelage, being slightly more olivaceous.

Two specimens taken September 9, at Fort Klamath, at the base of
the mountains, had assumed the complete fall pelage. They were
excessively fat, and were doubtless nearly ready to hibernate.

Specimens examined. — Total number, 16, from the following localities:

Oregon: Crater Lake (type locality), 9; Diamond Lake, 1; Fort Klamath, 2;
Mount Mazama (Anna Creek), 2; Mount Hood, 2.

ZAPUS ORARIUS sp. nov. Coast Jumping Mouse.

(PI. I, figs 4, 4a).

Type from Point Reyes, Calif., No. 250, ^ ad., collection of E. A. and O. Bangs. Col-
lected May 14, 1893, by Charles A. Allen. Original No. 618.

Geographic distribution. — Coast of California from Point Reyes north
to Mad River, Humboldt County; limits of range unknown.

General characters. — Size, medium; dorsal area and lower parts
strongly suffused with cttlor of sides. Skull rather small and peculiar
in shape.

Color. — Type in rather loorn spring pelage: Sides of body and head
rather dark ochraceous, moderately lined with black-tipped hairs;
dorsal area not sharply defined and strongly suffused with color of

30 NORTH AMERICAN FAUNA.

sides. White of lower parts so strongly suttased with ochraceous
that the white is almost obsolete, appearing only on lower parts of
forelegs and between thighs; hind legs ochraceous all around; sides of
throat deeper ochraceous than remainder of lower parts; feet yellowish
white; tail yellowish-white below and grayish above. A specimen
from Point Eeyes, in the collection of Dr. G. Hart Merriam, about one-
fourth grown, " has the upper parts almost uniform deep ochraceous
yellow, with only the faintest trace of the dorsal area."

Cranial characters. — Skull rather small; rostrum short and consider-
ably deflected; nasals very narrow anteriorly; greatest zygomatic
breadth on plane of middle molars; zygomata rather light, the maxil-
lary portion meeting nearly at right angles to main axis of skull; pro-
cess on lower border of maxillary arm of zygoma very pronounced and
nearer antorbital foramen than in any other species. Brain case high
and rather globular in shape; interorbital constriction narrow; inter-
pterygoid fossa narrow and shallow; incisive foramina rather small
and evenly elliptical in shape; audital bulhe small and rather near
together; upper incisors slender and more projecting than usual; man-
dibular symphysis short.

Measurements. — The type measures: Total length, 220; tail vertebrte,
127; hind foot (dry), 30. Sl'uU [ty\)e): Basilar length, 19; zygomatic
breadth, 12; mastoid breadth, 10.6; interorbital constriction, 1; incisor
to postpalatal notch, 8.5; foramen magnum to jjostpalatal notch, 7.1);
fronto-palatal depth at middle of molar series, 0.2.

General remarls. — Zapus orarins appears to be a very well-marked
species requiring close comparison with no other known form. The
type has the lower parts more intensely suffused with the color of the
sides than any other specimen of the genus that I have seen. A speci-
men from Mad Kiver, Humboldt County, resembles the type very closely,
and another from Eureka, without a skull, undoubtedly belongs to this
species. The latter is suffused beneath with ochraceous, though to a
lesser extent than the type. The skull of the type bears a slight resem-
blance in some respects to that of Z. imcijicus^ but the differences are
so great that unless the type specimen is abnormal orarins can not be
considered closely related to pacijicus. Much additional material is
needed to clear up satisfactorily the relationships of these and other
forms from this region.

Specimens examined. — Total number, 4, from the following localities:

California: Eureka, 1; Mad River (Carson's Camp), 1; Point Reyes (type
locality), 2.

ZAPUS PACIFICUS Merriam. Pacific Jumpinj;- Mouse.

«
Zapus pacijicus Merriam, Proc. Biol. Soc. Wash., XI, p. 104, April 26, 1837.

Type locality. — Prospect, Rogue River Valley, Oregon.
Geographic distribution. — Interior valleys of southwestern Oregon
and northwestern California; limits of range unknown.

REVISIOX OF THE JUMPING MICE OF THE GENUS ZAPUS. 31

Color. — " Dorsal area not sharply defined, but so strongly suffused with
yellowish that the yellow predominates over the black; sides buffy-
yellow, moderately lined with black hairs; inner side of legs only
slightly darkened; tail sliarply bicolor; grayish above, white beneath;
fore and. hind, feet soiled white." ' A topotype, younger than the type
from which the above description was taken, "has the dorsal area even
less distinct, the entire upper parts being ochraceous yellow."

Cranial characters. — Compared with that of Z. montanns (its nearest
relative geographically) the skull oi Z. pacificusi^ smaller; the rostrum
and nasals shorter; audital bulla' smaller; basi-occipital broader between
bullae; inter pterygoid fossa shorter; u[)per molar series more divaricat-
ing posteriorly.

Measurements. — The type measures: Total length, 225; tail vertebrjie,
14:1; hind foot, 31. SJcull : Basilar length, 17.5; zygomatic breadth,
12; mastoid, breadth, 10; interorbital constriction, 4.5; incisor to post-
palatal notch, 8.4; foramen magnum to postpalatal notch, 7.2; fronto-
palatal depth at middle of molar series, 0.

General remarlxs. — Owing to the small number of specimens available
from the immediate region, it is imi^ossible to discuss satisfactorily the
true relationship of Zajms pacijicus to the surrounding species. The type
and a topotype taken at the same time are in a i^eculiar washed-out,
almost albinistic, i»elage, and may possibly be abnormal, esijecially as
the locality is well within the humid, heavily foj-ested area, where most
of the animals are darker in color than their congeners to the eastward.
The species, however, has fairly well marked skull characters and is
undoubtedly a good form. An immature specimen from Siskiyou, Oreg.,
taken September 28, doubtless referable to this form, has dorsal area
more distinct; sides brighter ochraceous and with more black hairs;
upper surface of tail darker and fur of upper parts darker at base than
specimens from the type locality. Another, from Little Shasta, Calif.,
taken September 20, also provisionally referred to this form, resembles
the Siskiyou specimen, but has dorsal area finely flecked with yellowish
instead of ochraceous.

Specimens examined. — Total number, 4, from the following localities:

Oregon: Prospect, Rogue River Valley (tyjie locality), 2; Siskiyou, 1 (not

typical).
California: Little Shasta, 1 (not typical).

ZAPUS SALTATOR Allen. Stickeeu Jumping Mouse.
Zapus saltator Allen, Bull. Am. Mus. Nat. Hist., N. Y., XII, p. 3, Mar. 4, 1899.

Type locality. — Telegraph Creek, Northwest Territory, Canada.

Geographic distribution. — Telegraph Creek south to mouth of Skeena
Kiver and Tschimshian Peninsula; limits of range unknown.

General characters. — Similar in early fall pelage to Z. trinotatus, but
smaller and differing in cranial characters.

' From original description.

32 NORTH AMERICAN FAUNA.

Color. — Fall pelade: Sides yellowish, moderately lined with black-
tipped hairs; dorsal area well-defined, rather dark, and thickly flecked
with yellowish ; ears like back, narrowly edged with yellowish ; beneath,
pure white; tail dusky above and gray beneath. Compared with Z.
trinotatiis in corresponding pelage, the back is slightly darker, the tail
darker above, and gray, instead of yellowish- white beneath.

Cranial characters. — Compared vfith Z. trinotatus, the skull of Z. sal-
<rt<o>- differs as follows: Brain case narrower; zygomatic breadth con-
siderably less; nasals broader i^osteriorly. Compared with skulls of
Z. prince]) s from Field and Glacier, British Columbia, and Heniy House,
Alberta, the skull of Z. saltator has the rostrum longer; palate from
incisive foramina to postpalatal notch shorter; and molars, especially
last upper, smaller. All of the skulls of Z. saltator examined agree
very well among themselves, and all have the incisive foramina very
large and broad posteriorly. The zygomatic exj)ansion is sniaJl and
the brain case high and narrow.

Measurements. — An adult from Port Simpson, British Columbia,
measures: Total length, 245; tail vertebrne, 145; hind foot, 32. Average
of five young adults from same locality: Total length, 234; tail verte-
brae, 151; hind foot, 32. Skull: An adult skull from Port Simjisou,
British Columbia, measures: Basilar length, 20; zygomatic breadth,
11.8; mastoid breadth, 10.5; interorbital constriction, 4.5; incisor to
postpalatal notch, 9.4; foramen magnum to postpalatal notch, 8; fronto-
palatal depth at middle of molar series, 6.4.

General remarks. — Zapus saltator is a well-marked form, differing con-
siderably from its relatives to the southward. With Z. hudsonius alas-
eeiisis, which probably meets it on the north, it re(iuires no comparison,
the immensely larger incisive foramina and larger molars of the pres-
ent species distinguishing the two forms at a glance. A small series
of Z. saltator, which forms the basis of the present description, was
taken by the writer at Port Simj^son, British Columbia, in August,
1897. Some of these were taken in a grassy thicket near the edge
of the forest, a few feet above high-water mark, and the remainder in
a garden in the village. Z. saltator may be found to mtergrade with
either Z. trinotatus or Z. princeps (it seems nearer the latter), but until
its true relationship is proved it seems best to allow it full specific rank.
The tail is longer and the hind foot shorter, relatively, than in Z. trino-
tatus. Through the kindness of Dr. J. A. Allen, J have been enabled
to examine the type of Z. saltator. The skull of the type being imper-
fect, the cranial characters given above are taken from Port Simpson
specimens.

Specimens examined. — Total number, 8, from the following localities:

Northwest Territory: Telegrapli Creek, 1 (the type).

British Columbia: luverness (mouth of Skeena River), 1 ; Port Simpson, 6.

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 33

NAP^^OZAPUS^ subgenus novum.

Tjiije Zapus iiisigtils Miller, from Restigouche River, New Brunswick.

Suhgeneric characters. — Teeth: i.\'^, p.^^^, m.^-=16. Skull stouter
than in other subgenera; interorbital constriction
greater; frontal region more swollen ; middle molars
equaling first in size. Enamel pattern of molars
not essentially different from Zapus, but sulcus ou
inner side of upper molars deeper and persisting
longer in the wearing tooth (fig. 2). General color ^^^ ^ _,^^,^^ ^^^^^ ^^ ^„.
pattern as usual in the genus, bat tail tipped with ims (Napoeozapus) insig-

, .. nis (topotype), from

" " ' '-*'• Restigouche River, N. B.

CNo. 2333, coll. E. A. aud
O. Bangs) X 6.

KEY TO SPECIES OF THE SUBGENUS XAP.EOZAPUS.

Hind foot less than 33 mm. ; interorbital constriction usually more than 4.5 mm.
Larger and lighter in color; hind foot usually more than 30 mm. ; skull broader

and stouter insi(iii is.

Smaller and darker; hind foot about 30 mm. or less; skull rather slender.

roanensis.

Hind foot about 33 mm. ; interorbitiil constriction narrow, usually less than 4.5 mm. ;

size large ahietorum.

ZAPUS (NAP.^OZAPUS) INSIGNIS Miller. Woodland Jumping Mouse.

(PI. I, figs 1,1a.)

1856. Meriones labradorius Dawson, Edinburgh New Philos. Journ., new ser., Ill, p. 2.

1857. Jacitlus hiidsonius Baird, Mamm. N. Am., p. 430 (in part — specimen from Nova
Scotia).

1877. Zapus Inidsonius Coues, Mon. N. Am. Rodentia, p. 467 (in parf).
1891. Zapus insignis Miller, Am. Naturalist, XXV, p. 472, August, 1891.

Tyj)e locality. — Eestigouche River, New Brunswick,

Geographic (listribution. — Canadian Zone in eastern Canada and south
to western Maryland,

General characters. — Size rather large, larger than Zapus hudsoniiis^
with longer ears and paler, more fulvous coloration. Tail tipped with
white.

Color. — "Adult male No. if f f? collection of G-, S, Miller, jr., Peterboro,
N. Y., August 22, 1892; length, 250; tail vertebrfie, 154; hind foot, 31,6;
ear from notch, 18,6. Tij) of tail for 23 mm., dorsum of ruanus and pes,
and entire ventral surface pure white to base of hairs. Sides buft'-yellow,
tinged with clay color, except on cheeks, fore neck, and a narrow line
bordering white of belly, where the yellow is noticeably purer; the fur
plumbeous-gray at base and a trifie sprinkled with black, bristly hairs.
These black hairs predominate on the back, where they form a sharply

^ Xapo'ozajyus : 7^a7r azo?, belonging to a wooded vale or dell; -\- Zapus. This name
was suggested to me by Mr. Gerrit S. Miller, jr.
22095— No. 15 3

34 NORTH AMERICAN FAUNA.

defined dorsal stripe slightly mixed with the color of the sides, broad-
est just back of the shoulders, tapering gradually to base of tail, and
becoming indistinct on the head after passing between the ears. Ears
externally concolor with back, internally bufl'-yellow; muzzle grayish-
browu; whiskers mixed brownish and whitish, the longest hairs reach-
ing beyond shoulders; tail thinly haired, so that the aunulation shows
distinctly, sharply bicolor, dark brown, except ventrally and at tip."'
There is very little seasonal variation in this species, and the white of
lower parts is never suffused with the color of sides.

Cranial characters. — Skull rather broad aud stout; frontal region
much swollen; brain case broad and rather flat; interorbital constric-
tion broad as compared with that of all other species of the genus.

Measurements. — Nine specimens from Nova Scotia average: Total
length, 238; tail vertebra?, 146.7; hind foot, 31.1. Sixteen from Peter-
boro, N. Y., average: Total length, 236; tail vertebme, 147; hiiul foot,
30.25. One from Pocono Mountain, Pennsylvania: Total length, 224;
tail vertebme, 140; hind foot, 31. Two from Lake Hopatcong, New
Jersey, average: Total length, 231; tail vertebme, 140; hind foot, 31.5.
Sl'ull: Two skulls from type locality average: Basilar length, 18.7;
zygomatic breadth, 12.3; mastoid breadth, 10.2; interorbital constric-
tion, 4.9; incisor to postpalatal notch, 8.8; foramen magnum to post-
palatal notch, 7.8; fronto-palatal depth at middle of molar series, 6.
Four skulls from New York average: Basilar length, 19.6; zygomatic
breadth, 12.9; mastoid breadth, 10.8; interorbital constriction, 5; incisor
to postpalatal notch, 9; foramen magnum to postpalatal notch, S.6;
fronto-palatal depth at middle of molar series, 6.1.

General remarks. — Apparently the first published record which refers
unmistakably to this species is that of Prof. J. W. Dawson, who, in
1856, in the Edinburgh New Philosophical Journal, called attention to
the fact that two distinct species of Jumping Mice inhabited Nova
Scotia. The present species he wrongly referred to Meriones labradorius
(= Zapus hudsonius), and described true Z. liudsonius as a new species,
calling it, provisionally, Meriones acadiciis. In the National Museum
are two specimens of Zapus deposited by Dawson ; though mounted, they
are in good condition for examination and are plainly referable respec-
tively to Z. insignis and Z. hudsonius. Both Baird and Coues examined
them, or at least similar specimens received from Dawson, but did not
consider the two animals specifically distinct. Zapus hudsonius also
occurs throughout the range of the present species, but the two prefer
different situations — Z. hudsonius, shrubby fields and meadows, and Z.
insignis, deep woods near streams.

This species presents remarkably little variation throughout its range.
Si^ecimens from central Pennsylvania, in the collection of Mr. S. N.
Khoads, are practically indistinguishable from examples taken near
the type locality. Three specimens from Lake Hopatcong, N. J., have

1 G. S. Miller, jr., Proc. Biol. Soc. Wash., YIII, p. 2, 1893.

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 35

sligbtly smaller skulls than usual, but are otherwise perfectly typical.
I have also examined two specimens from western Pennsylvania, col-
lected by Mr, W. E. Clyde Todd, and have myself collected a speci-
men in the mountains at Finzel, in Garrett County, Md. jSToue of
these show any approach to Z. insignis roanensis from Eoan Mountain,
North Carolina. Mr. S. X. Ehoads writes me that he has also taken
the species at Summit, Cambria County, Pa.
Specimens examined. — Total number, 107, from the following localities:

Ne'w BrunsAwick : Restigoucbe Eiver (type locality), 3.

Nova Scotia: Halifax, 3; James River, 4.

Ontario : North Bay, 6.

New Hampshire: Antrim, 1; Chocorua, 8; Dublin, 3; Fabyaus, l; Fraiico-

nin, 4; Profile Lake, 10.
Vermont : Mount Mausfielcl, 4.
Massachusetts : Mount Greylock, 1.
New York: Cascacleville, 4; Catskills, 4; Elizabetbtown, 10; Glenville, I;

Lake George, 1 ; Peterboro, 30.
New Jersey: Lake Hopatcoug, 3.
Pennsylvania: Cherry Sjiring, Potter County, 1; Eaglesmere, 2; Kingston,

Westmoreland County, 1; Mount Pocono, 1.
Maryland: Finzel, Garrett County (6 miles north of Frostburg), 1.

ZAPUS (NAP.EOZAPUS) INSIGNIS ROANENSIS subsp. nov. Roan Mountain

Jumping Mouse.

Type from Magnetic City, foot of Roan Mountain, North Carolina, No. 66283, $ ad.,
U. S. Nat. Museum, Biological Survey Coll. Collected May 22, 1894, by A. G.
Wetherby.

Geographic distribution. — Known only from Roan Mountain, North
Carolina.

General characters. — Smaller and darker than typical Z. insignis, with
smaller, narrower skull.

Color. — Sides bright tawny ochraceous ; entire upi^er iiarts, including
ears, considerably darker than in typical Z. insignis. Beneath, pure
white; amount of white on tail averaging less than in Z. insignis.

Cranial characters. — Skull similar to that of Z. insignis, but smaller
and more slender throughout; mandible much more slender, especially
in region of molars; interorbital constriction very slightly less; frontal
region much less swollen; brain case narrower; fronto palatal depth
about the same.

Measurements. — Sixteen sj^ecimens from Eoan Mountain average:
Total leugth, 220; tail vertebne, 131 ; hind foot (dry), 29.5. Sl^uU: Six
adult skulls from Eoan Mountain average: Basilar length, 18.8; zygo-
matic breadth, 11.7; mastoid breadth, 10.1; interorbital constriction,
4.7; incisor to postpalatal notch, 8.8; foramen magnum to postpalatal
notch, 8; fronto palatal depth at middle of molar series, 6.2.

General remarl^s. — The collection of the Biological Survey contains
24 specimens of this form, collected from May to September. As in
typical Z. insignis. there is very little seasonal variation. A young
individual, about one-third grown, collected at the type locality Sep-
tember 11, 1892, by Dr. C. Hart INIerriam, resembles adults in color,

36 NORTH AMERICAN FAUNA.

except that the dorsal area is very iDdistinct aud the nose is white.
Several of the type series, including this young one, have the white
on the tip of tail reduced to the merest trace. While the material
examined from Maryland and Pennsylvania shows no indication of
iutergradation existing between typical Z. insigjiis aud the present
form, I have thought best to give it a trinomial name in the belief
that iutergradation does exist.

Sx^ecimens examined. — Total number, 24, all from Roan Mountain,
North Carolina.

ZAPUS (NAPiEOZAPUS) IXSIGNIS ABIETORUM subsp.uov.
Northern Woodland Jumping Mouse.

Type from Peninsula Harbor, north shore of Lake Superior, Ontario, No. 4268, J ad.,
collection qf Gerrit S. Miller, jr. Collected Sept. 27, 1896, by Gerrit S. Miller, jr.

Geograpliic distribution, — Probably throughout Hudsonian zone in
eastern Canada; limits of range unknown.

General characters. — Larger than typical Z. insignis, with shorter
ears aud peculiar skull.

Color. — Ai)parently not distinguishable from Z. insignis.

Cranial characters. — Compared with typical Z. insignis the skull of
the type has molar series longer and heavier; interorbital constriction
narrower than in any S[)ecimen of insignis examined; ventral border of
l^terygoids nearly on plane of palate; maxillary process of zygomata
shorter and zygomata much less curved than in typical insignis;
mandible much heavier.

Measurements. — Type: Total leugth, 1*55; tail vertebne, 160; hind
foot, 33; ear from meatus, 16.6. An adult male from Godbout, Quebec,
measures: Total leugth, 250; tail vertebne, IGO; hind foot, 32.5. iSJcull
(type): Basilar leugth, 19.1; i^ygomatic breadth, 12.2; mastoid breadth,
10.5; interorbital constriction, 1.3; incisor to postpalatal notch, 9.6;
foramen maguum to postpalatal notch, 8.4; fronto-palatal depth at
middle of molar series, 6.

General remarlcs. — This form is based mainly on a single specimen,
collected at Peninsula Harbor, Ontario, on the north shore of Lake
Suj^erior, in the Hudsonian zone, by Gerrit S. Miller, jr. Mr. Miller
recorded this specimen ' and described its peculiarities in some detail.
A specimen in the Bangs collectiou, from Lake Edward, Quebec, ami
two from Godbout, Quebec, in Dr. Merriam's collection, while not per-
fectly tyjtic'al, agree with the type in large size, narrow interorbital
coust'riction, and in other peculiarities. This makes it probable that
this form .will be found to occur throughout the Hudsonian zone, in
eastern Canada. The specimen of Z. insignis, recently recorded from
Labrador by Mr. Outram Bangs, ^ which he thinks is not typical, may

' Notes on the Mammals of Ontario <;Proc. Boston Soc. Nat. Hist , XXVIII, No. I,
p. 10, April, 1897.

"Am. Naturalist, XXXII, No. 379, p. 193, July, 1898.

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 37

belong to this form. The type of abietonim is slightly larger than the
largest specimens examined of the typical form, while the ear is slightly
shorter than usnal.

Specimens examined. — Total number, 4, from the following localities:

Ontario: Peuinsula Harbor, 1 (type).
Quebec: Godbout, 2; Lake Edward, 1.

EOZAPUS^ subgenus novum.

Type Zaptis setchuanus Pousargues, from Szechuen, Chiua.

Suhgeneric characters. — Teeth: i. |, j;. l^l, m. ^ = 18. Skull similar
to those of the subgenera Zapiis and Xa2)a'o.zapKS, but differing as fol-
lows: Skull relatively broader, shorter, and lower; nasals broad, dis-
tinctly forked posteriorly; antorbital foramina rather large; incisive
foramina medium; posterior palatine foramina mid-
way between incisive foramina and postpalatal
notch, the latter barely reaching posterior plane
of molars. Rostrum relatively shorter and broader;
zygomata relatively shorter, Mith jugal portion
almost perfectly straight, frontal region less swol-na. 3.— Molar teeth of
len. Enamel pattern of molars somewhat similar, ^^*"" (fozapm} setch-

^ > uamts (cotype), troui

but enamel folds not closely crowded; last lower xa-tsien-iou.Szeciiuen,
molar relatively larger (fig. 3). Color pattern differ- ^'i'°='- ^ ^•
iug from that of other subgenera in having i\ brown ventral stripe; tail
tipped with white, as in Napceozapus.

ZAPUS (EOZAPUS) SETCHUANUS Pousargues. Szechuen Jumping Mouse.

(PI. I, tigs. 2, 2a.)

Zapus setchuanus Pousargues, Anu. Sci. Nat., S" ser., I, No. 4, p. 220, Avril, 1896; Bull.
Mus. d'llist. Nat., Paris, II, p. 13, figs. 1-3, 1896.

Type locality. — Ta-tsien-lou, Szechuen, China.

Geographic distribution. — Known only fiom the type localit3\

General characters. — Differing from all other known members of the
genus in having a ventral stripe of brown.

Color. — Upper parts dark, talwny ochraceous; dorsal area sharply
defined and very thickly tiecked with black-tipped hairs, the sides mod-
erately lined with same. Pattern of color above as iu the American
species. Beneath white, suffused with ochraceous, the white nearly
obsolete on fore breast; a brownish stripe about 5 mm. wide extends
the entire length of breast and belly; feet, chin, and lips white; tail
strongly bicolored; brownish above and whitish beneath, with about
15 mm. of the tip pure white; ears dusky. »

Cranial characters. — Compared with Z. hi(dsonii(s the skull of setchu-
«n?<s differs as follows: In terorbital constriction relatively narrower;

'Eozapus: i)d}<i, dawn, /. £. eastern; -\-Zapus.

38

NORTH AMERICAN FAUNA.

Tig. 4.— Skull of Zapu&
(Eozapus) setchuanus (co-
type) from Ta-tsien-lou,
Szechuen, China. X IJ.

frontal region less swollen; rostrum shorter and less attenuated; zygo-
uiata shorter, tlie maxillary portion more nearly iierpendicular to main
axis of skull, tlie jugal portion heavier and nearly straight; palate
longer, the postpalatal notch not quite reaching
posterior plane of molars (in American S])ecies
commonly exceeding that point) ; nasals prolonged
farther posteriorly and forking (in American spe-
cies generally irregularly truncated) ; bulla? larger;
the coronoid process of mandible not ascending
so high, but longer, broader, and straighter, and
the coronoid notch deeper and less rounded than
in Z. hudsoniu.s (fig. 4).
Measurements. — The following measurements from Pousargues will
serve for comparison with other species : ' Vieux,' head and body, 100;
tail, 120; foot, 31; ' adulte,' head and body, 80; tail, 103; foot, 30;
'semiadulte,' head and body, 70; tail, 95; foot, 28. Skull No. 2:^
Zygomatic breadth, ll.G; interorbital constriction, 3.G; incisor to post-
palatal notch, 8.5; fronto-palatal depth at middle of molar series, 5.5.
No. 3: Zygomatic breadth, 11.5; interorbital constriction, 3.8; incisor
to postpalatal notch, 8.2; fronto-palatal depth at middle of molar
series, 5.G.

General remarks. — This very interesting species was described from
three skins in alcohol accompanied by imperfect skulls. Its describer,
M. 1]. De Pousargues, discusses its color and other peculiarities in con-
siderable detail, and presents a table showing the measurements of tlie
three specimens, together with the corresponding measurements of a
specimen of Z. hudsonius, taken from Coues. These measurements of
hudsonius (head and body, 85; tail, 135; foot, 27), if all from one indi-
vidual, must have been taken from a distorted specimen and do not
give a fair idea of the proportions, the foot measurement, 27 mm.,
being evidently too small for an animal with the tail measuring 135.
On this account the proportional differences between setclmanus and hud-
sonius are exaggerated. Through the courtesy af Mr. Gerrit S. Miller,
jr., I have been able to examine two of these skins and the skulls, gen-
erously loaned to him by M. Pousargues. The color of these skins is
apparently not much changed by the action of alcohol. The brown
ventral stripe and comparatively short, hairy tail with its white tip,
constitute the most striking peculiarities. The hind foot is about the
same size as in hudsonius; the ear shorter and broader than in the
American species.^ As already stated, the skulls are all imperfect.

^ These numbers are provisional, and owing to the imperfect condition of the skulls
only these measurements cbuld be taken.

- The following table will serve to show the approximate ratio of width to height
of ear in Zaptis insUivis, Z. hudbomus, and Z. seickuanus (alcoholic specimens) :

Z. insignis.
Two specimens.

Z. hudsonius.
Two specimens.

Z. setchuanus.
Two specimens.

Male.

Female.

Male.

Female.

Larger.

Smaller.

Height of ear from uotcli

15.5

15.5
9

14
9

14
9

12.5
10

11

Greatest width of ear

9

REVISION OF THE JUMPING MICE OF THE GENUS ZAPUS. 39

The largest is so badly crushed as to be almost worthless, aud the two
smaller ones lack the occipital portion of the cranium and the posterior
parts of the mandibles, the shape of the angular process, therefore,
being unknown. This species has the small upper premolar as in the
subgenus Zapus, that tooth being larger than in Z. hudsonius. The last
lower molar is also proportionately larger. All the molars differ from
those of any American species in having the enamel folds not closely
crowded, the resulting wide and deep sulci giving the teeth a very
different appearance, though the general pattern of enamel folding is
perhaps not essentially different. The incisors are lighter in color
than in the American species.

Specimens examined. — Two skins and • three skulls, from the type
locality.

I.:^DEX.

[New naniea in blaokface type; synoiiyniM in italics.]

Dipui. 9, 13.

americaniis, 10, 19.

canadensis, 10, 15.

hudionius, 11, 15.

labradorius. 11, 15.

mcllivorus, 11-12.
Eozapii!!) (.subgenus), 37.
Qerhillm, 9, 13.

brachyurus, 10.

canadensis, 15.

daviesii, 10.

lahradoriiift, 1.3.

leomtrus, 11.

macrourtts, 11.

megalops. 11.

niger, 12.

soriciniis, 12.

(■■ylvaticxi.s, 12.
Jaculun, 10, 13.

aiaericanvx, 15.

hudsonius, 15, 33.

labradorius, 15.
ileriones, 9, 13.

acadicxcs, 10, I.t.

labradorius, 15,33.

microceplialus, 12, 15.

ncmoralis, 12.
J/i(« lonrjipc.s. 11.
IVa|ia>ozapus (subgeuu.s), 33.

key to species, 33.
Zapus, 10, 13, 14.

distribution, 6-7.

22095— No. 15-

Zapus — Continued —

habits, 7-9.

historj", 5-6.

key to species, 14-15.

key to subgenera, 13.

list of species, 13-14.
Zapus alleni, 10, 27.

hudsonius, 15-17, 33.

hudsonius alasceu.sis, 10, 18-19.

hudsonius ainerieanus, 10, 19-20.

hudsonius CHin|»estri.s, 20-21.

hudsonius caiiadensin, 15.

hudsonius hardi/i, 10, 15.

hudsonius ladas, 11, 17-18.

imperator, 11, 26.

insignis, 11, 33.

iiisigiii!!! abictoi'iiiiB, 36-37.

iiisigiti.<< roaiieiiMiH, 35-36.

major, 24-25.

moutanus, 12,28-29.

■icvadeiisis, 25-20.

orariiiss, 29-30.

paciflcus, 12, 30-31.

priuceps, 12, 22-23.

priiicrps minor, 23.

princep!< orrg;onus, 24.

saltator, 12, 31-32.

setchuanus, 12, 37-39.

teuelhis, 12, 21.

trinotatus, 12-13, 26-27.

trinotatus alleni, 27-28.

41

PLATE I.

fOne and one-half times natural size.)

Fig. 1. Zapus (Xainrozaj^ns) insiinii-s Miller (Type). Restigoitclie River, New Bruns-
wick.

(Type Is J, Miller collection.)

2. Zapus {Eozapus) setchuaniis Pousargue-s (Cotype). Ta-tsien-lou, Szechueu,

China.

3. Zapus {Zapus) hudsonius (Zimmermann). James Bay, Canada.

(No. 60588, U. S. Nat. Mas.)

4. Zapus {Zapus) orarius Preble (Type). Point Reyes, Calif.

(No. 250, Collection E. A. & 0. Bangs.)

5. Zapus {Zapus) irinotatus Rlioads. I'ort Moody, British Columbia.

(No. 66928, U. S. Nat. Mus.)

6. Zapus {Zapus) princeps Allen (Topotype). Florida, La Plata County, Colo.

(No. 4139, Am. Mus. Nat. Hist.)

42

o

North Amencan Fauna No. 15

Plate

Skulls of Zapus.

1. Zapus insignis.

2. Z. setckuanus.

■i. Z. Iindsonius.
4. Z. orarius.

5. Z. trinotatiis.
(J. Z. princeps.

U. S. DEPARTMENT OF AGRICULTURE

DIVISION OF BIOLOGICAL SURVEY

NORTH AMERICAN FAUNA

:no. 16

[Actual date of i»ublicatioii, October 28, 1899]

RESULTS OF A BIOLOGICAL SURVEY OF MOUNT SHASTA
CALIFORNIA

BY

C. HART MERRIAM
CHIEF OF DIVISION OF BIOLOGICAL SUKVEY

WASHINGTON

GOVERNMENT PKINTING OFFICE
189 9

<1_

LETTER OF TRANSMITTAL.

TJ. S. Department of Agriculture,

Division of Biological Survey,

Washington, D. 0., May 19, 1899.
Sir: I liave the lionor to transmit herewith for publication, as Xorth
American Fauna No. IG, a report on the results of a Biological Survey
of Mount Shasta, California, made during the summer of 1898.
Eespectfully,

C. Hart MerriaMj
Chief, Biological Survey.
Hon. James Wilson,

Secretary of Agriculture.

CONTENTS.

Pagb.

Introduction 9

Itinerury 10

Persouuel 15

Previous pul)lications 15

New species 16

General features 17

Effects of scanty moisture 19

Glacial basins — , 20

Canyons 21

Streams 23

Rock slopes 25

Avalauches 26

Timbcrliue 27

Forests of Shasta 30

The yellow pine belt 30

The Shasta fir belt 36

The Avhite-bark pine belt 39

Forest iires 46

Eiiects of Inirus in changing zone positions 47

Slope exposure 47

Ert'ects of steep slopes 51

Basin slopes 52

Life zones of Shasta 52

Upper Sonoran zone 53

Transition zone 54

Canadian zone 61

Hudsonian zone 64

Alpine zone 67

The Boreal fauna and flora of Shasta contrasted with corresponding- faunas and

floras of the Sierra and the Cascades 69

Efficiency of Klamath Gap as a barrier to Boreal species compared with that

of Pitt River and Feather River gaps collectively 83

Sources of the Boreal faunas of Shasta and of the Sierra and the Cascades ... 85

Mammals of Shasta 87

Birds of Shasta and vicinity - 109

Notes on the distribution of Shasta plants 135

5

ILLUSTRATIONS.

PLATES. Facing

page.

Plate I. Mount Shasta from the west Frontispiece.

II. Shasta from northwest, showing great snowbank in head of Diller

Canyon 18

III. Southeast slope of Shasta, showing canyons of Mud Creek and

Clear Creek. (Kindness of J. S. Diller) 22

IV. Tongue of dwarf white-bark pines at extreme upper limit of tim-

berline. (Kindness of J. S. Diller). 28

V. Pine forest at west base of Shasta, showing yellow and sugar pines.

(Kindness of J. S. Diller) 30

TEXT FIGURES.

rage.

Fig. 1. Wagon Camp 10

2. Shasta from east brink of Mud Creek Canyon 11

3. Shasta from Inconstance Creek, near timberliue on north side 12

4. Shastina from north, showing great bank of snow which feeds Shas-

tina streams 13

5. Alpine hemlocks, Squaw Creek Camp 13

6. Heather meadow on Squaw Creek 14

7. Pumice sand strewn with gray volcanic shale. Young hemlocks in

foreground ; white-bark jiines in distance 18

8. Heather meadow bordering Squaw Creek. Shasta peak in distance

covered with fresh snow, September 22, 1898 19

9. Glacial meadow at head of Squaw Creek 21

10. Heather meadow on upper Squaw Creek, showing concentration of

vegetation near stream 23

11. Miniature cascade on upper Squaw Creek, frequented by water shrews

and ouzels 24

12. Characteristic rock slope on north side of Shastina 25

13. Track of avalanche invading forest of Shasta firs on Cold Creek, east

side of Shasta 27

14. High timberliue ridge, showing eft'ects of slope exposure. (Photo-

graphed by John H. Sage) 29

15. Manzanita chai)arral on south slope of Shasta 31

16. Cones of ponderosa and Jeffrey pines 32

17. Knobcone pine on Panther Creek 33

18. Incense cedar on south slope below Wagou Camp 35

19. Cone scales of Jhies shastensis and Abies concolor lowiana 37

20. White-bark pine 38

21. Dwarf white-bark pines on a high ridge 39

22. Pumice plain north side of Shasta, showing timberliue mats of

white-bark pines 40

7

O NORTH AMERICAN FAUNA. [no. 16.

I 'age.

Fig. 23. A large prostrate tree of white-bark pine, a little below timberliue . 41

24. Black alpiae hemlocks near Squaw Creek 42

25. Group of alpiue hemlocks near Deer Canyon 43

26. Trunk of alpine hemlock, showing thickening and curvature of base. 44

27. Group of alpiue hemlocks 45

28. Diagram showing average hourly march of temperature 48 ,

29. Dwarf jjines ending abruptly along cold east sule of ridge .50 i

30. Diagram of Shasta showing relations of life zones 52

31. Winter earth plugs of pocket gopher 96

32. Mound made by pocket gopher, (Photographed by W. H. Osgood). 96

33. Rock cony. (Photograplied by F. Stephens) 99

34. Miuk. (Drawn by Ernest S. Thompson) 105

35. Marten. (Drawn by Ernest S. Thompson) . 106

36. Red-tail. (Drawn by J. L. Ridgway) 112

37. Sparrow hawk, (Drawn by J. L. Ridgway) 113

38. Arctic three-toed woodpecker. (Drawu by L. A. Fuertes) 115

39. Clark crow. (Drawu by L. A. Fuertes) 120

40. Clark crow. (Photographed by Walter K. Fisher) 121

41. Western evening grosbeak. (Drawn by L. A. Fuertes) 122

42. White-crowued sparrow. (Drawn by L. A. Fuertes) 125

43. Audubon warbler. (Drawn by L. A. Fuertes) 129

44. White hellebore. (Photographed l)y Walter K. Fisher) 14C

45. Manzanita chaparial 151

46. Monument on summit of Shasta. (Photographed by W. H. Osgood. 16i

No. 16. NORTH AMEEIOAN FAUNA. October, 1899.

RESULTS OF A BIOLOGICAL SURVEY OF MOUNT SHASTA,
NORTHERN CALIFORNIA.

By C. Hart Merriam.

INTRODUCTION.

At the close of tlie field season of 1897 the Biological Survey had
nearly completed a recounoissance of Washinotou and Oregon, and in
previous years had carried its operations over extensive tracts in south-
ern, middle, and northeastern California, so that v\'ith the exception of
a rather large area in northern California fully two-thirds of the Pacific
States had been covered. In 1898, therefore, the nnworked part of
northern California, reaching from the Madeline Plains on the east to
the Pacific Ocean on the west, and from the Oregon boundary on the
north to Lassen Butte and adjacent parts of the Sierra on the south,
came to be the principal field of onr investigations. In this area Mount
Shasta occujjies a nearly central position.

All high mountains, particularly those that stand alone, are likely to
throw light on the problems of geographic distribution and are worthy
of careful study. Shasta, not only because of its great altitude, but
even more because of its intermediate position between the Sierrai and
the Cascades, promised an instructive lesson, and was therefore chosen
as a base station for part of the field work of 1898,

From work previously done in the Sierra Nevada of California and
the Cascade Kange of Oregon it was known that many species of ani-
mals and plants are common to both ranges, and many restricted lo
one or the other. Shasta, lying between tbe two, was expected to share
the common features of both, and in addition alford the northernmost
limit of Sierra species, the southernmost limit of Cascade species, or
an overlapping of both, so that its fauna and fiora, other things being
equal, should be richer than either. l>ut Shasta proved very much
drier than either the Sierra or the Cascades, and consequently many
species common to the two ranges were absent, and the total number
was less than was exi)ected. Nevertheless, the mountain shares a large
21753— No. 16 2 9

10

NORTH AMERICAN FAUNA.

[NO. 16.

percentage of the common species and is, as expected, a stepping stone
on wbich restricted Sierra and Cascade species overlap. But the rep-
resentatives of the two ranges are not equally apportioned. The most
evident gap is on the north, Shasta sharing many more species in com-
mon with the Sierra than with the Cascades. Indeed, the resemblance
to the northern Sierra is so exceedingly close, particularly in the mam-
mal fauna, that from the standpoint of geographic distribution Shasta
conld without violence be classed as part of the Sierra. This is the
more surprising in view of the fact that the geographical gaj) between'
Shasta and the Cascades is only half as broad as that between Shasta
and the Sierra. This subject is discussed in detail in the chapters
entitled 'The Boreal fauna and flora of Shasta contrasted with corres-
ponding faunas and floras of the Sierra and the Cascades;' 'Efhciency
of Klamath Gap as a barrier to Boreal species compared with that of
Pitt River and Feather River gaps collectively,' and 'Sources of the
Boreal faunas of Shasta and of the Sierra and the Cascades.'

ITINERARY.

Leaving the railroad at Sissou, at the west base of Shasta, we estab-
lished the first camp July 15, 1898, at a point known as Wagon Camp,

riG. 1. — Wagon Camp.

on the south slope of the mountain, about a quarter of a mile west o
Panther Creek, at an altitude of 5,700 feet (tig. 1). Wagon Camp issitu

OCT., 1899.]

ITINERARY.

11

ated in a descending tongue of Shasta rtrs between ascending tongues
of manzanita cliaparral, just above the uppermost grove of ponderosa
pines, on the boundary between the (Canadian and Transition zones.
It is abundantly supplied with water from several small springs, from
which tiny streamlets run short distances before disappearing in the
thirsty soil. Some of these springs unite to form a snmll marsh, in
which flourish a number of plants not found elsewhere on the mountain.
It is naturally a favorite spot for birds, and more sj^ecies were seen
here than elsewhere. Wagon Camp was occupied continuously by one
or more members of the party from July 15 to August 1, and at brief
intervals thereafter until October 3.

A few days after reaching the mountain I set out on a trip around
the peak in order to become familiar with the general features of the
region and lay plans for the season's work. On this trip I was accom-
panied by Vernon Bailey, my most experienced field assistant, and by a

Fig. 2. — Shasta from east brink of Mud Creek Canyou.

voluntary assistant, Lyman L. Merriam. We took saddle horses and
a pack animal, which were of material aid, although we had much dif-
ficulty in getting them across some of the deep canyons and over the
indescribably rough lava on the west side of the mountain.

Leaving Wagon Oamp on the morning of July 22, we ascended Pan-
ther Creek to its source, turned easterly through 'The [South] Gate,'
north of Gray and Red buttes, crossed Squaw Creek near its head, and
kept on among the timberline white-bark pines to the rim of Mud
Creek Canyon (pi. iii), which we followed down into the Shasta firs.
The first night was spent in the bottom of this canyon at an altitude
of 5,600 feet — some distance below the lower fall. The second day we
climbed the steep east bank of the canyon, here 1,000 feet deep (fig. 2),
crossed Cold Creek and Ash Creek Canyon below timberline, and reached

12

NORTH AMERICAN FAUNA.

[NO. 16.

Brewer Creek Canyon in tlie upper part of the wbite-bark pines. Find-
ing absolutely no grass or other feed for the animals here, we crossed
the canyon lower down (a little below the forks) and continued on
over rough lava ridges in the upper edge of the forest until dark,
when we camped on Inconstance Creek (fig. 3). The third day we pushed

'r^:<

I'lO. 3. — Shasta from Inconstance Creek, near tiiuberline on north side.

on around the north end of the mountain, keeping a little below the
great glaciers, and in the main near timberline. We climbed over a
number of lava ridges, availed ourselves of a natural passageway
('North Gate') at the upper end of a pair of conspicuous lava buttes,
traversed a curious pumice plain covered with timberline mats of
prostrate white-bark pines (fig. 22), crossed the fearful canyons of Whit-
ney and Bolam creeks, aud finally reached Shastina, where, after a very
severe day, we camped on some small streams of snow water on the
north side (fig. 4). The fourth morning we climbed the rough slide rock
of Shastina to an altitude of 10,000 feet, in order to get around a high
impassable lava ridge, and then, after encircling a great amphitheater '
of rough slide rock, descended by some immense masses of perpetual
snow to the white-bark pines, in which we continued to the great can-
yon on the west side of Shastina (pi. ii), which I named Diller Canyon,
in honor of J. S. Diller of the U. S. Geological Survey, in recognition of
his admirable researches on the geology of Shasta. After crossing
Diller Canyon we kept in the upper part of the Shasta fir forest all the
■way to Panther Creek, which we followed down to Wagon Camp. This

on-., 1899.

ITINERARY.

13

was the most trying day of all — sixteen miles of continual climbing,
removing blocks of lava, and building trail. Our animals suffered

Fig. 4.— Sliastiiia

'I til, showing great bank of snfiw which feeds Shastiiia streams.

severely, and one of them gave out entirely. However, the mountain
was completely encircled after four long days, and the desired informa-

FlG. 5. — Alpine hemlocks, Squaw Creek Camp.

tion "was obtained. In the main we kept near tiiuberline, climbing over
the bare rock slopes above, or descending into the dark forest below,

14

NORTH AMERICAN FAUNA.

[no. 16.

as occasion required. And since all the canj^ons of Shasta radiate from
the summit, all were crossed on this trip.

A base camp was next established in a grove of black alpine hem-
locks near the head of the west branch of Squaw Creek, close to and
just east of the upper end of lied Butte. Here one or more of tlie party
remained continuously from August 1 till September 24. All things
considered, this is probably the best camping ground on Shasta, though
I am not aware that it had been used before our visit. It is close by
the three upper 'meadows' on Squaw Creek and within reach of the
best feed for horses found on the mountain, with the possible exception

Fig. 6. — Heather meadow ou Squaw Creek.

of a small area near Cold Creek, and it is by far the most conven-
ient base from which to work the timberline region of tire southern
slopes.

Temporary camps were established at the head of Panther Creek, in
Mud Creek Canyon at the mouth of Clear Creek, in Ash Creek Can-
yon a little below timberline, and high up between Mud Creek Canyon
and the head of Clear Creek, from which point the main peak was twice
ascended. At the base of the mountain, work was done at Sisson on
the west side, in Squaw Creek and McCloud valleys on the south, and
in Shasta and Little Shasta valleys on the north, and finally a trip was
made completely around the mountain, mainly within the belt of yel-
low pines Avhich clotlies its lower slopes.

OCT., 1899.1 PREVIOUS PUBLICATIONS. 15

PERSONNEL.

Ill the iield work on wliicli the present report is based, I was aided
by Vernon Bailey, chief held naturalist of the Biological Survey, and
my assistants, Wilfred II. Osgood, Walter K. Fisher, and Eichard T,
Fisher. Vernon Bailey had charge of the work at the Shasta base camps
and on a trip around the base of the mountain; Walter K.Fisher
had charge of the work in Mud Creek and Ash Creek canyons and
near timberline east of Mud Creek, and afterwards took a party to Fall
River Lake and Lassen Butte; W. H. Osgood had charge of the work
on Lassen after Walter Fisher's departure, and also visited Squaw
Creek and Shasta and Little Shasta valleys; E. T. Fisher spent the
season working from the various camps on Shasta and at Sisson, and
accomi^anied Osgood on the trip to Little Shasta Valley.'

Our camps on Shasta were visited by Henry Gannett, chief geo-
grapher of the V. S. Geological Survey; John H. Sage, of Connecticut,
secretary of the American Ornithologists' Union ; and two or three
others, all of whom rendered important assistance.

In addition to the work on and near Shasta covered by the present
report, field work was done in various directions. Three cross sec-
tions of the Sierra Nevada, north of latitude 39"^, were made by Bailey,
Osgood, and myself; Bailey and Walter Fisher ran a line from Black
Eock Desert, Nevada, to Shasta, by way of Madeline Plains; Bailey
and I, accompanied by Henry Gannett, carried the work across the
wild and little known mountains from Shasta to the ocean, which
we reached at Humboldt Bay; and later in the season much work was
done farther south, chiefly in the inner and outer Coast Eanges.

PREA'IOrS PUBLICATIONS.

Only two publications have been found relating to the zoology and
botany of the Shasta region. The first is a report by Charles H. Town-
send, of the U. S. Fish Commission, who, fifteen years before our visit,
was stationed at Baird, a fish hatchery on McCloud Eiver. While
thereMr.Townsend visited Berryvale (now Sisson Tavern) and accom-
panied Major Gilbert Thompson, who was in charge of a triangulation
party of the U. S. Geological Survey, in his field work on Shasta. The
results of Mr. Townsend's work are contained in an important report
entitled 'Field Xotes on the Mammals, Birds, and Eeptiles of Northern
• California,' published in the fall of 1887.^ In addition to the records in
this report, Mr. Townsend has kindly placed his manuscript catalogue
at my disposal, and has in several instances given me important sup-

' While this report Ava.« passing through the press (July, 1899), I sent "Walter K.
Fisher to Mount Shasta and Shasta A'alley to obtain supplemental inforniatiou, some
of which is incorporated in the mammal, l)ird, and plant rejiorts at the end. — C. H. M.

-Proc. U. S. National Museum, X, pp. 1.59-241, Nov., 1887.

16

NORTH AMERICAN FAUNA.

[no. 16.

plementaiy iiiforniatiou respectiug the exact localities at wliieli speci-
mens were collected, all of which is duly credited in the body of the
present report.

The second publication referred to is a brief i)aper by Miss Alice
Eastwood on 'The Alpine Flora of Mount Shasta," containing the
results of a hasty trip to the summit made in August, 1S93.

So far as I have been able to ascertain, this is the first and only pub-
lication relating directly to Shasta plants, although a number of species
collected there during a brief visit by Prof. Wm. H. Brewer in the
early sixties are mentioned in the Botany of California (by Brewer and
Watson, 1876-1880).

NEW SPECIES.

In working up the collections it was found that several of the plants
and mammals belonged to undescribed species. Some of the new
plants have been described by Prof. E. L. Greene;^ others remain
unnamed. The new mammals are here described. Tlie new species
are:

Plants. Mammals.

Agoseris monticola.
Aruica merriami.
Campanula wilkiusiaua.
Pbacelia frigida.
Pyrola pallida.

Lepns klamathensis.
Lyux fasciatus pallesceus.
Neurotricbiis gibbsi major.
Procyon psora pacitica.
Reitliiodontomys klamathensis.
Sorex shastensis.
Thomoiiiys mouticola piuetorum.
Urocyon californicua townsendi.

> Erythea, IV, No. 9, pp. 136-142, Sept., 1896.
-Pittonia, IV, pp. 36-10, March 17, 1899.

GENERAL FEATURES OF SHASTA.

The snowy peak of Shasta, the pride of California, is one of the
highest and most accessible of the snow-clad glacier-bearing moun-
tains of the United States. It is an old volcano, 14,450 feet in alti-
tude, and is completely cut otf from neighboring mountains — from
Lassen Peak, at the north end of the Sierra proper, by the valleys of
the McCloud and Pitt riv^ers; from the south end of the Cascade Eange
in Oregon by a broad lava i)]ateau and the valley of Klamath River.
The breadth of the gap on the north is diminished by a cluster of low
volcanic mountains known as the Goose Nest Group.

Shasta is the best-known landmark in California. Seen from the
north, south, and east it appears as a single cone pushing its lofty
crown upward six or seven thousand feet above apparent timberline.
Seen from the west and southwest its summit is elongated and looks more
like the crest of a ridge (frontispiece). This appearance is due in part
to a large secondary volcano, Shastina, which rises from the northwest
shoulder of the mountain, and in part to a long ridge which pushes out to
the south. This west side, the one seen by tourists in traveling over the
Shasta route from San Francisco to Portland, is in many respects the
least interesting. From its exiiosure to the direct rays of the afternoon
sun it is the hottest slope, and consequently the one on which timber
reaches highest and on which the ice and snow are most reduced.

Like most isolated mountains, Shasta is seen to best advantage from
a distance. The most imposing view to my mind is from the northeast,
the region of the Modoc lava beds, from which the peak looms up in all
its icy grandeur — a single massive cone buried from toj) almost to bot-
tom in continuous glaciers, below which it is encircled by a dark belt
of coniferous forest. It is also very imposing as seen from the distant
Trinity Mountains.

The north and east sides of the peak are completely, and the south side
partly covered by glaciers, but not a glacier is to be found on the west,
where the large masses of white seen froni Sisson are banks of snow,
more or less permanent. The only glacier visible from the railroad is
Whitney Glacier, which occupies the notch between Shasta and Shas-
tina, and may be seen from points north of Edgewood. The higher
slopes, between the lower edge of the ice and snow and the upper edge
of the forest, are steep and rocky. In the main they consist of radi-
ating ridges alternating with glacial basins and precipitous canyons.
As a rule the surface is light pumice and pumice sand thickly strewn
with fragments of gray volcanic rock, interrupted here and there by
masses and cliffs of darker lava, often reddish brown in color.

2175.3— No. 10 3 17

II

NORTH AMERICAN FAUNA.

[NO. 16.

The south and east sides, except the deep canyons of Mud, Ash, and
Brewer creeks, are fair traveling- for mountain horses. The north side,
below the great glaciers, is interrupted by exceedingly rough lava
ridges and the terrible canyons of Bolam and Whitney creeks. The
west side, though scored by only a single notable canyon — Diller Can-
yon (pi. II) — is by far the most difticult. After crossing the tremendous
sloi^es of steej) and sharj) slide rock, very dangerous for horses, on the
northwest side of Shastina, and surmounting the two principal lava
ridges west of Shastina Creek, the way to Diller Canyon is comparatively
easy. But between Diller Canyon and Casca<le Gulch, a mile or so north
of Horse Camp, and extending from timberline downward several thou-
sand feet, is a chaos of lava the like of which I have never seen. It
suggests the worst parts of the Snake Eiver and Modoc lava beds
turned up on end — basins, ridges, and tumultuous piles without order
or direction, without beginning or ending — dry basins that empty
nowhere, drier ridges that lead nowhere, until one is worn out with
thirst and efforts to escape. The whole is hidden in a dark forest of
Shasta firs whose hardy trunks force themselves out between the lava
blocks in ways that almost surpass belief. Finally all this stops as
suddenly as it began, and one emerges from the dark inferno to slake
his thirst in the refreshing pools of Cascade Gulch — known only to the
deer — and, with a sense of infinite relief, reenters the area of pumice

sand and gray shale
whicli stretches away
to the southeast and
thence onward around
three-quarters of the
mountain.

The timbered valley
at the west base of
Shasta falls away both
to the south and to the
north. On the south
it drains immediately
into the Sacramento
Kiver; on the north
into the Shasta Eiver,
which traverses Shasta
Valley and empties
into Klamath River.

' /fl^^ ^^^^^^^^^hP

r.'&?»»''?.*-~."fc } '^

Fig. 7.- rmiiice sand sticwji witli gray volcauic shale. Yonug gli-ista VallCV IS ail
In-nilijcks in foregroimd: white-bark pines in distance. < ^ i ., x

open plain northwest
of the mountain; it is lowest at the north, and its northwestern cor-
ner ends in a pocket or basin containing the mining town of Yreka,
which is doubtless the hottest part of northern California west of the
axis of the Sierra-Cascade system.

North American Fauna, No. 16.

Plate II.

OCT., 1899.]

EFFECTS OF SCANTY MOISTURE.

19

EFFECTS OF SCANTY MOISTURE.

The flora of Shasta, contrasted with that of moister mountains imme-
diately north and immediately south, is poor in species and individuals;
and the same is true in less degree of the fauna. At least nineteen
characteristic genera and numerous additional species of jilants com-
mon to the Sierra and the Cascades, are unknown (p. 80) ; and to these
must be added the distinctive species of each range which fsiil to reach
Shasta. The luxuriant mountain meadows and flower beds that form
such conspicuous features of the timberline region in the Cascades, the
Olympics, the High Sierra, and the Eocky Mountains are wholly
absent, and the only areas that in any way resemble them are the

Fio. 8.— Heather meadow bordering Squaw Creek. Shasta peak in distance covered with fresli snow,

September 22, 1898.

insignificant patches of mountain heather and accompanying plants
that carpet the moist bottoms of the glacier basins and form narrow
beds along the tiny streams, where they are concentrated by the local
distribution of soil moisture. The only real soil above timberline is
restricted to the borders of the streamlets, where the decomposing
heather has left a shallow covering. Everywhere else are pumice,
broken lava, and barren clifts.

The summer rainfall amounts to little or nothing, and when rains
occur they sink and vanish in the thirsty pumice sand. The streams
from melting snows are exceedingly small, averaging hardly more than
a foot or two in width, and most of them disappear before reaching the
base of the unmntain. The turbid streams from the glaciers are larger,

20 NORTH AMERICAN FAUNA. [no. 16.

but tliey have cut for tliemselves deep gorges where they run their
rapiil courses ],()00 feet below the surface, aud cousequently are use-
less for i)urposes of geueral irrigation. They exert a local influence, it
is true, since far down in the damp bottoms of the canyons and along
their cool eastei^ly lower slopes a number of moisture-loving plants occur
that are not found elsewhere except about the few and widely scattered
springs in the forest — serving by contrast to accentuate the general
aridity. Even the black alpine hemlock, which in the Cascades forms
so attractive a feature of the upper slopes, is of local occurrence on
Shasta, where its distribution is interesting as furnishing an index to
soil moisture. It is associated with the white-bark pine {rinm albi-
mulis), which requires less moisture and is the dominant timberliue
tree. In our circuit of the peak we found the range of the white-bark
pine practically continuous; that of the alpine hemlock discontinuous
and greatly restricted. As a rule the hemlock is confined to narrow
strips along the streams and gulches, or to tongues along the cool east
sides of buttes and ridges, where the soil, sheltered from the hot after-
noon sun, is able to retain more moisture than elsewhere. Below the
alpine hemlocks and occupying the middle belt of the mountain is a
magnilicent forest of Shasta firs; but the humbler vegetation of this
belt is scanty and irregular.

From what has been said it is obvious that excessive dryness pre-
vents nuiny of the characteristic zone species from filling their appro-
priate belts, restricting them to scattered spots, where, as in the desert,
succulent vegetation is concentrated about springs and streams. Hence
Shasta is a poor place to study the broad general facts of zone distri-
bution, but, as shown later, an admirable place to study detailed effects
of slope exposure and humidity.

GLACIAL 35ASINS.

As in most parts of the Sierra and many parts of the Cascades, gla-
cial basins are conspicuous on the higher slopes of the mountain. They
occupy the deep depressions between the radiating ridges, and their
terminal moraines are usually clearly defined. In some of the valleys,
as along the upper part of Squaw Creek, two or three such moraines
may be found at intervals, marking successive stages in the retreat of
the glacier. The glacial basins usually contain small streams, at least
during spring and early summer, and they receive additional moisture
from the melting snows, which linger long in the shadows of the ridges.
This moisture permits the growth of a more abundant vegetation than
occurs elsewhere on Shasta, save only along the streams. The bottoms
of the basins therefore are usually carpeted with red heather {Bryanthus
or Phyllodoce cmpetriformis) and a variety of small plants, the majority
of which are inconspicuous except when in flower. Among the most
noticeable of these, each contributing its mite to the general verdure
of the heather beds, are the dwarf huckleberries, white alpine anteu-

OCT., 1899.

CANYONS.

21

narias, sileiies and ligusticuins, yellow monkey flowers, violets and
hieraciimis, blue veronicas and asters, cream-colored feathery lutkeas
and parnassias, pink epilobiums, red alpine laurels, and scarlet painted
cups.' True grasses are scarce, but grass-like carices abound.

The mammals inhabiting the heather meadows are the rare alpine
phenacomys ( Fhenacoinyn orophiius), the white-footed mouse {J'eromi/scxs
(jambeli)^ the long-tail mountain vole {Microtus mordcw), and the Sierra
pocket gopher {Tlioiuomi/s monticola). The gophers throw up their
characteristic mounds about the edges of the heather beds but are
commoner on the adjacent pumice slopes.

1"IQ 9. — Glacial meadow at head of Squaw Creek.

CANYONS.

All the canyons of Shasta radiate from the ice-covered summit and
take remarkably straight courses down the steep sides of the moun-
tain. Most of them are iirofound gorges cut by swift-flowing glacial

^ The plants of the glacial basins in the timberline region vary somewhat with the
raoi.stnre of the soil. The commonest species in moist spots and along the borders
of the streamlets are: Arnica merrkimi, CastiUeja miiiiata, EpUohinm claratum, Jliera-
ciinii (jracile, Hypericum anafjalloides, MimuJus implcrus (growing in the water), Mimu-
hm primnJoides, Mitella pentandra, Parnaasia califorynca, Veronica citsicki. The com-
monest species in the drier parts of the heather beds are : Antennaria meilin, CastiJleJa
tniniaia, Hicracium (jracile, luiJmia i/lanca microphyJlu, Lufkea pcclinata, Lit/iisiicnin
grnyx, PlnjUodoce empetriformis, Sibhaldia procumhens, Silenc f/rayi, Vaccinium casjii-
tosum, J'iola jnirpurea.

22 NORTH AMERICAN FAUNA. [no.16.

streams. They average about a tbousaiid feet in depth, aud tlieir
slopes are as steep as permitted by the unstable material through
which they are cut — usually pumice, gravel, and fragments of lava.
Owing to the fact tliat all of the great glaciers are on the north, east,
or southeast sides of the mountain, the canyons made by their rivers
are necessarily on these sides also. The only one of any size which
does not come from a glacier is Diller Canyon, on the west side of
Shastina.

In most, if not all, cases the bottoms of the canyons in their upper'
courses are bridged for long distances by masses of ice and snow — the
dumps of avalanches. Below these snow bridges are vast accumula-
tions of loose stones, which in several instances, as in Brewer, Bolam,
and Whitney canyons, are piled up in a curious manner. During
periods of high water the rocks that fall in are carried down by the
torrent and deposited on each side in banks several feet high, so that
the traveler on reaching the bottom has to climb up over a ridge of
loose stones and down again before coming to the stream. These lat-
eral ridges form miuiature canyons in the bottoms of the big ones.
Most of the canyons have falls several hundred feet high in their upper
courses, and some have other falls farther down. Rotable falls are
found high up in the canyons of jMud Creek, Ash Creek, Bolam Creek,
and Whitney Creek. While difilicult of access, they are well worth the
efibrt of a visit.

Mud Creek Canyon (pi. in), the only one likely to be seen by the
ordinary visitor to Shasta, is not easy to cross except near the mouth
of Clear Creek, which comes into it from the east. Its east bank is a
precipitous single slope about 1,000 feet in height. Its west bank,
except above timberline, is broken by a forest- covered terrace or bench,
and both descents are likewise steep, though less difficult than the oppo-
site side. The canyon of Ash Creek is better timbered and a little less
precipitous than that of Mud Creek. The canyons of Bolam and Whit-
ney creeks, like that of the upper part of Mud Creek, are terrific naked
chasms, very deep and so steep that in most places the loose material
of their sides will not sustain the weight of a man — much less that of a
horse — and when disturbed dashes in avalanches to the bottom.

Diller Canyon is peculiar (pi. ii). It is a tremendous gash on the west
side of the otherwise symmetrical cone of Shastina, which it cleaves
from top to bottom before taking its practically straight westerly
course down the rest of the mountain. It is the only canyon on Shas-
tina, the only notable one on the west side of Shasta, and the only oue
anywhere on the mountain that does not emanate from a glacier. Its
stream comes from enormous banks of perpetual snow.

While the upper parts of the canyons are exceedingly steep and
barren, and practically devoid of vegetation, the middle and lower
parts are invaded by the trees of the adjacent sloi)es, and in marshy
and springy spots contain patches of willows, alders, and a multitude

North American Fauna, No. 16.

Plate III.

W
o

c

H

w
>

(/)

b
tn

o

- >

5 o

- o

OCT . 1899.]

STREAMS.

23

of smaller plants. These places, in Mud Creek and Ash Creek canyons,
are the homes of the mountain showt'l or sewellel {Aplodontia niajor)^ a
curious bob-tailed rodent resembling- a large muskrat, which lives in a
labyrinth of subterranean passages in wet ground, and cuts and drags
to its burrows bundles of coarse plants on which it feeds. Weasels
[Putorius arizonenshs) are usually found in the aplodontia colonies
and it is safe to assume that their presence there is the most serious
factor in the life of the rightful owners of the land.

STREAMS.

The streams that come from glaciers are rapid, turbid, and muddy,
and have cut deep V-shaped canyons down the steep slopes of the
mountain. Those that come from melting snow are clear as crystal

Flo. 10.— Heatlier meadow on unper Squaw Creek, sbowini; lomentration of vej:etation near stream.

and usually flow on the surface or in shallow channels hardly more
than a foot or two in depth. They are smaller and less constant than
those from the glaciers, and in times of high water carry so much
gravel and pumice that they often block their own shallow channels
and overflow, cutting new courses near the old ones. During the fluc-
tuations incident to the irregular melting of snow they often reopen the
older channels and at the same time retain the new, so that on the
higher slopes it is not unusual for a mountain rivulet to occupy several
beds at the same time. These are commonly separated by intervals of
a few feet or a few rods, and the spaces between are often covered with
patches of red heather, dotted with flowering plants of many kinds.

24

NORTH AMERICAN FAUNA.

[NO. 16.

Tlie bauks of the more permanent streamlets are so well supplied
with moisture that the heather and other plants, often mixed with
alpine mosses,^ form a sod which, growing thicker each year, gradually
comes to overhang and finally bridge the swiftly running water. Even
away from these sod bridges, which for long distances completely hide
the water, the space between the constantly protruding banks is so
narrow that only the middle part of the stream can be seen.

The icy rivulets abound in cascades, miniature waterfalls, and crys-
tal pools, bordered by overhanging banks of moss, heather, and dwarf
alpine laurel, and adorned by the waving heads of the scarlet painted

•^TV,

r'^;

rf^<r

Fig. 11.— Miuiature caaciule on upper Sciuaw Creek, trequented by w.iter shrews and ouzels.

cups and cream-colored parnassias. They are ideal homes for the water
ouzel {Cinclus mexicanus), the large water shrew {Xeosorex navk/ator),
and the mountain vole {Microtus mordax). The latter animal is par-
ticularly abundant, and its dark burrows, almost hidden among the
plants, may be found opening out on the vertical overhanging banks
only a few inches above the water, so that whenever the occupants
come out they may ])luuge in the icy stream before ])roceeding on their
journey. Minks occur lower down along the streams, and still lower,
otters.

iThe prevailing moss on the banks of the alpine streams is Anhicomninm androgy-
num.

OCT.. 1899.]

ROCK SLOPES,

25

EOCK SLOPES.

The whole upper part of the mountain between the glaciers and snow
banks above and the forest belt below consists of bare rocky sloi^es,
broken at intervals by precipitous cliffs and small heather meadows.
The slopes are largely pumice sand, strewn and mixed witli fragments of
gray volcanic rock, among which the individual i^lants are so scattered
as to disappear in the general view.^ White-footed mice {Pcromyscus
(jamheli) are common on these slopes, feeding on seeds of Polygonum
neicherryi and other timberline i)lants. Pocket gophers [Thomomys
monticoJa) occur here and there and throw up their characteristic mounds
in the pumice sand between the rocks. They subsist on the tough roots

Fig. 12. — C'liaiacteristii' roi-k slope on north siilc of Sba.-^tiiiii.

of alpine plants, and were observed at intervals up to an altitude
of 9,000 feet. The pika or rock cony (Ochotona scMsUceps) inhabits
steep rock slides at distant points around the j^eak, and lays up stores
of plants for winter use. Another mammal inhabiting the rock slides

'The commonest plants of the bare stony pnmice slopes are: Afjoaeria monticola,
Antennaria media, Jrabis platijsperma, Cha'nactis )ierad()isis, Chrysothamnus hloomeri,
Cymopterus tcrehinthhuis, Eriogonum pohjpodum, E. pyroht folium, Erigeron compositus
irifidus, Hulsea larseui, If. nana, Luikea pectinata, Lupinua ' ornatus,' L. lyalli, I'ent-
siemon menziesi, J'hlox douylasi diffusa, Polygonum newherryi, P. shaslense, Saxifraga
tolmiei, Senecio canus, Silene grayi, S. suksdorfi, Spraguea umhellata, Strepianthua
orhieulatus, and Viola purpurea. Besides these, several ferns occur very sparingly
on the rock slojies. These are Dryopterls aculcata scopulina, Cy>itopteris fragilia,
Clieilanthes gracillima, and Phegoptcris alpentvis.
21753— No. 16 4

20 NORTH AMERICAN FAINA. fmie.

and cliffs, but for some unaccountable reason even rarer than the cony,
is the bushy-tailed wood rat or pack rat (Net>t(>)iia dnerea), which, if
my memory serves me correctly, is less connaoii on Shasta than on any
other mountain I have visited in the West. The mountain chipmunk
(Eutamias scnex) and golden ground squirrel {GaUoKperinopMliis chryso-
deini.s) inhabit the tongues of ])iues on the ridges, and not infrequently
live in burrows among the bare rocks. Marmots, it is safe to assert, are
altogether absent. We completely encircled the peak in the neighbor-
hood of timberliue, and examined innumerable ledges and rock slides,
such as on other mountains are inhabited by marmots, but without tind-
ing so much as a track or sign or even a bleached bone to indicate that
any member of the genus ArciomyH had ever inhabited Shasta. In
former days the bighorn [Oris canadensis) was commo)i here, but now
the occasional fragment of a skull or the scattered parts of a skeleton
are all that remain. In fall the old bucks of the Columbia black-tail deer
wander up on the higher ridges. Here and there, particularly in the
shelter of the prostrate white-bark pines, tracks and dung of rabbits
were seen, but in spite of all our efforts no member of the party succeeded
in liuding a rabbit on the mountain. The species is probably the Sierra
rabbit [Lepus llam((t]iensis), though from the large size of some of the
dung pellets I was inclined to suspect the i)reseuce of Lepus campestris.

AVALANCHES.

During the loosening of the S7iow in spring, avalanches must be very
common on the higher vslopes, and it is probable that they exert a
controlling influence in determining the timber areas above the limit ol
continuous forest. iS^othing forces itself on the observation more firmly
than the peculiar way in which the white-bark pines are restricted to
the long radiating ridges where they form narrow tongues, sei^arated
by broad intervals of steep slopes and basin- shaped valleys. While it
might be hazardous to assume that the absence of trees from these
extensive slopes and basins is due mainly to avalanches, the fact
remains that the tracts they occupy along the tops and upper slopes
of the ridges are entirely out of reach of these resistless engines ol
destruction.

Now and then, however, an avalanche, taking an unusual course,
reaches the outskirts of one of these tongues of alpine pines and
snatches up and carries below all that lie within its path. This h
evident from the weathered trunks and roots often found at the bot
toms of slopes where trees have never grown.

The most conspicuous path of a recent avalanche observed is on CoU
Creek, between the deep canyons of Mud and Ash creeks (tig. 13). Hen
an avalanche of unusual size must have shot down the higher slope.-
until it reached the upper edge of the continuous forest of Shasta firs
where, instead of stopping, it cut a broad swath through the hug(
trees, tearing them up by the roots or snapping them off' and carrying

OCT., 1899.]

TIMBERLINE.

27

them oil over an almost level tract with such resistless force that the
i'ew now left standing are deeply scarred at a height of 10 to 15 feet
above the ground, showing where they were struck by other trees in
passing over the deep snow. Hundreds of huge trunks 75 to 100 feet
long and 3 or -4 feet in diameter arc strewn in desolate confusion over
the broad area that marks the jjlace where this terrific avalanche
slowed up and finally stopped. The accompanying illustration shows
a part of this area, and also the gate cut by the avalanche when it
struck the upper edge of the forest.

Fli.i. i:3.— Trai-k of avalauclie iiivadiu^, loie.-^l i>l' Slia^ta liis on Cold Creek, east side of Shasta.

TIMBERLINE.

Timberliiie is the upper or boreal limit of tree growth, as determined
by temperature. It varies somewhat according to the particular species
of tree, for even Hudsonian species differ in the degree of cold they
are able to endure. Thus in the northern Cascades where the alpine
hemlock and alpine fir are the dominant tiraberline trees, the tir pushes
up to higher altitudes than the hemlock. So on Shasta, where the
alpine tir is replaced by the white-bark pine, the latter is the true tim-
berliiie tree and always attains higher elevations than the hemlock.

Theoretically, nothing is easier than tracing timberline on a moun-
tain whose upi)er slopes are bare or dotted with alpine flowers and
whose middle slopes snpport a continuous forest. Yet on Shasta, and
on most high mountains, it is exceedingly difficult to tix the boundary
of timberline or indicate its exact position on a map. Of course, it is

28 NORTH AMERICAN FAUNA. [no. 16.

possible to follow tbe uppermost trees wherever tliey may lead, but a
map showin<^- such a route would resemble a saw from which alternate
teeth had been removed, the remaining teeth indicating the way the
dwarf trees push up on the summits of ridges, the broad spaces
between the teeth, the treeless gaps, usually the intervening valleys
or basins. Trees always occur at some point in the bottoms of these
valleys, and usually extend completely across them, but at an altitude
a thousand feet or more lower than that reached on the ridges, and
there is a material difference in the trees themselves. If of the same
species, those in the valleys are much larger and taller; if of other
species, as is frequently the case, they belong to the upper part of the
belt below — the middle forest belt. On Shasta, the trees that push up
highest on the ridges are always the dwarf white-bark pines, while
as a rule those that bridge the intervening valleys below are full-
grown alpine hemlocks or Shasta firs, the upper limit of which must
not be mistaken for timberline. The difficulty lies in determining
what ought to be considered true timberline, and the reason why
in the absence of obvious barriers the white-bark pines do not fill more
than a third or a fourth of the belt to which they properly belong.
If a mountain could be found whose upper slopes form a true cone
instead of a series of alternating ridges and valleys, so that suc-
cessive transverse sections would be circular in outline, instead of
irregularly scalloped, it is probable that timberline would form almost
a true circle around the peak, rising a little on the southwest and
dipping down a little on the northeast. But in the absence of such
ideal conditions, actual visible timberline is usually confined to the bor-
ders of the tongues of dwarf trees that occupy the summits of the radi-
ating ridges (pi. iv). The explanation of the absence of trees from the
intervening valleys is not always easily found; still, if the valleys are
studied with reference to the details of their several slope exi^osures
and other local conditions, the position of the hypothetical timberline,
in most cases, will be obvious. Let us take, for instance, one of the
numerous glacial basins on the south side of Shasta, bordered on each
side by lofty ridges which are capped by tongues of white-bark pines.
The bottom of the valley, whenever its axial slope is steep enough to
be regularly swept by avalanches, can not, of course, contain trees. The
broad basin slope of the ridge on the west faces east and is in its own
shadow in the afternoon; as a consequence it is too cold for trees, but
is well sprinkled with alpine plants. Its summit is covered with dwarf
white-bark i^ines, which come up from the other side and end abruptly
along its eastern crest. The cold eastern slope is, in its zone position,
actually above timberline, although the tongue of dwarf trees along
its crest may stretch up a thousand feet above the lowest alpine
plants.

On the opposite or eastern side of the basin the slope faces west or
southwest, and receives the warm rays of the afternoon sun. The

North American Fauna No. 16.

Plate IV.

OCT., 1899.1

TIMBERLINE.

29

result is tliat this slope, unless too steep or otherwise uusuited to tree
growth, or within the track of avalanches, is commonly covered with
white-bark pines. As a rule the timbered area on these westerly slopes
takes the form of a broadening tongue, beginning- at the highest alti-
tude attained by trees on the crest of the ridge and increasing in width
at lower altitudes until the l)ottom of the valley is reached and skirted,
and the limit of avalanche movement jjassed, when the trees again
strike out boldly. The pines in the basins are much larger and more
erect than those on the summits of the ridges; they decrease in size
with increase in elevation. The long oblique line which on the west
side of each ridge marks the lower limit of tree growth commonly

'^i4%

'0ff\

V

.Ji

Fig. 14.— lli-h tiiuljL'rluiu iiil,i;e, .-^liuwiii.u t-lRcts of slope exi)osure. The (lurk [lati'lics on the left
(west) siJe of the riilge are dwarf w)iite-hark piues. (Photographed by John H. Sage.)

marks also the upper limit of the area in shadow during the late
afternoon. In local spots other factors may account for the absence of
trees. Thus, they are always absent from the avalanche-swept bot-
toms of the valleys, and from ground kept wet by springs or melting
snow. Studied with these facts in view, comparatively few treeless
areas will be found which can not be explained, and the position of
true timberline may be fixed with some degree of certainty. This is
really a very important matter inasmuch as it affects the zone position
of a great many species.

It is necessary to remember that the reason trees are absent from
the cold east and northeast slopes of the ridges whose summits are

30 NORTH AMERICAN FAUNA. [N...16.

covered with dwarf trees is that these slopes are iu their zone position
truly alpine and above timberline, as alrea<ly ex])lained.

Nothing is easier than to refer to the wrong zone species found in the
treeless basins between the pine-covered ridges. But when it is under-
stood that parts of each basin, regardless of tlie distance below the
highest tongue of timber, are unquestionably above timberline (and con-
sequently Alpine) and that other parts, regardless of the distance above
the nearest trees in the basin, are unquestionably well beloir timberline
(and consequently ffiKlsonian), mistakes of this kind will be less
frequent.

THE FORESTS OF SHASTA.

Shasta rises from a forested region (pi. v), and the mountain itself is
continuously forest-covered up to an altitude of 7,500 or 8,000 feet. The
trees of the lower slopes are those of the surrounding region, but those
of the middle and upper slopes belong to such widely different species
that it is necessary to divide the mountain forest into three belts, which,
from their most distinctive trees, may be designated (1) the lower or
yellow-pine belt; (2) the middle or Shasta tir belt, and (3) the upper or
white-bark i)ine belt. It is interesting to observe that these forest
divisions, as shown later, coincide with the tliree Life zones — the
Transition, Canadian, and Hudsonian.

(1) The Low^er Belt or Belt of Yellow or Ponderosa Pines (T'dihs ponderoxa)

The most abundant and characteristic tree of the lower slopes and
surrounding region is the yellow or ponderosa pine, which forms a con-
tinuous open forest up to an altitude, on the south and west sides, of
about 5,500 feet. The only material gap in the i)ine belt of the moun-
tain i^roper is a strip about 8 miles iu length on the cold northeast
quadrant, which is occupied by lodge-pole pines belonging to the zone
above (Canadian zone).

On the south and west the open pine forest of the basal slopes is
interrujited by extensive parks, which from a distance appear to be
meadows of waving grass. A nearer view shows this to be an illusion,
the broad fields of green being in reality impenetrable thickets of
chaparral — a chaparral of unyielding manzanita and buck brush
(.Aretostaphijlos paiula and Ce<inothns velKtuiiis, see tig. 15).

Northwest of Shasta the yellow pine forest is interrupted by the open
plain of Shasta Valley, which on the southwest ends abruptly at the
town of Edgewood. North, northeast, and east of Shasta the ponder-
osa pine forest continues with unimportant interruptions to Devils
Garden, (loose Lake, and the Madeline Plains; on the south it is prac-
tically continuous to the base of Lassen Butte, and thence along the
flanks of the Sierra for 350 miles ; on the southwest it follows the canj^on
of the Sacramento River to a little below Delta, where, in the bottom
of the canyon and on its warmer slopes, the curious digger i)ines ot
the Upper Sonorau zone mix with and soon replace the ponderosa pines

Motth American Fauna No. 16.

Plate V.

Pine Forest at west base of Shasta
Showing yellow and sugar pines.

OCT., 1899.

YELLOW PINE BELT,

31

of the Transition zone. On the cooler and liigher canyon slopes and
adjacent foothills the ponderosa pines continue to tlie border of the
Sacramento Valley. West of Shasta they cover all but the highest
elevations of the Scott IMountaius, completely surround Scott Valley,
and reach up a considerable distance over the east arm of the Salmon
Mountains, where, mixed as usual with Douglas firs, incense cedars,
and sugar pines, they fill the Transition zone. Still farther west they
occur in greater or less abundance in the valleys of Russian Creek,
North and South forks of Salmon River, Trinity River, and Klanuxth
Eiver, and at appropriate altitudes on the west arm of Salmon ]\ruun-
tains, Trinity Mountains, and the mountains between Iloopa Valley
and Redwood Creek. Hence the Shasta forest of ponderosa pines is

Flu. 15. — Mau/.anitii chapaiTal ou .south -slope of Slia.sia.

directly continuous — either broadly or by narrow and tortuous tongues — •
with corresponding' forests of southern Oregon, northeastern Califor-
nia, northwestern California, and the flanks of the Sierra.

The ponderosa pines of the Siiasta i)lateau and adjacent region are
peculiar — peculiar in the extent of their variability — and deserve care-
ful study. Not only do the cones of adjacent trees ])resent an unusual
degree of variation in size and com[)actness (particularly noticeable in
Scott Valley), but the cones of trees subjected to apparently slight
differences of temperature, moisture, and soil present certain average
differences that are quite surprising. Moreover, on higher parts of
the Scott Mountains, and also along their cool east base, fairly tyi)ical
Piniifi jejf'reiii grows within a short distance of pondero.sd. Whether or

32 NORTH AMERICAN FAUNA. [no. 16,

not tbe two actually intergrade, while an interesting question, is of
little consequence conq)ared with the fact that here, as in the Sierra,
the two trees occupy adjoining but distinct belts — pondcrosa the warmer
and normally the lower; Jeffrey! the colder and normally the higher. It
sometimes hai)pens, however, as in places along the cold east base of
the Scott Mountains, where local conditions produce abnormal tempera-
tures, that a strip of Jeffrey pine is sandwiched in between two areas
of ponderosa pine. In this instance the low temperature comes in part
from the cooling effects of cold streams, and in part from the afternoon i
shadows of the mountains.

The ponderosa forest is nowhere pure over any large area, but is
sprinkled in varying proportion with sugar pines, incense cedars,
Douglas tirs, and white firs, and at lower altitudes with black oaks.
The stately sugar pines are so valuable for lumber that the best have
been already cut, but enough remain to show that the sj^ecies Avas

formerly common in most
parts of the ponderosa for-
est. The incense cedars
also are scattered over the
whole region, but the Doug-
las and white firs require
more moisture and conse-
(juently are less evenly dis-
tributed. They are most
abun<lant on the borders
of streams, in cool canyons,
and along the well- watered
east base of Mount Eddy and the Scott Mountains, where they become
the dominant trees, the ponderosa j)ines being comparatively scarce.
On drier and warmer soil, away from the cooling inlluence of the Scott
Mountains, the ponderosa i>ines rapidly increase, and in the area
between Black Butte, Shasta Valley, and the mountain, although
sprinkled with incense cedars and black oaks, they form the purest
l^onderosa forest of the region.

Another conifer of the yellow-pine belt is the knobcone pine [Pinxs
attenuata), a narrow interrupted tongue of which i)ushes up Panther
Creek. The deciduous trees of this belt are the black oak {Quercus
calif 0 mica), Oregon maple [Acer macrophylluni), tree alder {Ahuis
temiifolia), and Oregon dogwood [Cornus nuttalli). The maple and
dogwood are restricted to the lowest levels and do not occur in very
dry places; the alders are confined to the neighborhood of water: the
oak ranges more widely over the lower half of the pine belt and thrives
on dry as well as on moderately moist soils.

The conifers will be considered as individual species.
SuciAR Pine {Finns lamhertiana). — The sugar pine is the largest,
handsomest, and noblest of our western pines, and its wood is so

Fid. 16. — Cones of poiulerosa and Jett'rey pines.

OCT., 1899.]

YELLOW PINE BELT.

33

valuable for lumber tliat, except iu iuaccessible places, the best trees
have been cut. The liuge trunks, often G or 7 feet in diameter, rise as
straight symmetrical pillars to a lieight of 150 or 200 feet, and are cov-
ered with fine beautiful bark. The long and graceful branches are
usually confined to the upper parts of the trees, and the cones they
carry are the longest known, frequently attaining a length of a foot and
a half and sometimes of 2 feet. They are very light, however, and
when falling are by no means
so dangerous to the passer
below as the shorter and
more massive cones of the
digger pines.

Around the base of Shasta
the sugar pines reach from a
point on the northwest slope
about 4i miles southeast of
Edgewood, near the south
end of Shasta Valley, south-
erly and westerly all the way
around to Ash Creek, where
they cease at an altitude of
about 5,000 feet. They are
fairly common in McCloud
Valley and at Sisson, whence
they extend south along the
Sacramento Canyon to 'The
Loop.' They are at present
more abundant iu the neigh-
borhood of Black Butte than
elsewhere about the moun-
tain. In the Shasta region
they are not so large as on
the west slope of the Sierra
iu central California; still
the stump of a sugar pine measured by me in McCloud Valley was 7
feet 7 inches in diameter G feet above the ground.

Knoboone Pine [Pinus attenuafa, fig. 17). — The knobcone pine is
a tree of erratic distribution. On Shasta it is confined to the lower
slopes on the south side, from Panther Creek easterly to a point
between the branches of Mud Creek, where it ranges irregularly from
an altitude of 3,800 up to 5,G00 feet. The latter limit is attained in a
gully a little east of Wagon Camp, in a continuation of the Panther
Creek strip. Lower down on Panther Creek, where the original forest
of ponderosaand sugar pines has been removed by the combined work
of lumbermen and forest fires, and the slopes are now covered by
an impenetrable jungle of manzanita, this singular pine remains, com-
21753— Xo. 16 5

Fig. 17. — Knobcone pine on Panther Creek.

34 NORTH AMERICAN FAUNA. [no. 16.

monly growing- in narrow lines. The trees are rarely more than 50
feet in height, and mo.st of them are mnch smaller. They bear a mar-
velous load of slender curved cones, which on the limbs grow close
together in whorls or rows, and on the trunks are scattered or grow in
circles. They remain on the trees for many years, as in the case of
few other species, and their large size, extraordinary numbers, and
peculiar arrangement give tlie tree a singular and unusual appearance.
Vernon Bailey has given me the following note on some knobcone
pines examined by him on Panther Creek, September 27, 1898 :

The trees were loaded with coues, in whorls of three to seven around the hranches,
and down the trnnks to 10 or 12 feet from the ground. Some of the cones must have
been 20 or 30 years ohl, and perhaps much older. I cut off a lot of the old lower
cones to see if the seeds were good, and put them on a bowlder and cracked them
with a few hard blows of the as. All of them were full of worm dust, with only
now and then an undiscovered seed or a fat white worm. Cones of medium age (5
or 6 years back from the end of the branch) were invariably occupied by worms and
worm dust, and usually contained few good seeds. Coues only 1 or 2 years old
were rarely wormy. A great many of the old cones had been dug into by wood-
peckers, either for seeds or, more likely, for the fat white grubs that live on the
seeds. The cones are too hard to be broken or split ajiart by the woodpeckers, and
are opened by a smooth hole drilled into the middle, or sometimes to the opposite
wall. Usually the opening is long and narrow. Sometimes the whole inside of the
cone has been drilled out, leaving only the shell ; sometimes a small round hole has
been drilled just through the outer shell.

White Fir {Abies concolor lon-iana.) — The white fir ranges from
Sisson, at the bottom of the west slope of Shasta, up to the lower edge of
the Shasta fir belt, which it slightly overlaps. At Wagon Camp (fig. 1,
alt. 5,700 feet) both species are common. The Avhite flr requires more
moisture than the other conifers of the lower timber belt, and con-
sequently its distribution is discontinuous. It is most abundant aloug
the well-watered eastern base of Mount Eddy, north of Sisson. The
highest elevation at which it was observed is a warm ridge on the
east side of Mud Creek Canyon, between the mouths of Mud and Clear
creeks, where, with a number of other Transition zone species, it occurs
at an altitude of 6,7U0 to 7,000 feet. This is 1,000 feet above its usual
limit, and its presence here is due to the angle and steepness of the
slope, as explained elsewhere (p. 49).

Ahies lou-iana is easily distinguished from A. shastensis and A. mag-
nijica by the bark, which is very thick and deeply furrowed, so that it
resembles that of Douglas spruce [Pseudotsnga mueronata) much more
closely than that of the other Ahies of the region. The cone scales are
broad and rather short, and the bract is short and tricuspidate (fig. 19).

Douglas Fir or Spruce {Pseudotsuga mueronata). — Douglas fir is
scattered irregularly through the ponderosa pine forest. Like the white
fir, it prefers a moister soil than suits the jjonderosa pines, and there-
fore thrives best in the gulches and near the streams. Thus along the
cool well-watered east base of Scott Mountains the forest consists maiuly
of Douglas and white firs, with scattered incense cedars and sugar

OCT., 1899.]

YELLOW PINE BELT„

35

and ponderosa pines, while on the drier ground a little farther east
the pines increase and the tirs decrease or disapj)ear. Most of the
larger trees on the south and west slojies have been cut for lumber. A
stump near McCloud Sawmill measures, at 6 feet above the ground, 8
feet in diameter.

Incense Cf.dar {Libocedrus decurrens)^ — The incense cedar occurs
in greater or less abundance in all parts of the ponderosa forest, on
both dry and wet ground, and from the bottom of the valley at Sisson

Fio. 18. — liiceuse cedar ou south slope below Wagou Camp.

up to the edge of the Shasta firs at Wagon Camp. But it is common-
est near the cool east base of the Scott Mountains. In moist places
the trees often grow in groups, but in the dry forest they are usually
scattered at intervals among the pines. Ou Shasta the bark of the
cedars is generally smooth and free from scales, except ou the very
youngest trees. In the more humid area between Scott Mountains and
the coast the scaly bark persists for many years, so that the trunks of
middle-aged trees look very different from those of corresponding size
m the dry interior.

36 NORTH AMERICAN FAUNA.

[NO. 16.

(2) The Middle Belt or Belt of Shasta Firs {.Ihirs shastensis).

Above tlie forest of poiiderosa pines, and therefore not connected
with similar forests elsewhere, a belt of Shasta firs averaging 2 or 3
miles in breadth and 2,000 feet in vertical range completely encircles
the mountain. It is the distinctive forest of Shasta — a forest of tall
stately trees, dark, somber, and free from underbrush, thragh here and
there beds of the low mountain manzanita {Arctostaphylos nevadensin)
aftbrd a pleasing relief from the uniform dark brown of the surface
carpet — usually a shallow layer of fir needles mixed with decayed cones
and wood. The massive trunks, which on the steep slopes are often
swollen jnst above the ground to give greater strength to resist the
pressure of snow, average from 4 to C feet in thickness and some attain
a diameter of 7 or 8 feet. Above the level of winter snow their northern
sides are usually covered with the handsome bushy yellow lichen,
Evernia vtilpina, which also clothes many of the branches; and in the
denser parts of the forest the trees are draped with pendant masses of
the long black-beard lichen, Alectorla fremonti. The forest almost
everywhere consists of large mature trees, and is free from evidences of
fire; but in one place, between Mud Creek Canyon and Cold Creek, a
considerable area, evidently an old burn, is covered with young trees
averaging perhaps 20 feet in height.

As a rule, the Shasta firs stop abruptly where the white-bark pines
begin, and trees at their upper normal limit are of full size. But now
and then on the steep and relatively warm southwesterly slopes of
the ridges, dwarf Shasta firs occur. The highest point at which such
were observed was at an altitude of 8,000 feet on the east rim of Mud
Creek Canyon, where a few stunted trees 3 or 4 feet in height were found
mixed with white-bark pines. On a similar warm slope west of Squaw
Creek a scattered line of these trees was noted at an altitude of 8,100
to 8,300 feet. Here the largest were 20 feet in height. Their bark dif
fered materially from that of trees lower down, being i)ale grayish
instead of dark brown, and thin, smooth, and full of blisters, instead of
thick and deeply furrowed. These fir trees were mistaken for another
species until 1 had the good fortune to find a few bearing cones, which,
on August 1, were two-thirds or three-fourths grown. To my surprise,
they had long exserted bracts which, as in the young cones, stood
straight out horizontally. Very young cones (2 to 3^ inches long) always
appear to consist of bracts alone, the scales being hidden inside. On
breaking open the cones the tiny scales are seen encircling the axis;
they are less than half the size of the bracts and occupy the inner half
of the diameter or section of the cone.

The normal bark of the Shasta fir resembles that of the alpine hem-
lock. It is rather regularly furrowed vertically, and the plates between
the furrows are cracked horizontally, so that it suggests that of the
ponderosa pine, though the plates are smaller and less red. Along the
lower edge of the Shasta fir belt the bark is darker and the cracks
and furrows are narrower.

OCT., 1899.]

SHASTA FIR BELT.

37

The upper edge of tlie Shasta fir belt meets the alpine hemlocks and
white bark pines of the belt above; the lower edge the pouderosa pines,
incense cedars, Douglas spruce, and white firs of the belt below.
The firs are easily distinguished by bark, branches, and cones. The
Shasta fir has very dark and relatively thin bark, regularly furrowed
so as to form 'plates' like those of the pouderosa pines, only smaller,
narrower, and transversely cracked. The branches are irregular, droop
at first (from the weight of Avinter snow), and then curve upward, and
the branchlets are small and terete, and stand out with mathematical
precision ; the cones are huge, and their green, tongue-like, single-xwinted
bracts protrude far beyond the scales, as in the noble fir of the north-
ern Cascades. In young cones the bracts
stand out straight; in old cones they are
strongly deflexed. The white fir [Abies
lou'iana) has much thicker and grayer
hark, deeply furrowed at base and not
forming regular scales or plates; the
branches are more regular and more nearly
horizontal, the branchlets flatter, more
spreading, and lacking the mathematical
lines of the Shasta fir ; the cones ar^} more
slender, and the tricuspidate bracts are
short, reaching less than half-way across
the scale. The cone-scale differences
are shown in the accompanying diagrams.
(See fig. 19. ) The year 1898 was an ' off year '
for cones, but plenty of old scales were
found on the ground, and broken cones
were discovered in holes in logs, where
they had been carried by pine squirrels.

The Shasta fir forest is mainly pure,
but in places, particularly on the east
and northeast sides of the mountain, silver pines are scattered through
it, and in one place along its lower border (between Ash and Incon-
stance creeks) the firs are replaced by lodge-pole pines, the only ones
on the mountain.

Whether or not Abies magnijica occurs on Shasta is a question on
which we can throw no light. I do not know how to tell magnifica
from shastensis except by the cones, and the trees did not bear cones the
year of our visit.^ Still, we found great numbers of old cones tucked
away by the scpiirrels in decayed logs, and disconnected scales under
most of the trees where search was made, and among all these failed to
find a single bract which was not strongly exserted. And yet Miss

' While this paper was passing through the press (July, 1899), Walter K. Fisher
revisited Shasta. He found the firs heavily laden with cones, and althougli thou-
sands of trees were examined he failed to find a single cone without the exserted
bracts.

Fig. 19. — Cone scales of (a) Ahies shas-
temis and (b) Abies cnncolorlowiana.

38

NORTH AMERICAN FAINA.

[NO. 16.

Alice Eastwood showed me, in the herbarium of tho California Acad-
emy of Sciences, a cone, said to have come from Wagon Camp, iu which
tlie bracts, except a few at the base, are not exserted.

Silver Pine or Mountain White Pine {PinuH monticola). — Silver
pines occur here and there on Shasta, scattered among the Shasta firs.
They were found in greatest abundance on a pumice slope south of
Brewer Creek Canyon, where they are the dominant trees up to an
altitude of 7,200 feet, and where the ground was strewn with cones of
the previous year — cones in which the scales are strongly reliexed.

V.

l'"iG. 20. — Wliitebark piue {Finns albicaulis).

In this area, along Brewer Creek, they meet and slightly overlap the
alpine hemlocks and white-bark pines of the zone above. They are
common also on the steep lava ridges on the north side of Shasta, par-
ticularly in the neighborhood of Inconstance Creek and in Mud Creek
Canyon, especially on the west side of the ridge between Mud and
Clear creeks. A few trees occur near the top of Bed Cone, east of
Wagon Camp.

Lodge-Pole Pine {Pinus murrayana). — The lodge-pole pine was
not found on Shasta except on the northeast quadrant, where Ver-

OCT., 1899.]

WHITE-RARK PINE BELT.

39

non Bailey, in following the wagon road around tlie mountain, passed
through a belt of it about 8 miles in length. It begins 3 miles
northeast of Ash Creek at an altitude of about 5,400 feet and reaches
northerly to about 3 miles northwest of Incon stance Creek, where
it ends abruptly at an altitude of o,(»00 feet. Here it is the dominant
tree, and in half of it the only tree. This area is covered during the
latter part of the afternoon by the shadow of the mountain, and conse-
quently is colder than places of equal altitude farther north or south.
The soil is sandy and barren and the trees are of small size.

(3) The Upper Belt or Belt of White-Bark Pines (Pinus alUcaulis).

Still above the forest of Shasta firs, braving its way upward over the
bare rocky ridp-^s into the very teeth of the domain of perpetual snow,
is another timber belt — an c pen belt of straggling, irregular trees, whose

Fig. 21.— Dwarf wliite-bark pines on a high ridge.

whitened, twisted trunks with tlieir storm-beaten heads of green are
among the most weirdly picturesque objects on the mountain (fig. 20).
The tree is the timberline white-bark pine, which, wherever found,
pushes its way over steep and barren slopes to the extreme upper limit
of tree growth.

At the lower part of its range it forms an almost continuous though
narrow belt around the mountain, and often attains a height of 30 or 40
feet and a diameter of 2 feet. In the higher parts of its range it soon
becomes restricted to the ridges, leaving the intervening basins and
gulches bare, and as it climbs higher and higher becomes more and
more reduced in size and undergoes material changes of form and posi-
tion. At certain altitudes the slanting trunks, only 4 or 5 feet in
height, serve as pillars to support the flattened tops which form a
canopy of intertwined and matted branches (fig. 21).

40

NORTH AMERICAN FAUNA.

[no. 16.

These dwarf groves ofter attractive shelters from wind and storm,
and we usually camped among them when working the upper sloi)e8.

The tree is one of exceptional hardihood, and as it pushes on still
farther into the realm of cold the trunks become completely prostrate
and the branches hug the ground, forming among the rocks dense mats
which sometimes rise a foot or two above the general level, but at their
upper limit usually occupy depressions, or, if growing in the lee of a
bowlder, crouch behind it and continue its surface level to the adjacent
slope, as if trimmed to lit. Indeed, one is amazed at the way these
uppermost pines avoid exposure by flattening themselves into the hol-
lows, as if afraid to lift a finger above the general level. Their life is a
perpetual straggle — not against competing plants, but against a hostile
environment. In summer they are buffeted by the winds and pelted by
sand and gravel; in spring they are swept and torn by the resistless
avalanches, and in winter they arc deeply buried under heavy banks of
snow. The prostrate trunks in young and middle-aged healthy trees
are completely concealed, often half covered by stones and pumice sand,
and hidden by the density of their own branches and foliage; but in very
old trees, and those injured by passing avalanches or laid bare by the
washing away of protecting rocks during violent storms, the trunks are
partly exposed and their extraordinary features may be easily examined.
As a rule they are not only naked, but the strongly twisted wood,
gnarled, contorted, and, ironlike in firmness, has been eaten into by
the sand blast till the knots and hardest parts stand out in prominent
ridges.

A little below timberline on the north side of Shasta, between I^orth
Gate and Shastina, is an extensive gently sloping pumice plain, strewn
with fragments of gray shaly lava, and thickly sj)otted with rather

Fig. 22.— Pumice plain north side of Shasta, showing timberline mats of white-bark pines.

large mats of pines, averaging 2 to 4 feet in height, which give a most
curious aspect to the region (fig. 22). This area, which is about a mile

OCT., 1899.]

WHITE-BARK PINE BELT.

41

and a half across (horizontally), must contain thousands of acres of the
dwarf, flattened pines. Along its lower edge, singularly enough, trees
of the same species suddenly staiul upright and grow to large size,
forming a rather solid forest, perhaps 30 feet in height, with an abrupt
front facing the dwarf pines above. The suddenness of the transition
is unusual and difficult to explain.

The forest just mentioned is probably the largest continuous area of
Pinus (dbicaulis on Shasta. Situated a little below timberline, it
stretches, a^jparently without interrui^tion, from North Gate Buttes
to Diller Canyon, a distance of fully 5 miles, thus encircling the north-
west quadrant of the mountain, including Shastina.

Perhaps the most attractive grove of white-bark pines on Shasta is
one that fills an open gulch or glade on the east side of i^orth Gate
Bnttes. Here, in the lower i:>art of their belt, the trees are large and
uncommonly symmetrical, and the gray pumice soil is covered with
silvery lupines. In ascending the gulch the pines gradually decrease
in size until at 'The Gate' (alt. 8,500 feet) they are dwarfed and their
tops are broadly flattened.

The normal altitudinal limits of the white-bark pines on Shasta are
hard to fix. On the south and southwest sides the trees descend in
places to 7,500 feet and range thence upward on the hottest ridges to
an extreme limit of 0,800 feet. But this extreme altitude is attained at
two jjoints only — on the long ridge above 'The [South] Gate ' (near Bed
Butte) and on a ridge about a quarter of a mile west of Mud Creek
Canyon. On the west rim of the canyon the pines stop at 9,500 feet

Fig. 23. — A lari;<- jimstratf tree of wl!itf-l)ark i)iiie. a little below tiiiiljerline.

and on the ridge on the east side at 8,600. Probably 9,300 to 9,500
would be a fair average for their upper limit on the warmer southerly
slopes.
On the cold northeast slope, just south of Brewer Creek, they descend
21753— No. 16 6

42

NORTH AMEKICAN FAUNA.

on a barren pumice slope to 7,000 feet, where, si)ariugly mixed with
alpine hemlocks, they meet the upper limit of Shasta firs and silver
pines.

Black Alpine Hemlock {T.siiga vwriensiana'^ ). — But the white-
bark pine, although the dominant and most widely distributed tree of
the upper timber belt, is not the only tree, for in places it is mixed
with or replaced by the black alpine hemlock. Shasta is a very dry
mountiiin, and yet the white-bark pine thrives on its driest slopes and
grows among the bare, naked blocks of lava where tree life seems
imj)ossible. The hemlock requires more moisture, and therefore is at a
decided disadvantage. It never reaches as high as Finns aJhieauUs
and attains its best development along the lower border of the Hud-

FlG. 24. — ISlaik alpine hemlocks near Squaw Creek.

sonian zone, where it occurs in disconnected sheltered localities —
usually iu canyons or on the shady east or northeast sides of buttes or
ridges, where there is more moisture than on the exposed slopes. Since
these shady easterly slopes are always cold, the hemlocks that occupy

'This is the sjiecies heretofore commonly l^nowu as Tsuga pattoni or Tsuga patto-
niana. It has heeu recently discovered that the name Tsuga meriensiana, commonly
applied to the Pacific lowland hemlock, was lirst given to the present alpine species,
necessitating a most nnhappy change of name. Fortunately, however, the common
English names of the two and their Avidely different zone ranges — one restricted t"
the low Transition belt near the coast, the other to the high Hudsonian zone on the
loftiest mountains — may prevent the confusion that otherwise would result from the
change of name.

OCT., 1899.]

WHITE-BARK PINE BELT,

43

ttem (leseeiid in tongues considerably below tlie usual lower limit of
the belt to whioli tbey belong. The most extreme case of tlie kind
observed is on the east side of the series of hills and ridges known on
the map as 'Gray Butte,' where a gulch, sheltered from the warm after-
noon sun and moistened by seepage from melting snows, carries the
hemlocks to a lower altitude than they reach elsewhere. On suitable
slopes they usually begin about 7,200 or 7,300 feet and range up to
jvbout 8,000 feet. The highest altitude at which they were observed is
8,700 feet, a little east of Mud Creek Canyon, where a few stunted trees
were found among the white-bark pines. Their extreme upper limit is
thus a thousand feet lower than that of the white- bark pines. This is

Fig. 25. — Group of alpine, hemlocks near Deer Canyon.

due, in part at least, to the character of the upper slopes, where no
trees can grow except on the ridges — as explained under the head of
Timberline (pp. 27-30) — and here the ridges are too exposed and too
dry for hemlocks.

On Shasta the alpine hemlock does not grow in such luxuriance
or attain such dimensions as in the Cascade Eange. The average
height of mature trees seems to be 80 or 100 feet; the average diameter
a little less than 3 feet. Trunks 4 and 5 feet through are by no means
rare and the one shown in the accompanying i^hotograph (fig. 26) meas-
ured C feet. It is a characteristic habit of hemlocks on sloping ground
to grow in clusters, 3 to 7 springing from a common base. In this way,
when young, they are better able to withstand the pressure of the snow.

44

NORTH AMERICAN FAUNA.

[NO. IC.

Those that grow singly usually supi)ort tlieniselves by having the
trunks strongly curved downward just above the ground, as shown in
fig. 20.

The alpine hendock is one of the most ])icturesque and attractive
trees of our western mountains. Its beauty is due in part to the hand-
some trunks and irregular drooping branches, but mainly to the dense
and peculiarly tufted foliage which falls in graceful masses in such
manner as to conceal the branches and upper parts of the trunks. The
twigs or ultimate branchlets curve upward and the needles stand out
on all sides and point outward — away from the body of the tree — i)ro-
ducing a tufted appearance very unlike that of other conifers. This

Fig liG.— Trunk of alpinr henilock, .sliowiiig tliiikrniiig and curvature of hase.

effect is heightened during the latter part of summer by the light green
tips of the new growth.

The hemlock forest is dark, somber, and silent, and its drooping
branches are draped with the dangling beards of the black-beard
lichen [Alectoria fremonii). The only color that breaks the otherwise
uniform darkness is the bright yellow lichen [Evernia vulpina) which
covers the north or northwest sides of the trunks and upper sides of
the branches— a conspicuous feature and one seemingly out of harmony
with the general tone of the forest. In all these respects the hemlock
forest accentuates the features of the Shasta fir forest of the belt below,
with which it is directly continuous. The tree trunks and bark also
resemble those of the Shasta firs. The bark is in long scales which, as

OCT., 1899.

WHITE-BARK PINE BELT.

45

the trees grow, thicken into irregular jtlates transversely broken at
intervals of S to 12 inches.

In the growing tree the branches die from below upward in a curious
way. First a subdivision of a low branch dies and the tips curl down-
ward and inward, drawing together until they form a close curl or
tail which can be set on tire by a single match. Dozens of these curls
can be seen on most of the young trees, and also on the lowermost
remaining branches of the middle-sized and some of the old ones. This
process of dying and curling continues until all the lower branches are
dead. Meanwhile, the curl-tails gradually drop off and litter the
ground, leaving the bare dead branches hanging down at a sharp angle.
These dead branches hug the trunks closer than the living ones and
cling on until the bark comes off, when they form an armature of

Fig. 27. — Group of alpine hemlocks.

unsightly bleached and brittle sticks poiuting downward around the
trunk. These in time break off, too, so that as the tree grows into
maturity the handsome trunk finally becomes clear and clean.

The alpine hemlocks are prolific bearers and the ground is always
• strewn with their cast-off cones, which average about 2^ inches in
length, and have a dark streak down the middle of each scale. When
young the cones are conical, when old and the scales become fully
retlexed they are slender, subcylindrical and only three-fourths of an
inch in diameter. Year after year the cones fall to the ground in
such prodigious numbers that they form a very important part in
the layer of felting that covers the surface in the hemlock forests —
a loose dark felting composed of disintegrating needles, twigs, and cone
scales pressed firmly together by the weight of the snow iu winter, and

46 NORTH AMERICAN FAUNA. [no.16.

only rarely dotted by living i)l;ints. In moist spots, particularly along
the borders of the tiny sparkling streams, the red heather [Phyllodoce
empetriformis) forms little beds aiid the delicate feathery Lutkea peeti-
nata spreads a faint veil of green over the dark soil. In the drier
parts of the forest hardly a plant is seen save now and then a solitary
clump of prince's pine [ChimapMla menziesl) or painted wintergreeii
[Pyrola picia).

Late in September the hemlocks molt and the wind brings down show-
ers of needles that falling on the tent at night sound like rain. Their
color has now changed from green to golden brown and they sprinkle
the black floor of the forest so thickly as to change its appearance.

FOREST FIRES.

During the past ten years the country about Shasta, particularly on
the west aiid south, has been repeatedly devastated by forest fires.
Here, as elsewhere, lumbermen and fires have destroyed the greater
part of the timber on the lower slopes and adjacent plain, which are
now covered by a dense chai)arral of manzanita and buckbrush, dotted
with scattered pines. Fortunately, the fires have not as yet spread
upward far enough to do much damage to the Shasta firs of the middle
timber belt. Whether the character of these trees and the freedom of
the ground beneath from combustible material will prevent the spread
of fire remains to be seen. Thus far the greatest harm has been done
in the forests of ponderosa and sugar pines, where lumbering opera-
tions are being carried on with painful vigor.

While we were on the mountain, from the middle of July until the
end of September, one or more fires, the result of vandalism or neglect,
were raging continuously on the south and west sloj)es, and two of them
did irreparable injury. One began near some woodcutters' shanties, 3
or 4 miles below Wagon Camp, on the road to Sisson; the other and
more destructive originated in the area covered by the lumbering opera
tions from McCloud Mill and pushed swiftly up the Panther Creek
slope, consuming the greater part of the only area of Pinus attemiata
on Shasta and burning great tongues into the handsome fir forest on
both sides of Wagon Camp, which it closely and almost comj^letely
surrounded.

The fire that lasted longest in the summer of 1898 did the least harm.
It consumed a worthless tract of manzanita chaparral between Black
Butte and the mountain, and gave off a surprisingly enormous quantity
of smoke, hiding the country to the west for a full month. During its
continuance the entire mountain was often enveloped in smoke and
when the wind was northwest, as it was a great deal of the time,
showers of burned leaves fell daily at our camps. On August 2, when we
were at work on the rocky slopes above the head of Squaw Creek at an
altitude of 9,500 feet, charred leaves fell so abundantly that we caught
many in our hands. Great clouds of smoke rolled up between us and

OCT., 18911.] SLOPE EXPOSURE. 47

tbe sun, which became deep red like the full uioon and then disappeared.
At 5 o'clock the smoke began to settle back, as it always did when the
chill of the evening came on, and the sun reappeared, to set as usual
behind the dark outlines of the distant mountains.

Fires ou the south, in the valley of the McCloud, cut off the view in
that direction, and it was only at rare intervals, and usually at sun-
set, that we were able to see the snowy crown of Lassen Butte 00
miles away. Even Castle Crags, almost at our feet, were rarely visible.
This experience is frequent in the west; and of the hundreds of persons
who visit the Pacific slope every summer to see the mountains, few see
more than the immediate foreground and a haze of smoke which even
the strongest glass is unable to penetrate.

Along the railroad between the head of Sacramento Canyon and
Shasta Valley one traverses desolate tracts which a few years ago were
covered by a noble forest of ponderosa and sugar pines.

EFFECTS OF BURNS IN CHAN(rING ZONE POSITIONS.

A burn in the lower part of the Shasta fir forest a little above
Wagon Camp affords an excellent illustration of the way fires some-
tnnes change the zone relations of particular areas. The area in ques-
tion was well within the Canadian zone before the fire, which occurred
only a few years ago. Since the fire. Transition zone species have crept
up the ridge, and now Ceanothvs velufinus, Arctostaphylos j>atula, Hot-
l-elia pseud occqnta ta , Apoci/n n m p7( m il ion , Gayophyt nm ramosi.ssim um, and
others are common. The manzanita and buck-brush are young and
small but are spreading, so that in a few years the ridge, whi(;h has a
warm southerly slope, will be mainly Transition. But in the mean-
time a new growth of Shasta firs has started, and in ten or twenty
years is likely to overtop and drown out the Transition zone species,
enabling the Canadian zone to reclaim the burn.

Such cases of alternation of zones resulting from the clearing of
forest land are not uncommon, particularly when deforestation is
caused or accompanied by fire. But on steei)er sloi)es, especially rock
slopes, if the vegetable layer is burned oft", the (lower) zone wliich
creeps up to replace the (higher) one destroyed becomes permanent or
nearly so.

It may be laid down as a general rule that the destruction of forests,
by admitting the sun and wind, lessens the moisture in the soil and
increases the temperature, thus inviting animals and plants to come in
from adjacent warmer areas. Deforestation of an area therefore tends
to lower its zone position.

SLOPE EXPOSURE.

By slope exposure is meant the inclination of the earth's surface at
a particular point with reference to the angle at which it receives the
sun's rays. The amount of heat, were it not for the retarding effect of

48

NORTH AMERICAN FAUNA.

[no. 16.

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-Diagram .showing average hourly march
of temperature.

the atmosphere, would be greatest at noon, when the sun's position
with reference to the earth is most nearly vertical. But, as everyone
knows, the atmosphere becomes heated slowly and does not attain its
highest temperature until the middle or latter part of the afternoon,
the hour varying with the locality. Hence slopes that face the sun
most nearly at a right angle at the time of day when the atmosphere
is hottest are naturally the hottest slo])es — those that carry the zones
highest; while conversely, slopes that face the opposite direction are
naturally the coldest slopes — those that depress the life zones lowest.
At Portland, Oregon (about 275 miles north of Shasta), the hottest
part of the day in summer is a little after 4 o'clock in the afternoon,
at which time the sun is nearly due west. Consequently the hottest
uncomplicated slopes are those that face west or a little south of west.
The accompanying diagram shows the actual mean hourly march of
atmospheric temperature at Portland, Oregon, for June, July, and
August :

The influence of sloj^e exposure on the faunas and floras of moun-
tainous regions is profound. Measured by a scale of altitudes, it
amounts on ordinary slopes to nearly a thousand feet, and on steep
slopes is still more marked. Thus on mountains it is usual for plants
and animals of i)articular species to occur on warm southwesterly
slopes at elevations 800 to 1,000 feet higher than on cool northeasterly
slopes. Similarly on north and south ridges, the faunas and floras of
the warm west slopes often belong to lower zones than those of equal
elevations on the cool east slopes.

Shasta aftbrds innumerable examples of the eftects of slope exposure,
both simifle and complicated by canyons.

An excellent illustration of the latter is to be found in Mud Creek

OCT., 1899.] SLOPE EXPOSURE. 49

Canyon, aear the mouth of Clear Creek (see pL iii). The altitude of
the bottom of the canyon at this point is 6,700 feet, which would nat-
urally place it in the middle of the Canadian zone. The steep west
side of the narrow ridge between the two creeks receives the afternoon
sun at nearly a right angle, and is in consequence an unusually warm
slope for the altitude. The result is that seeds of plants carried upward
by winds and birds from the Transition zone, more than 1,000 feet
below, have here found a favorable resting place and have grown into
a colony of Transition zone species, among which are Abies concolor
loiciana, Arctostaphylos patula, Kunzia trideutafa, Amelanchier alnifoUa,
Kiibus parvifloru.Sj Sorhus namhueifolia, Sytnplioricarpos pilosus^ Rihes
viscosissiniuin, E. (unictum, Samhucus melanocarpa, Apocynum pumilum^
Spira'a douglasi, Yaccimmn {arhuscula ?), Lupinus elmeri, Eriogomim
marifoliiimy Gilia agf/regata, Pteris aquiluia lanuginosa, and the large
Transition zone form of Castilleja miniata. Just across the canyon,
and in one place less than 100 feet from the lower edge of this isolated
Transition colony and at a lower level, are species belonging to the
Hudsouian zone — such as Pentstemon neicherryi and Pulsatilla oeciden-
talis. Thus, growing close together on opposite slopes of the same
canyon, are species characteristic of zones both above and below the
one to which the altitude i)roi)erly belongs. This case is by no means
peculiar and is a good illustration of the simpler effects of slope exposure
commonly shown in mountain canyons.

Another class of cases is found on the buttes and hills. A mile
and a half northeast of Wagon Camj) is ;i conspicuous red cinder cone
about 1,000 feet in height, known as Ked Cone. In zone position it is
Well within the lower part of the Canadian belt where it is surrounded
by the forest of Shasta lirs, scattered trees of which push up on the
basal sloi)es and on the south side reach the toj). Just below the sum-
mit are a few silver pines belonging to the same zone. The warm south-
west slope brings up irom the Transition zone below thickets of man-
zanita, a few incense cedars, and several humbler plants. The cold
northeast slope, although reaching an altitude of only about 0,800 feet,
which would place it in the middle of the Canadian zone, is cold enough
to bring down from the Hudsouian zone several characteristic plants,
such as Polygonum neu-berryi, Cymoptcrus terehinthinus, Cycladenia
humilis, Eriogonum polypodum, and Viola purpurea.

On the south side of Shasta an interrupted ridge or series of buttes,
known collectively as ' Gray Butte,' reaches up the mountain side from
Ked Cone, east of Wagon Camp, northward to the gap known as 'The
Gate.' In a gulch or canyon on the east side of this butte the black
alpine hemlock descends more than 1,000 feet below its usual lower
limit. The reason is obvious. The row of buttes cuts off the warm
afternoon sun, and seepage from melting snows keeps the soil moister
than in more exposed places, so that the gulch meets the two require-
ments of the alpine hemlock — a moist soil and a cold atmosi)here. For
its entire length this long tongue of hemlock is flanked by Shasta firs
21753— No. 10 7

50

NORTH AMERICAN FAUNA.

[NO. 16.

from the zone below, so that the usual zone relations are chanj>ed,
parallel strips of Canadian and Hudsouian running up and down the
mountain — instead of encircling it in the usual liorizontal belts. Along
Squaw Creek another tongue of iilpine hemlock descends to the liead
of the main fall, at an altitude of about 7,25() feet, and is simihirly
sandwiched between ascending tongues of Shasta firs.

Between 'The [South] Gate' and the grove of alpine hemlocks on
upper Squaw Creek is a prominent mass of lava 700 or 800 feet high,
known as 'Eed Butte.- It is about 2,000 feet below the altitude of
extreme timberline and its summit is covered with trees; nevertheless
its precipitous northeast side is so cold that its zone i^osition is well
above timberline, as shown by the presence there of such distinctively
alpine plants as Oxyria digyna and Saxifraga tohniei. In this case the

Fiii. 'M. — i)\\ all jjiiR's oiidiiig abruptly along cold east .sido uf ridge.

effect of a very cold mass of rock is added to that of the coldest slope,
and the result is a lowering of alpine zone species 2,000 feet below their
normal elevation on the hottest southwest slopes.

The high north and south ridges afford i^erhaps the simplest example
of the direct influence of slope exposure. The warm west sides of these
ridges usually bear trees in proportion to the availability of their slopes,
while the cold east sides remain naked and alpine (see tig. 14). The way
the dwarf pines stop along the east crest of the ridges is shown in the
accompanying figure (fig. 20).

Finally, the glaciers of Shasta afford impressive evidence of the
effects of slope exposure. ^ly party did not take the altitudes of the
glaciers, but according to the Shasta map sheet of the U. S. Geological
Survey those on the cold east and northeast slopes descend below
9,000 feet, and one of them, at the head oi" Ash Creek, below 8,r)00 feet,
while the only one having a south exposure (at the head of Mud
Creek) stops at 11,000 feet,' and there are no glaciers at all on the west

' There is another glacier on the south side, tributary to Mud Creek, which
descends lower than the one marked on the map as 'Konwakiton glacier,Mnit it
is coni])letely hidden by a high ridge and is not exposed to the late afternoon sun.

o<T., 1890.] EFFECTS OF STEEP SLOPES. 51

side. Hence if the altitudes to which glaciers descend on the various
slopes be accepted as indicating the course of a sinuous line of equal
temperature, it follows that the difference in temperature dependent on
the angle and conditions of slope exposure, as measured by the glaciers,
is e(iuivaleiit to a difference of upward of 2,000 feet in altitude. But
this is doubtless excessive and due in part to local influences.

EFFECTS OF STEEP SLOPES.

Steep slopes, particulaily those that face the southwest and west,
exaggerate the effects of slope exposure. Those that ftice the hot after-
noon sun at nearly a right angle receive the greatest quantity of heat,
but this alone is not sufficient to account for the very extraordinary
degree to which the fauna and flora are sometimes affected. When it is
remembered that the hottest ordinary slopes carry up the zones only
800 to 1,000 feet, one is startled to find that on some favorable steep
slopes they are pushed up more than 2,000 teet above their normal
limits. The explanation did not occur to me until, in discussing the
matter with the geologist, G. K. Gilbert, he suggested the diurnal ascend-
ing current as the missing factor.

It is well liUown that in ordinary calm weather the air currents on
mountain sides and in deep canj-ons ascend by day and descend by
night. The ascending currents are warm, the descending currents cold.
The night current, being in the main free from local influences that
affect its temperature, must exert an essentially equal effect on all sides
of a mountain; but the temperature of the ascending day current, being
constantly exposed to and in fact created by the influence of the sun,
must vary enormously on different slopes. The activity and effective-
ness of this current increases with the steepness of the slope and the
directness of its exposure to the afternoon sun. Hence the hottest
normal slopes — those that face the sun at nearly a right angle during
the hottest part of the day — are rendered still more potent by increased
steepness, the direct exposure to the sun keeping up the supply of heat
while the steepness of the slope accelerates the rate of movement of the
diurnal ascending current, carrying the heated air upward a very great
distance before it has time to be cooled by the general temperature of
the stratum it penetrates. Thus it is that species characteristic of the
Transition zone on Shasta — species which on normal southwesterly
slopes attain their upper limits at an altitude of 5,500 to 5,700 feet — are
in favorable places enabled to live at elevations of 7,900 and even 8,000
feet, considerably more than 2,000 feet above their normal upper limits.

The steep slopes of Diller Canyon furnish instructive illustrations of
the effects of these ascending hot-air currents. Here, on the hot stony
pumice slopes, such distinctive Transition zone species as Arctostaphylos
patula, Kunzia tridentata, Ceanothus velutinus, and Chrysothamnus-occi-
dentaUs flourish among the Shasta flrs and white-bark pines at an alti-
tude of nearly 8,000 feet in the belt where the Canadian and Hudsonian
zones overlap, and more than 2,000 feet above the extreme upper limit
of their normal distribution on uncomplicated hot southwesterly slopes.

52

NORTH AMERICAN FAUNA.

[NO. 16.

J5AS1N SLOPES.

Many of the glacial basius of the timberline region are broad U-shaped
depressions with gently sloping bottoms, ending abruptly in terminal
moraines, below which they may or may not continue on to other
moraines. Tbey were excavated by glaciers at a period when the ice
cap of Shasta was much larger and more complete than at present.
The upper ends of most of these valleys abut against the steep upper
slopes of the peak, and are bordered on both sides by lofty ridges, so
that they are walled in on three sides and thus converted into basins.
Such basins, when they face the southwest, appear to promote the
reflection of heat and retard the escape of hot air, so that they some-
times become hot pockets characterized by species belonging to the zone
below.

LIFE ZONES OF SHASTA.

Shasta stands on a Transition zone plane, with a dilute tongue of
Upper Sonoran approaching its northern base by way of Klamath and
Shasta valleys. Its forested slopes rise quickly through the Boreal
zones to timberline, above which its ice-clad summit towers to a height
of 5,000 feet. The life zones of Shasta, therefore, beginning with the
Upper Sonoran element of Shasta Valley, are —

Upper Sonoran Hudsonian

Transition Arctic-Alpine.

Canadian

In a generalized diagrammatic north and south section of the moun-
tain the relations of these zones may be shown somewhat as fellows:

VaVaV/^Va'^aVa

V A

Fig. 30.— Diagram of Sliaata showing relations of life zones.

The altitudes of zone boundaries here given are intended to repre
sent their average or mean elevation on normal southerly slopes. The
aridity of the mountain as a whole, with consequent scattered or
'spotty' instead of 'continuous' distribution of most of its zone species,
complicated by the influences of hot and cold slopes, springs, and air
currents, elsewhere discussed, which frequently carry species 1,000 feet
or more above or below their normal limits, makes it almost impossible

OCT., 1899] UPPER SOXORAN ZONE. 53

to fix with certainty the normal zone altitudes. Hence those here given
must be regarded as approximate only. The average width of the belt
of overlapping of adjoining zones appears to be about 800 feet; so that
the normal vertical distribution of the distinctive species of each zone
overreaches in both directions the altitudes given by about 400 feet.

The zone positions accorded the various species are based on per-
sonal field experience, and in the great majority of cases are believed to
be correct; in a few instances, however, the evidence is inconclusive —
hence the zone lists must be regarded as provisional and subject to
correction. Species recently exterminated or driven away, as the moun-
tain sheep, elk, and grizzly bear, are included in the tables.

UPPER SONORAN ZONE.

The Upi^er Sonorau element in the region about Shasta is dilute and
is limited to Shasta Valley at the north base of the mountain, which it
reaches by way of the Klamath country on the north and northeast.
It has no connection whatever with the Upper Sonorau of the Sacra-
mento Valley on the south, which ends near the point where Pitt Eiver
joins the Sacramento, about 50 miles below Shasta.

Shasta Valley is an arid plain about ii5 miles in length by 10 or 12 in
breadth, studded with small volcanic buttes and lava flows. It varies
in altitude from a little less than 4,000 to about 2,700 feet, and is lowest
at the north, where the Shasta Eiver, by which it is traversed, flows
into the Klamath Eiver. Shasta Kiver is in places bordered by decid-
uous trees, mainly Poj)uIhs U'ichocarpa, Betiila occidcnfali.s, and Alnus
rhomhifolia. The valley is sparsely covered with rabbit brush of sev-
eral species, mainly Chrysothnmnus occidentalis and C. viscidiflorus, and
in some places, chiefly on the east and south, by the true sagebrush
[Artemisia tridentaia). The southeastern part contains an open forest
of junipers {Juniperns occidentalis), which meets the pine forest of the
basal slopes of the mountain. The western part is an open plain very
scantily dotted with rabbit brush and a few small and mainly incon-
spicuous plants, among which were observed: Mentzelia hvincaulis,
Xanthium strnmarium, Heliotropium curassavicum, Verbena bracteosa,
Datura meteloides, barcohatus rermiculatus and Nicotiana sp. f Along
the western border of the valley, near the Scott Mountains, several
characteristic shrubs occur, among which are Ceanothus cxneatus, Arcto-
staphyJos [viscida?), Cercocarpus parrifolitis, Eriodictyo7ighitinosum, and
Garrya fremon ti. '

The zone position of this region is further indicated by the presence
of such birds as the chat {Icteria virens lonyicauda), bush-tit {Psaltri-
pariis minimus californiciis), California towhee [Pipilo fuscus crissaUs),
California jay {Aphelocoma caiifornica), aud California valley quail
{Lophortyx californicus vallicola). Formerly antelope were common
here, but now they are rarely seen. The most abundant mammals are
jack rabbits {Lepus californicus), sagebrush cottontails [Lepus nut

' Garrya fremonti is both Upper Sonorau and Transition.

54 NORTH AMERICAN FAINA. [m.u.\

talli), kangaroo rats {IHpodomys californicns), i)ocket mice {Perof/mttlim
parvus f), ^YOO^[Tixts {Neoioma fuscipes), coyotes {Canin ochropus), and
raccoons {Procyon psora pacifica).

So little work was done in Shasta Valley that the species mentioned
are of course only a fraction of those present.

MAMMALS OF I'PPEU SONOKAN ZONE (IN SHASTA VALLEY).

liestrictcd to Vjrper Sonoran.

Lepus calilornicus. Perognatbus (parvus'!').

Lepus uuttalli. Peroiiiyscns trnei.

MicTotus califoinicus. Eeitbrodontoiuys klamp^thensis.

Oomtnon to V'j}per Sonoran und Transition.

Antilocapra americana. Procyon psora pacifica.

Canis ocLropus. Scapanns califovnicns.

Dipodomj's califoruicus. Speriiiophilns douglasi.

IMeplaitis occidental is. Si)ilogale latifrons.

Myotis evotis. Taxidea taxus.

Neotoma fnscipes. Vespertilio fuscus.

Peromyscus gambeli. ?Zapu8 pacificus.

BIKDS OF TTPPER SONORAN ZONE (IN SHASTA VALLEY).
Restricted to V-pper Sonoran.
C'atberpes luexicanus pimitubxtus. Lanius liidovicianus excnbitorides.

Chondestes grammacus strigatus. Otocoris alpestris merrilli.

Clyanocepbaliis cyauocepbalus.' Pipilo fnscns crissalis.

P"'alco mexicanus. Psaltripanis minimus calil'ornicns.

Coninion to Upper Sonoran and Transition.

Apbelocoma califoruica. Melanerpes formicivorus bairdi.

Astragaliuns psaltria. Phabi'uoptilus uuttalli.

Piutco swaiusoni. Pipilo maculatns megalonyx.

Calypto auna. Scolecopbagus cyanocepbalus.

Carpodacus mexicanus obscurus. S^jeotyto cuuicularia bypoga^a.

Catbartes aura. Spizella socialis arizonae.

( "bondestes grammacus strigatus. Sturnella magna neglecta.

Cbordeiles virginiauus. Tbryomanes bewicki spilnrus.

Cyanospiza ama-ua. Troglodytes acdon parkmaui.

Dendroica a'Stiva. Tyranuus verticalis.

Geothlypis tricbas occidentalis. Vireo gilvus swainsoni.

Icteria virens longicauda. Zeuaidura macroura.
Lopbortyx calitbrnicus vallicola.

TRANSITION ZONE.

Exceptinu' Shasta Valley, the Transition zone covers the whole coun-
try about Shasta and pushes up over the basal slopes to an altitude of
5,000 or 0,000 feet. To the northeast and east it reaches and extends
beyond the Klamath country and Goose Lake in Oregon, and the
^Madeline Plains in extreme northeastern California, interrupted only
by narrow tongues of Upper Sonoran in the ui)per Pitt liiver Valley,
and by small islands of Canadian on the highest mountain summits.

' Tbe pifion jay visits the juniper forests in Shasta Valley in fall to feed on the
juniper berries, but whether or not it breeds there is not known.

OCT., 1899.1 TRANSITION ZONE. 55

To the south the Transition zone tills the McCloud and Pitt River val-
leys, embraces the canyon of the Sacramento, aud stretches onward
along the flanks of the Sierra all the way to southern California. To
the west it overspreads the wild mountain region l)etween Shasta and
the Pacitic Ocean, changing gradually from Arid Transition to Humid
Transition, and surrounding the Upper Sonoran bottoms of Scott and
Hoopa valleys, and the Boreal summits of Salmon, Trinity, and Siski-
you mountains. It covers the lower slopes and eastern ])art of the
Siskiyous, and passes around the southern ends of the Salmon and
Trinity mountains continuously to the sea.

On the lianks of Shasta the Transition zone forms a broad continuous
belt covering the basal slopes on the northwest, west, and south, but
interrupted on the cold east and northeast sides by the Boreal, which
here pushes down to the actual base of the mountain, crowding the
Transition out to the east around a group of low volcanic hills.

On the southwest and west it pushes up on ordinary slopes to
5,500 or 6,000 feet, rising on steep pumice canyon slopes a couple of
thousand feet higher, and everywhere embracing tongues of Canadian
which descend along the cold streams and on the cold easterly slopes
of ridges.

On Shasta the study of the Transition zone is complicated by strong
local ditierences of soil-moisture and humidity — differences that exert
a profound effect on the distribution of plants, and to a less degree on
that of animals also. It has been shown elsewhere (Life Zones and Cioj)
Zones of the United States, p. 28, September. 1898) that in some places
the Arid Transition of the Rocky Mountains and Great Basin passes
gradually into the Tlumid Transition of the Pacific coast. On Shasta
similar changes occur in such small compass that they may be studied to
excellent advantage. Thus, near the south end of Shasta Valley the
dominant types of vegetation are Pinus ponderosa^ (^uerviis californica^
Artemisia tridentata, Kunzia tridentata^ Arctostaphylos pafiila, Chryso-
thamnuH oceidentalis, Rhus trilohata, (hirri/a freinonti, and Pruiius snb-
cordata, all characteristic Arid Transition species. On moister soils
near by, particularly in shady canyons, the dominant types are Pseu-
dotsit(/(( mucronata, Abies coneoJor lowicDut, Acer glah rum, Cornus 'mdtaUi^
Riihus parviflorus {=nKUam(s Auct.) Spirwa douglasr and other Humid
Transition species.

In this connection it is important to bear in mind that the extreme
bottom of the west slope of Shasta, between the Sacramento River and
Shasta Valley (elevation about 3,400 feet), is decidedly cooler and more
boreal than tlie middle part of the slope J, 000 or 2,000 feet higher.
The reason is twofold: The bottom ])art of the west slo])e, from the
head of the Sacramento Canyon northward, lies close to the east base of
Mount Eddy and the Scott Mountains, by which it is shielded from the
late afternoon sun, and consequently receives less heat than higher
parts of the same slope. Furthermore, it is well watered, an<l the rapid
evaporation caused by the dry atmosphere tends to lower the tern-

56 NORTH AMERICAN FAUNA. [no. 16.

peratiire still more. Higher parts of tbe same slope are not only very
much drier, but are exposed to the direct rays of the afternoon sun,
which, owing to the increasing steepness, strike the ground more nearly
at a right angle than at lower elevations, the result being a material
increase in the amount of heat received. It is obvious, therefore, that
on tbe southwest and west slopes the middle part of the Transition
zone is the hottest and driest, the part in which low Transition or even
Upper Sonoran species are most apt to be found, while the up^jer and
lower parts are coolest and dampest, the parts most likely to be invaded
by Canadian zone species. Hence it is not surprising that Douglas and
white lirs, willows, alders, elder, snowberry, red spinea, osier, and thim-
bleberry flourish at the top and bottom of the Transition slope, but are
absent from the middle part. At the extreme foot of the slope, along
the cool streams near Sisson Tavern, two normally boreal plants occur
which were not found at all in the boreal belts on the mountain. These
are Limuva borealis and Pachi/stima myrsinites. With them is associ-
ated the mountain chinquapin {CasUtnopsis sempervirens). There is
something peculiar about the distribution of rachystima. It is abund-
ant in the neighborhood of Sisson Tavern and occurs here and there,
apparently on dry soil, up to an altitude of 4,700 feet, but not above.
Since all three of these species are boreal, it is possible that the forms
here referred to are Transition zone subspecies.

MAMMALS OF TKANSITION ZONE.

(1) Species known to occur in the Transition zone of Shasta.

Autilocapra ameiicana. Odocoileua columbianus.

Bassariscus astutus raptor. Perognathus nioUipilosua.

Callospermopbihis cbrysodeirns. Peromyscus boyli.

Cauis oehropus. Peromyscus gambeli.

Castor cauadensis.' Procyon psora pacifica.

Cervus occidentalis. Scapanns cnlifornicus.

Dipodoiuys calilViruicus. Sciuropterus (klamatbensis?).

Eretbizon epixantbus. Scinnis albolimbatus.

E'ltaniias auid-nus. Sciurus fossor.

Eutauiias senex. Sorex montt'reyensis.

Felis oregouensis. Sorex vagrans amd'uus.

Lxitra budsonica. Sperniopbilus dougbisi.

LutreoLa vison. Spilogalc latifrous.

Lynx fasciatus pallesoens. Taxidea taxus.

Mepbitis occidentalis. Tbomomys monticobi piuetorum.

Microtns montanus. Urocyon califomicus towuseudi.

Microtus mordax.' Ursns americanus.

Myotis evotis. Ursus borribilis.

Myotis hu'ifngns longicrus. Vespertilio fuscns.

Neotoma cinerea. Vulpes macronrus.-

Neotonia fuscipes. Zapus paciticus.

Neurotricbns gil)bsi major.

' Restricted to cold streams whicb tboiigb traversing tbe Transition zone afford
Boreal temperatures.

- Probably does not breed below Canadian.

OCT., 1899.]

TRANSITION ZONE.

67

MAMMALS oi" TRAN.siTiON ZONE — Continued.

{'2') Restricted to Transition ~ove.

Bassariscus astntiTS raptor.
Lynx fasciatus palU-sceus.
fMepliitis occideutalis.

fMicrotns luontanns.

tSciurus I'ossor.

I'rocyon californions townscndi.

(3) Common to Tra7isition and Upper Sonoran zones
[See p. 54.]
(4) Common to Transition and Canadian zones.

Callo.«!permophilns chrysodeiriia.

Cervus occidentalis.

Eretliizon epixaiitlius.

Entainias amcpniis.

Eutamias senex.

Felis oregonensis.

Lutra hudsonica.

Lutreola vison energiimenos.

Mierotus mordax.

Myotis lucifugns longicrus.

Neotoma ciuerea.

Neurotriclius gibbsi major.

Odocoileus columbiauus.

Peroguathns mollipilosus.

Peromysfus Itoyli.

Peromyscus gambeli.

Scapanus califoruicus.

Sciuroiiterus alpinus klauiathensis?

Scinrus an>olinibatn.s.

Sorex moutereyensis.

Sorex vagrans amo'iius.

Taxidea taxns.

Ursus americauus,

Ursns borribilis.

Vulpes macrourus.'

BIRDS OF TRANSITION ZONK.

(1) Species known to occur in Transition zone of Shasta,

Aphelocoma californica.
A(juila chrysai'tos.
Astragaliuns psaltria.
Astragalinus tristis salicauians.
Bubo virginianus.
Biitco borealis calurus.
Buteo swuinsoni.
C'alypte anua.

Carpodacus uiexicanns obscurus.
Catbartes aura.

Ceophla'us pileatus abieticola.
Certhia fannliaris occidentalis.
Chiptura vauxi.

Choudfstes grammacus strigatus.
Chordeiles virginianus.
Circus budsonius.
Colaptes cafer.
' Contopus borealis.
Contopus ricbardsoni.
Cyanocitta stelleri.
Cyauospiza amn-na.
Dendragapns obscurus fuliginosus.
Deudroica a-stiva.
Dendroica auduboni.
Dryobates pubescens gairdneri.
Dryobates villosus hyloscopns.

Empidonax difflcilis.
Enipidonax bammondi.
Empidonax wrigbti.
Falco sparvorius.
Geotblypis tolmiei.
Geotblypis trichas occidentalis.
Glancidium gnoma califoruicuni.
Helminthopliila celata lutt'scens.
Helmintbopbila rubricapilla gutturalis.
Hirundo erythrogastra.
Hylocichla aonalascbka- audul)oui.
Icteria virens longicauda.
Lopbortyx californicus vallicola.
Melanerpes formicivorus bairdi.
Melanerpes torquatus.
Melospiza lincolni.
Melospiza melodia moutana.
Merula migratoria propinqua.
Oreortyx pictus plumiferus.
Oreospiza cblornra.
Passerella iliaca megarbyncba.
Petrocbelidon lunifrons.
Pbala-noptilus uuttalli.
Pipilo maculatus megalonyx.
Piranga ludoviciaua.
Scolecophagus cyanocepbalus.

' Supposed to breed in Canadian and wander over Transition.
21753— No. 16 8

58

NORTH AMERICAN FAUNA.

fxo. 16.

BIRDS OF TRANSITION zoNK--continne(l.
{!) Species known to occur in Transition zone of Shasta — Continued.

Selasphorns rufus.

Sialia mexicana ofcidentalis.

Sitta carolinensis acnleata.

Sitta pygmaja.

Speotyto cnnicnlaria hypoga-a,

Spliyrapicns rnhcr.

Spizella sociali.s arizona".

Stellula calliope.

Stnrnella magna neglecta.

TacliYciueta. bicolor.

Tachycineta thalassina.
Thryomaues bewicki spilnrus.
Troglodytes ai'dou parkinani.
Tyrannus verticalis.
Vireo gilvus swaiiisoui.
Vireo solitariiis cassiiii.
fWilsouia pnsilla pileolata.
Xenopicns albolarvatus.
Zam(dodia nielanocephala.
Zenaidura. macroura.

(2) llestricted to Transition zone.

fChjetura vauxi.
Contopus ricb-ardsoui.
Dryol)ates piibescciis gairdneri.
Glaucidiuin gnoiiia caliloruicnui.
^Melant'ippis tonpiatu.s.
Oreospiza chlornra.

Passerella iliaca megarhyncha.
Sialia mexicana oci-identalis. '
Sitta pygma'a.
Sphyrapicus ruber.
Vii"eo solitari'is cassini.
Zamelodia melanocepbala. '

(3) Common to Transition and Upper Sonoran zones.

[See p. 54.]

(4) Common to Transition and Canadian zones.

Aqnila cbrysai'tos.
Bubo virginianus.
Butco borealis calurns.
Ceophlceus pileatus abieticola.
Certhia familiaris occidentalis.
Chordeiles Anrginianus.
Colaptes cafer.
Contopus borealis.
Cyauocitta stelleri.
Dendragapus obscurus fuliginosus.
Dendroica auduboui.
Dryobates villosus byloscopus.
Empidonax difficilis.
Enipidonax bammoudi.
Empidonax wrighti.

Falco spar\crius.

Geotblypis tolmiei.

Helmintbopbila celata lutescens.

Helmiutbophila rubricapilla guttnralis.

Hylocicbla aonalascbka' auduboui.

Melospiza lincolni.

Merula migratoria propinqua.

Oreortys pictus plumiferus.

Piranga ludoviciana.

Selaspborus rufus.

Sitta carolinensis acnleata.

Stellula calliope.

Wilsonia pnsilla pileolata.

Xenoi)icu8 albolarvatus.

PLANTS OF TRANSITION ZONE.

(1) Species Inown to occur in the Transition zone of Shasta.

Abies concolor lowiaua.
Acer circinatum.
Acer glabrnin.
Acer macrophyllum.
Acbillea lanulosa.
Ahius tenuifolia.
Amelanchier alnilolia.
Anapbalis margaritacea.

Antenuaria geyeri.
Apocynum pnmilum.
Aquilegia truncata.
Arctostapbylos patnla.
Artemisia ludoviciana.
Artemisia tridentata.
Asarum bartwegi.
Botula occidentalis.

Probably common to Transition and Upper Sonoran.

OCT., 1899.]

TRANSITION ZONE.

51)

PLANTS OF TRANSITION ZONE — COUtiuUcd.

(1) Species k7iotin) to occur in Transition zone of Shasta — Contiimctl.

Carnm gairdneri.

Castanopsis sempervirens.

Ceanotlius cordulatiis.

Ceauotlius iutegerrimvis.

Ceanotlius (Cerastes) prostratus.

Ceanothus velutinus.

Cerasus dcmissa.

Cerasiis emarginata.

Cerasus glandulosa.

Ceieocarpiis ledifolius.

CtTcocarpns parvifolius.

Cliaiiia'Siiracha uana.

Cliimapbila menziesi.

Cbimaphila umbellata.

Cbrysamphora californica.

Cbrysotbamuiis bloomeri angustatns.

CbrysotbaiuDiis occidentalis.

Coiuus iiiittalli.

Coriuis pubescens.

(. orylus rostrata californica,

Crataegus riviilaris.

Cryptauthe geminata.

Epilobiuni brevistylum.

Eiiilobium oregoneuse.

Eiiilobiuiii spicatuin.

Eriogonniii marifolium.

Eriogonum uudiim.

Eupatdrium occidentale.

Fragarta bracteata.

Fragaria. cbiloensis.

Fraxinus oregaua.

Fritillaria atropnriuirea.

Gayopbytum raniosis.siuuim.

Gilia aggregata.

Gilia (Colloniia) granditlora.

Hastiiigsia alba.

Heleniastriim rivulare.

Heracleum lanatum.

Hieraciiim albiflorum (large form).

Hieracium cynoglossoides nudicaule.

Hieracium greeuei.

Horkelia pseudocapitata.

Jnniperns occidentalis.

Kuuzia trideutata.

Lajipula nervosa.

Liboceilrus deciirrens.

Lilium wasbingtoniauum.

Liniuca borealis.

Limini lewisi.

Lotus americauus.

Lupiuus elnieri.

Lupiuus minimus.

Macb;i'rantbera sbastensis (large form)

Mimulus moniliformis.

Mimulus tilingi.

Osmorrbiza nuda.

Pacbystima myrsinites.

Fieonia browni.

Pentstemou confertus (form not typical).

Pbai'elia magelbmica.

Finns attenuata.

Pinus lambertiaua.

Finns ponderosa.

Polygala cornnta.

Populus tricbocarpa.

Potentilla glandulosa (large form).

Frunns subcordata.

Pseudotsuga mncronata.

Ptiloria lactuciua.

Fyrola picta.

Fyrola secunda.

Quercus californica.

Rbamnns californica (or rubra).

Rbus trilobata.

Ribes amictum.

Ribes cereum.

Ribes klamatbense.

Ribes viscosissimum.

Rosa californica.

Rosa gyninocar})a.

Rubus parvitlorus.

Rubus vitifolius.

Salix lasiandra.

Salix uuttalli.

Salix sitcbensis.

Sanrbucus melanocarpa.

Sisyrincbium bellum.

Smilax californica.

Solidago elongata.

Sorbns sambucifolia.

Spira'a douglasi.

Stacbys ingrata.

.Sympboricarpos pilosus.

Symi)boricarpos racemosus.

Tritclia ixioides.

Vaccinium arbuscula.

Vagnera stellata.

60

NORTH AMERICAN FAUNA.

[NO. 16.

PLANTS OF TRANSITION ZONK — continued.

(2) (S^)ec('es restricted to Tra)iiiHlun zone.

Abies concolor lowiana.

Acer circinatuni.

Acer glabnim."

Acer niacrophylluni.

Achillea lannlosa.

Aluns tennifolia.

Amelancbier alnifolia.*

Anapbalis niargavitacea.

Autenuaria geyeri.

Apocynuni pumilum.

Aquilegia truncata.

Arctostapbylos jjatula.

Artemisia Indoviciaua (form).

Asarnni Iiartwcgi.

Ceanotbus cordulatus.

Ceanotbns integerrimus.

Ceanotbns (Cerastes) prostratus.

Ceanotbns velutinus.

Cerasns deuiissa.

Cerastis emarginata.

Cerasns glandnlosa.

Cercocarpus ledifolins.

Cbrysampbcra calilornica.

Cbrysotbamnns bloomeri augustatns.

Cornus nnttalli.

Coruns pubescens.*

Corylns rostrata ealifornica.

Cratii'gns rivularis.*

Cryptantbe geminata.

Fragaria bracteata.

Fragaria cbiloensis.

Fritillaria atropnrpnrea.

Gayopbytnni ramosissimnm.

Gilia fCollomia) granditiora.

HaHtingsia alba.

Hieracium albitlornm Harge form).

Hieraciuni cynoglossoides nudicaule.

Hieracium greenei.

Horkelia psendocapitata.

Lapjiula nervosa.

Liboccdrus decurrens.

Lilium wasbiugtonianum.

Linna-a borealis.* '

Linum lewisi.

Lotus aniericanus.

Lnpiuns minimus.

Osmorrhiza nuda.

Facbystima myrsinites.*

Pu'onia browui.

Pinus attenuata.

Finns lambertiana.

Finns ponderosa.

Folygala coruuta.

Fseudotsuga mucronata.

Quercus californicii.

Rhamnns californiia (or rubra).

Ribes klamatbense.

Ribes viscosissimum.

Rosa californica.

Ro.sa gymnocarpa.

Rnbus parviflurns.*

Rnbns vitifolins.

Salix lasiandra.

Sambncus melanocarpa.

Sisyriucbinm bellnui.

Smilax californica.

Solidago elongiita.

Sympboricarpos pilosus.

Sympboricarpos racemosus.

In otber mountains the species marked with an asterisk (*) are believed to occur
in the Canadian zone as well as the Transition.

(3) Species common to Transition and rppev Sonoran zones.

Alnus rboml>itolia.
Artemisia tridentata.
Betula occidentalis.
Cercocarpus parvifolius.
Cbrysotbamnns occidentalis.
Eriogouum nudum.

Fraxinns oregana.
Juniperns occidentalis.
Kunzia tridentata.
? Popnlns trichocarpa.
Prunus subcordata.
Rhus trilobata.

^Linnaa horeaUs is ordinarily a Boreal plant, but it was not found in the Boreal
belt of Shasta, though common in places at Sisson. Tlie Sisson form has been
separated as var. longijioru, and may be worthy of recoguition as a Transition zone
subspecies.

OCT., 1899.]

CANADIAN ZONE.

Gl

PLANTS OF TUANSiTiON /ONE — Continued.

(4) Sjtecieii common to Transition and Canadian zones.
(Or growinj; ou boundary between.)

Canim gairdneri.

Castanopsis sempeivirens.

Castilleja rainiata.

Chareiit'saracha nana.

Chimapliila uienziesi.

Cliimapbila umbellata.

Epilobimn brevistylum.

Epilobiniu oregouense.

Epilobium spicatum.

Eriogonum niarifolium.

Eupatorium occidentale.

Gilia aggregata.

Heleniastrnm rivulare.

Heracleum lanatum.

Lupin u.s elmeri.

Macbicrantbera shastensis (large form).

Mimulus moniliformis.

Mimulus tilingi,

Pentstemou confertns (form not typical).

Phacelia magellauica.

Poteutilla glandulosa (large form).

Ptiloria lactucina.

Pyrola picta.

Pyrola secuuda.

Kibes amictum.

Ribes cereum.

Salix nuttalli.

Salix 8itchen.si8.

Sorbus samlnicifolia.

SpiriPa douglasi.

Stacbys ingrata.

Tritelia ixioides.

Vaccinium arbnscula.

Yasnera stellata.

CANADIAN ZONE.

The Canadian zone on Sliasta forms a broad forest belt, usually sev-
eral miles in width and about 2,000 feet in vertical depth, which com
pletely encircles the mountain. On southwesterly slopes it begins at
an altitude of about 5,000 to 5,500 feet and pushes up to 7,500 to 7,800
feet.

Excepting" an area of lodge-pole pines {Pinus murrayana) on the
northeast base of the mountain, the Canadian zone is everywhere
marked by a continuous forest of stately trees, consisting of Shasta lirs
{Abies shasttmsis), mixed in places with silver pines {Pimts monticola).
It is a dark, somber forest, growing on a blackish soil, with very little
noticeable vegetation except scattered patches of dwarf nianzanita in
the dry woods, and lines of more succulent plants at widely distant
intervals along the borders of streams.

MAMMALS OP CANADIAN ZONK.

(1) Species known to occur in Canadian -one on Shasta.

Aplodontia major.

Callospermophilus obrysodeirus.

Canis lestes.

Cervus occidentalis.

Eretbizon epixantbus.

Eiltamias amoenus.

Eutamias senex.

Evotomys mazama.

Felis otegonensis.

Lepus klamatbcnsis.

Lutra hudsonica.

Lutreola visou energumenos.
Microtus mordax.
Mustela caurina.
Mnstela ijcunanti.
Myotis lucifugus longicrus.
Myotis yumanensis saturatua.
Neotoma cinerea.
Neurotricbus gibbsi major.
Odocoileus columbianus.
Perognatbua mollipilosus.
Peromyscus boyli.

62 NORTH AMERICAN FAUNA.

MAMAIALS OF CANADIAN ZONE COUtinUcd.

(1) S2)ecie>^ kuoirn to occur in Canadian zone on Shasta — Contiuued.

[no. 16. I

Peromyscus gambeli.
Putorius arizoneusis.
Scapauns califoinicus.
Sciuroptcrus alpinus ]<laujathensis?.
Sciurus albolimltatns.
Sorex luontereyeiisis.
Sores (Neosorex) navigator.
Sorex shasteusis.

(2) Species restricted to Canadian cone

Sorex vagraus anid'nus.
Taxidea taxus.
Tliomomys montieola.
Ursus americauus.
Ursns horribilis.
Vulpes macrourus.
Zapus triiiotatus alleni.

Aplodontia major.

? Sorex (Neosorex) navigator.

Sorex shastensis.
Zapus trinotatns alleni.

(3) S2)ecies common to Canadian and Transition zones.

[See p. 57.]

(4) tSjye.cies common to Canadian and Hndsonian zones.

Callospermophiliis clirysodeirus.
Cams lestes.
Eretbizon epixautbus.
Eutamias amd'uus.
Eutaruias seuex.
Evotomys mazama.
Lepus klamatbensis.
Microtus mordax.
Mustehi caurina.
Mustela pennauti.

Myotis yumaneusis satnratus.
Neotoma ciiierea.
Odocoileus cohimbianus.
Perognatbus luollipilosus.
Peromyscus gambeli.
Putorius arizonensis.
Sorex vagraus amcrous.
Tbomomys montieola.
Vulpes macrourus.

BIRDS OF C.\NADIAN ZONE.

(1) Species Iniown to occur in Canadian zone on Shasta.

Accipiter atricapillus striatulus.

Accipiter velox.

Aquila cbrysai'tos.

Bubo virginianus.

Buteo borealis calurus.

Carpodacus cassini.

Ceopblo'iis pileatus abieiicola.

Certhia familiaris occidental is.

Cbordeiles virginianus.

Cinclus mexicanns.

Coccotbraustes vespertinns montanus.

Colaptes cafer.

Contopus borealis.

Cyanocitta stelleri.

Dendragapus obscurus fuliginosus.

Deiulroica auduboni.

Deudroica occidentalis.

Dryobates villosus bylosco])us.

Empidonax difficilis.

Empidonax bammondi.

Empidonax wrigbti.

Falco sparvcrius.

Geothlypis tolmiei.

Heiminthopbila celata lutesceus.

Helmintbopbila rubricapilla gutturalis.

Hylocichla aonalaschka^ auduboni.

Junco hyemalis tburberi.

Loxia curvirostra bendirei.

Melospiza lincolni.

Mernla migratoria propinqua.

Myadestes tovrnsend:.

Oreoi'tyx pictus plnmiierus.

Parns gambeli.

Perisoreus obscurus,

Picoides arcticus.

Piranga ludoviciana.

Regulus calendula.

Regulus satrapa olivaceus.

Selaspboriis rnfiis.

Sitta canadensis.

Sitta carolineusis aculeata.

Spbyrapicus thyroideus.

Spinas pinus.

Stellula calliope.

Wilsouia pusilla ]iileolata.

Xenopicus albolarvatus.

i OCT.. 1899.]

CANADIAN ZONE.

63

BIRDS OF c.\NAi>iAX /.( )Ni':— Continued.
(2) Species restricted to Canadian zone.

Loxia cnrvirostra bendirci.
Melospiza lincolni.

Picoidea iircticus.

(3) Species common to Canadian and Iranhiition zonen.

[See p. 58.]

(4) Species common to Canadian and Hudsonian zones.

Accipiter atricapilliis striatulus.

Accipiter velox.

Aquila chrysai'tos.

Carpodacus cnssini.

Cinclus mexicanus.

Coccothrauste.s vesi)ertinns montanns.

Dendragapus obscurns fnliginosus.

Dendroica auduboni.

Dendroica occidentalis.

Falco sparverius.

J unco liyemalis thvirlieri.

? Loxia cnrvirostra bendirei.
Myadestes towuseudi.
Parus gambeli.
Perisoreus obscurus.
Regulus calendula.
Regulus satrapa olivaceus.
Selasphorus rnfus.
Sitta canadensis.
Sphyrapicu.s thyroideus.
Spin us pinus.
Stellula calliope.

PLANTS OF CANADIAN ZONE.

(1) Species known to occur in Canadian zone on Sltasta.

Abies shastensis.

Aconitum columbiamun.

Allium validum.

Alnus sinuata.

Arctostaphylos nevadensis.

Arnica longifolia.

Campanula wilkinsiana.

Carnm gairdneri.

Castanopsis sempervirens.

Castilleja miniata.

Chamaesaracba nana.

Cbiniapblla menziesi.

Chimapbila umbellata.

Chrysotbaninus bloonieri,

Corallorbiza bigelovi.

fCrepis intermedia.

Cymopteris terebintbinus; Marge form).

Delphinium sonnei.

Drosera rotundifolia.

Epilobium brevistylum.

Epilobium oregonense.

Epilobium spicutum.

Erigeron inornatus.

Eriogonum niarifohum.

Eupatorium occidentale.

Gentiana simplex.

Gilia aggregata.

Habenaria leucostacbys.

Habenaiia unalascbeosis.

Heleniastruui rivulare.

Pleracleum lanatum.

Holodiscus discolor (large form).

Hypericum anagalloides.

Ligusticum grayi.

Lilium parvum.

Lupinus elmeri.

Madia bolanderi.

Macb;eranthera shastensis (large form).

Mimulus moniliformis.

Minuilus primuloides.

Mimulus tilingi.

Monardella odoratissinia.

Peutsfcenion confertus (lorm not typical).

Pentstemou deustus.

Pentstemon gracilentus.

Phacelia magellanica.

Phlox douglasi diffusa.

Pinus monticola.

Pinus murrayana.

Potentilla glandulosa (large form).

Ptilor.a lactucina.

Pyrola pallida.

Pyrola picta.

Pyrola secunda.

Ribes amictum.

Ribes cereum.

Salix uuttalli.

Salix sitchensis.

Senecio trigonophyllus.

Sorbus sambucifolia.

64

NORTH AMERICAN FAUNA.

[NO. 16.

PLANTS OF CANADIAN ZONE — Continued.
(1) Species known to occur in Canadian zone on Sliasta — Continued.

Spirppa douglasi.
Spraj^uea uuibellata.
Stachys ingrata.
Stellaria crispa.
Tofieldia occidentalis.

Tritelia isioides.
Vaccinitim occidentale.
Yagnera stellata.
Veratrum californicuni.
Viola blanda.

(2) Species restricted to Canadian zone.

Abies shastensis.
Acouitum columbiauum.
Allium validum.
Alnus siuuata.
Arnica longifolia.
fCorallorbiza bigelovi.
Delphinium sonuei.
Drosera rotundifolia.
Erigeron iuornatus.
Gentiana simples.
Habeuaria leucostachys.
Habeuaria unalascbensis.

Lilium parvum.
Madia bolauderi.
Peutstemon deustus.
Peutstemoii gracilentus.
Pinua monticola.
Pinus niurrayana.
Pyrola pallida.
Seuecio trigonophyllus.
TofielJia occidentalis.
Yaccinium occidentale.
Yiola blanda.

(3) Species common to Canadian and Transition zones.
[Seep. 61.]

(4) Species common to Canadian and Hudsonian zones.

Arctostaphylos nevadensis.
Campanula wilkiusiana.
Castanopsis sempervirens.
Castilleja miniata.
Cbrysothamnns bloomeii.
Holodiscus discolor.
Hypericum anagalloides.

Ligusticum grayi.
Mimulus primuloides.
Monardella odoratissima.
Phlox douglasi diffusa.
Ribes cereum.
Spraguea uuibellata.
Stellaria crispa.

HU13SONIAN ZONE.

The Hudsonian zone is the highest of the timber belts. Its sinuous
upper border rises ou the high ridges to Inclose the narrow tongues of
dwarf prostrate trees that push uj) on the warmest southwesterly expo-
sures to an extreme altitude of 9,800 feet, but between the ridges it
dips down a thousand feet or more, and is difticult to fix with precision.
The lower border slightly overla^js the upper limit of Shasta firs. Two
species of trees, and only two, grow in this zone — the black alpine hem-
lock {Tsuga mertensiana) and the white-bark pine {rinus alhicaulis).
The hemlock is restricted to local spots, while the white-bark pine forms
a practically continuous belt, as already explained (see p. 42). Ou
warm southwesterly slopes the Hudsonian reaches from 7,500 or 7,800
up to 9,500, or in extreme cases to 9,800 feet.

OCT., 1899.]

HUDSONIAN ZONE.

65

MAMMALS OK HUD.SONiAN ZONE.

(1) Species known to occur in Uudsonian zone of Shustn.

Callospenuopbilus chrysodeirus.

Cauis lestes.

Erethizon epixanthns.

Eutamias amtenus.

Eutamias eenex.

Evotomys raazama.

Lepiis klamatbensis.

Microtus mordax.

Mustela canriiia.

Mustela pennanti.

Myotis ynmanensis saturatns.

Neotoma cinerea.

Ochotona schisticeps.
Odocoileus colnmbianus.
Ovis canadensis.
Perognathus mollipilosus.
Peromyscua ganibeli.
Pheuacomys orophihis.
Putorius arizonensis.
? Sorex vagrans amceuus.'
? Sorex (Xeosorex) navigator.
Thomomys mouticola.
Vnlpes macronrus.
? Zapns trinotatus alleni.'

(2) Species restricted to Hiidsonion zone.

Ochotona schisticeps.

(3) Species common to Hiidsonian and Canadian zones.

[Seep. 62.]

(4) Species common to Hndsonian and Alpine zones.

Canis le.stes.-
Microtus mordax.
Ovis canadensis.
Perorayscns gambeli.

Phenacomys orophilus.
Thomomys monticola.
Vulpes macrourus. -

BIRDS i^V HUD.SONIAN ZONE.

(1) Species knonn to occur in Hndsonian zone on Shasta.

(Species queried are not positively known to breed in the Hudsonian, tliough frequently .seen in this

belt in Jnly .and August.)

Accipiter atricapilhis striatuhis.

Accipiter velox.

Aquila chrysii-tos.

Carpodacns cassini.

fCerthia familiaris occidentalis.

Cinclns mexicanus.

?Coccothraii3tes vespertinus inontanus.

Dendragapus obscurus fuligiiiosus.

fDendroica andnboni.

Dcndroica occidentalis.

Falco sparverins.

Jiinco hyemalis tburberi.

?Loxia curvirostra bendirei.

Myadestes townsendi.

Nucifraga colnmbiaua.
Parus gambeli.
Perisoreus obscurus.
Eegulus calendula.
Regulns satrapa olivaceus.
Salpinctes obsoletus.
Selasphorus rufus.
Sialia arctica.
Sitta canadensis.
?Sitta carolinensis aculeata.
Sphyrapifus thyroidens.
Spinus pinus.
Stellula calliope.
Zouotrichia leucophrys.

(2) Species restricted to Hudsonian zone.
Nucifraga Columbiana. Zonotrichia leucophrys.

' These three species occur along the lower edge of the Hudsonian zone, but were
not obtained in its upper part, and it is not certain whether or not they should be
included.

-Does not breed above Hudsonian.

21753— :N^o. 16 9

66

NORTH AMERICAN FAUNA.

[NO. 16.

Biups OK HUDsoNiAN /ONE — Continued.

(3) Species common to Hudnonian and Canadian zones.

[See p. 63.]

(4) Species common to Hndsonian and Alpine zones.

Salpiuctes obsoletus.' Sialia arctica.

Stellula calliojie.

PLANTS OF HUDSONIAN ZONK.

(1) Species known to occur in Hudsonian zone on Shasta.

Agoberis mouticola.

Allinni sp. ?

Antennaria media.
Arabis platyspeiiiia.
Arctostaphylos nevadensis.
Arnica merriami.
Arnica viscosa.
Campanula wilkinsiana.
Carex breweri.
Castanopsis semijervirens.
Castilleja affinis.
Castilleja miuiata (small form).
Cbeirantlins perennis.
Cbrysotbamnus bloomeri.

Crepis intermedin.
Cycladeiiia Immilis.

Cyniopterus terebintbinns.
Epilobiuni tdavatum.

Epilobinni obcordatum.

Epilobiuni pringleanum.

Erigerou armeri:i'folium.

Eriogonuni polypodum.

Eriogonum pyrolafolinm.

Hieracium albitlorum (alpine form).

Hieracium gracile.

Hieracium borridum.

Holodiscus discolor.

Hypericum anagalloides.

Juncus parryi.

Juniperus niina.

Kalmia glauca microphylla.

Ligusticum grayi.

Lupinus albil'rons.

Lupinus 'ornatns.'

Lntkea pectiuata.
Macb;erantbera sbastensis.
Miuiulus implexus.
Mimnlus ])rimuloides.
jVIitella pentandra.
Mouardella odoratissima.
Oreastrum alpiginum.
Oreobroma triphylla.
Ortbocarpus pilosns.
Parnassia californica.
Pentstemon glaber vitahensis.
Pentstemon menziesi.
Pentstemon newberryi.
Pblox douglasi diffusa.
Pbyllodoce empetriformis.
Piuus albicaulis.
Polygonum newberryi.
Polygonum sbastense.
Potentilla flabelli folia.
Potentilla ])8euilorupestris.
Pulsatilla occidentalis.
Kibes cereum.
Saxifraga bryopbora.
Scutellaria nana.
Sibbaldia procumb«ns.
Silene grayi.
Sitanion cinereum.
Spraguea umbellata.
Stellaria crispa.
Streptanthns orbiculatus.
Tsuga mertensiana.
Vaccluium ca^spitosum.
Veronica cusicki.
Viola purpurea.

(2) Species restricted to Hndsonian zone.

Allium sp. ?

Arnica merriami.
Arnica viscosa.
?Castilleja affinis.

Castilleja miniata (small form).
Cycladeuia humilis.
Epilobiuni clavatum.
Epilobium obcordatum.

1 On Shasta the rock wren is restricted closely to the Alpine and Hudsonian zones,
but in other places it occurs much lower down, and was recently found in Sbasta
Valley by AV. K. Fisber.

OCT., 189'.).]

ALPINE ZONE.

67

PLANTS OF UUDSONIAX ZOXK — COUtiuTied.

(2) Sjtecies restricted to Hndsouian zone — Continued.

Pentstemon "laber utiiheiisis.

Ei)ilo1jinai priugleanum.

Holodisciis discolor (typical form)

Jnniperns nana.

K;dinia glauca niicrophylla.

Miniiilus implexus.

Mittdla peutandra

Oreobronia triphylla.

Ortliocarpiis pilosiis.

Paniassia califoniica.

Pentstemon newberryi.
Phyllodoce enijietrifbriius.
Pinus albicaulis.
Poteutilla flabellifolia.
Potentilla psendornpestris ?
Scutt llaria nana.
Tsnga niertensiana.
Vaccininm cjTjspitosuin.

(3) Species common to Hudsonian and Canadian zones.
[See p. lU.]

(4) Species common to Hudsonian and Alpin<- zones.

Agosciis nionticola.
Aiiteunaria media.
Arabis platyspermu.
Carex breweri.
Clieirautbus perenni.s.
Clirysothamnns bloomcri.
Cymopterus terebinthinus.
Erigeron armeria' folium.
Eriogounm polypodum.
Eriogonura pyrola-folium.
Kieracium albitloruiii (al])inc form).
Hieracium gracile.
Hieracium horridnm.
Juncns parry i.
Ligusticnm grayi.
Lu;Mnu8 'oruatus.'

ALPINE ZONK

Lutkea pectinata.
Macbarauthera sbasteusis.
Oreastrum alpiginum.
Pentstemon menziesi.
Plilox douglasi difii'usa.
Polygonum newberryi.
Polygonum sbasteuse.
Pulsatilla occidentalis.
Sibbaldia procumbens.
Silene grayi.
Sitaniou cinereum.
S]iraguea nmliellata.
St rep taut bus orbiculatus.
Veronica cusicki.
^'ioIa ])urpnrea.

Tbe Alpine zone occui)ies the irregular belt of pumice and lava
between timberline and the upper limit of plant growth. On the
warmer southwesterly slopes its lower limit may be found at 9,500 to
9,800 feet, but on ordinary slopes it is considerably lower. The great
majority of its species stop at or below an altitude of 11,000 feet, but
on the relatively warm southwesterly slopes Huhea nana was found at
11,1)00 feet, and two species. Draba brnceri and Pnlemonlion pnlchelliimy
as high as 13,000 feet — the extreme limit of plant growth on iShasta.

MAiMMALS OF ALPINF. ZONK.

(1) Species knouin to occur in Alpine zone on Shasta.

Canis lestes.'
Microtus mordax.
Ovis canadensis.
Peromysciis gambeli.

Pbenacomys oropbilus.
Tbomorays nionticola.
Vulp<'.s macrourus.'

(2) Species restricted to Alpine zone.
None.

' The coyote and fox range up into the Alpine zone, but do not breed above the
Hudsonian and possibly not above the Canadian.

68

NORTH AMERICAN FAUNA.

MAMMALS Of ALPINE ZONE — contimied.

(3) Species common to Alpine and Hudsonian zonts.
[See p. 65. J

BIRDS OF ALPINE ZONE.

(1) Species known to occur in Alpine zone on Shasta.
Anthus pensilvanicus. Salpinctes obsoletus.

Leucosticte tepbrocotis.
Nucifraga coluiubiana.

Authiis pensilvanicus.

Sialia arctica.
Stellula calliope.

(2) Sjyecies restricted to Alpine zone.

Leucosticte tepbrocotis.

(3) Species common to A Ipine and Hudsonian zones.
[See p. 66.]

PLANTS OF AI.PINE ZONE.

(1) S2)ecies known to occur in Alpine zone on Shasta.
Lupinus 'ornatiis.'

Acbillea borealis.

Agoseris luonticola.

Anteunaria media.

Arabis platysperma.

Bikukulla imitlora.

Cardainiiie liellidifolia pacbypbylla

Carex breweri.

Cbitnactis iievadeusis.

Cbeirantbus pcreunis.

Cbrysotbarunus bloomeri.

CymopteriLs terebintbinus.

Draba breweri.

Erigerou armeri;clbliniu.

Erigeron compositns trifidns.

EriogouuDi polypoduiii.

Eriogomim pyrobi- folium.

Hieraciiini albiliorum (alpine form),

Hieracium gracilc

Hieracium borridum.

Hulst-a larseni.

Hulsea nana.

Jnncus parryi.

Ligustioum grayi.

Lupinus lyalli.

Lutkea pectiuata.
Macba^rantbera sbastensis.
Oreastrum alpiginum.
Oxyria digyua.
Pentstemon menziesi.
Phacelia frigida.
Pblox douglasi diti'usa.
Polemonium iiulcbellum.
Polygonum uewberryi.
Polygonum sbastcuse.
Pulsatilla occidentalis.
Sagina saginoides.
Saxifraga tolmiei.
Senecio canus.
Sibbaldia procumbens.
Silene grayi.
Silene suksdorti.
Sitanion cinereuni.
Spraguea nmbellata.
Streptantbus orbiculatus.
Veronica cusicki.
Viola purpurea.

(2) Species restricted to Alpine zone.

Acbillea l)orealis.

Bikukulla unitli)ra.

Cardamiue liellidifolia pacbypbylla.

C'ba'nactis uevadensis.

Draba breweri.

Erigeron compositus tritidu«.

Hnlsea larseni.

Hulsea nana.

Lupinus lyalli.
Oxyria digyna.
Pbacelia frigida.
Polemonium pnlcbellum.
Sagina saginoides.
Saxifraga tolmiei.
Senecio canus.
Silene suksdorti.

(3) Species common to Alpine and Hudsonian zones.
[See p. 67. ]

THE BOREAL FAUNA AND FLORA OF SHASTA CONTRASTED
"WITH CORRESPONDING FAUNAS AND FLORAS OF THE SIERRA
AND THE CASCADES.

In considering tbe relations of the boreal faunas and floras of Shasta
to those of other parts of the Sierra-Cascade system it is necessary at
the outset to have a clear conception not only of the extent of the
range as a whole, but also of the number and magnitude of the breaks
or gaps in the continuity of its boreal fauna and flora. The Cascade
Range enters the State of Washington from British Columbia in lati-
tude 49^^ and pushes ..outhward completely across "Washington and
Oregon; its continuation, the Sierra Nevada, traverses California
for a distance of 500 miles, ending a little south of Mount Whitney, in
about latitude 36°. The Cascade-Sierra system, therefore, extends over
13 degrees of latitude, and has a total length of fully 1,000 miles. For the
whole of this distance it rises abruptly from a low region, whose faunas
and floras are in the southern part Sonoran, in the northern part Tran-
sition. The field work of the Biological Survey has shown that the
narrow boreal band which occupies the higher parts of the range is not
continuous, but is interrupted by four important gaps, through which
Transition zone species pass freely in broad belts from one side to the
other. These gaps, begining at the north, are:

(1) The Gohonbia Gap,ov gorge of the Columbia River, on the bound-
ary between Washington and Oregon, where the breadth of the Tran-
sition zone seems to be less than 50 miles.^

(2) The Klamath Gap, on the boundary between Oregon and Califor-
nia, extending from a little south of Mount Pitt in Oregon to Mount
Shasta in California, a distance of about 50 miles. This interval is
iuterrui)ted by one or two detached groups of low mountains on the
California side, and by long ridges on both sides, whose summits are
inhabited by boreal species, materially decreasing the actual breadth
of the gap.

(3) The Pitt Biver Gap, between Mounts Shasta and Lassen in north-
ern California. The breadth of the Transition zone here is about 00
miles.

'Although not bearing on the fauna of Shasta, it is interesting to note, in connec-
tion tt-ith the effects of the Columbia River Gap, tliat a number of species charac-
teristic of the northern Cascades, in the State of Washington, do not occur in the
southern Cascades, iu Oregon. Among the mammals the most notable species of
this kind are Arctomys caVujains, CallospcnnophUiis xatitratus, Krotomys gajiperl .latii-
rati(8, Oreamnos montanns, Feromysciis oreas, Pntorins ivashingtoni, Zapus trinoiatus.

69

70 NORTH AMERICAN FAUNA. [no. 16.

(4) The Feather River or Quincy Gap, between Mount Lassen and the
high ridg^e northwest of Honey Lake — the northern end of the Sierra
proper. This gap is the shallowest, narrowest, most irregular, and
least effective of all, and is the only one which has not been cut deeply
and com])letely through the range by a large river. The Boreal zones
of the two sides, at the points where they come nearest together, which
is southeast of the southeastern extension of the boreal plateau on
which Lasseu stands, are not separated, apparently, by more than 15
miles. The distance between the Hndsoiiian elements ap[)ears to be
several times greater. This region needs further exi)loration.

Fully half of the boreal species of Shasta are common to both the
Sierra Nevada and the Cascade IJange, and some of them extend over
the entire length of the Sierra-Cascade system, inhabiting the princi-
pal boreal summits all the way from British Columbia to ]Mount Whit-
ney: others are restricted to particular parts of the mountains, and
each of the lour gaps mentioned forms a barrier beyond which certain
species do not pass. Therefore, in contrasting the boreal faunas and
floras of Shasta with corresponding faunas and floras of the Sierra
Nevada and Cascade Range it is necessary to fix definite limits to the
terms employed. The term 'Sierra,' as used in the table headings and
following discussion, is restricted to the lofty range extending from
Mount Whitney northward a little beyond Honey Lake; the 'Cas-
cades,' to the Cascades of Oregon. In other words, the term ' Sierra' is
restricted so as not to include Shasta or Lassen;' the term 'Cascades'
so as not to include the Cascatlcs of Washington.

The paucity of animal and jjlant life on Shasta, contrasted with that
of the Sierra and Cascades, has been already noted and is clearly
shown in the following tables. The explanation, briefly stated, is that
Shasta, on account of its aridity and relatively small area, is incapable
of supporting so rich a fauna and flora as either of the extensive ranges
between which it is situated. It is not assumed that all of the boreal
species inhabiting Shasta were discovered by us, but in the case of the
Canadian and Hudsonian mammals and birds, and the Hudsonian and
Alpine plants it is believed that the number which escaped detection
is too small to materially alter the results here given. ^

In grouping the species for study it seems most logical to arrange
the mammals, birds, and plants in two principal categories: {a) Boreal

' The flora and fauna of Lassen are not known in sufficient detail to admit of com-
plete comparisons in either direction; hence this mountain is omitted from consid-
eration in tbe accompanying tables. At the same time it should lie stated that
Lassen is clearly a part of the Sierra, so far as its fauna is concerned.

'-'The accompanying percentages and lists of species are provisional and snbject
to revision. They are based on present information and will, of course, be corrected
and supplemented by future field work. They are sufficiently near the truth, how-
ever, to demonstrate certain facts and warrant certain deductions and generaliza-
tions of very great interest in connection with the origin of the l)oreal faunas and
floras of the Sierra Nevada and Cascade Range.

OCT., 1890] BOREAL SPECIES OF SHASTA AND SIERRA-CASCADES. 71

species that occur on Shasta, with reference to their occurrence in the
Sierra or Cascades or both; and (/>) Boreal species that occur on the
Sierra or the Cascades or both, but which, so far as known, are absent
from Shasta.

BOREAL SPECIES OF SHASTA CONSIDERED WITH REFERENCE
TO THEIR PRESENCE OR ABSENCE IN THE SIERRA AND THE
CASCADES.

The boreal mammals, birds, and plants of Shasta have been grouped
in four categories • (1) species common to Shasta and the Sierra-Cas-
cade system as a whole: (2) species common to Shasta and the Sierra,
but not known from the Cascades; (3) species common to Shasta and
the Cascades but not known from the Sierra, and (4) Shasta species not
known from either the Sierra or the Cascades.

Thirty-six distinctively Boreal mammals are known from Shasta,
including the boreal species which range down into or through the
Transition zone. Of these, 20 are common to the Sierra on the south
and the Cascades on the north, 7 are common to Shasta and the Sierra
but are not known from the Cascades, 1 is common to Shasta and the
Cascades but is not known from the Sierra, and 2 are peculiar to
Shasta.

Of the 3G distinctively Boreal mammals of Shasta, 17 are believed to
be exclusively boreal. Of these, 12 are common to the Sierra and the
Cascades, 4 are common to Shasta and the Sierra but are not known
from the Cascades, 1 is peculiar to Shasta, but not one is common to
Shasta and the Cascades which does not occur also in the Sierra.

Forty-seven distinctively Boreal birds are known from Shasta, includ-
ing tlie boreal species which range down into or through the Transition
zone. Of these, 41 aie common to the Sierra and the Cascades, 4 are
common to Shasta and the Sierra but are not known from the Cas-
cades, and 2 are common to Shasta and the Cascades but are not known
from the Sierra.

Of the 47 distinctively Boreal birds of Shasta, 22 are believed to be
exclusively boreal. Of these, 18 are common to the Sierra and the Cas-
cades, 2 are common to Shasta and the Sierra but are not known from
the Cascades, find 2 are common to Shasta and the Cascades but are
not known from the Sierra.

One hundred and twelve distinctively Boreal plants are known from
Shasta, including the boreal species which range down into or through
the Transition zone. Of these 55 are common to the Sierra and the Cas-
cades; 31 are common to Shasta and the Sierra but are not known
from the Cascades; 10 are common to Shasta and the Cascades but
are nol. known from the Sierra, and 8 occur on Shasta which are not
known from either the Sierra or the Cascades.

Of the 112 distinctively Boreal plants of Shasta, 101 are believed to
be exclusively boreal. Of these, 47 are common to the Sierra and the
Cascades; 28 are common to Shasta and the Sierra but are not known

72 NORTH AMERICAN FAUNA.

[no. 16.

from the Cascades; 15 are common to Shasta and the Cascades but
are not known from the Sierra; and 8 are restricted to Shasta.

Three mammals, 5 birds, and OS plants are believed to be restricted
to the Hadsonian and Alpine zones. Of these, 2 mammals, 3 birds,
and 30 plants are common to the Sierra and the Cascades; one mam-
mal, 1 bird, and 18 plants are common to Shasta and the Sierra but
are not known from the Cascades; and no mammal, 1 bird, and 12
plants are common to Shasta and the Cascades, but are not known
from the Sierra. Five Hudsonian-Alpine plants from Shasta are not
known from either the Sierra or the Cascades.

These comparisons show:

(1) That of the boreal species known from Shasta, including those
which range down into the Transition zone, 87 percent of the birds,
72 percent of the mammals, and only 50 percent of the i)lants are
common to the Sierra and the Cascades.

(2) That of the exclusively boreal species known from Shasta, 85 per-
cent of the birds, 70 percent of the mammals, and 4G jiercent of the
lilants are common to the Sierra and the Cascades,

(3) That of the exclusively Iludsonian and Alpine species known
from Shasta, GO percent of the birds, 07 percent of the mammals, and
44 percent of the plants are common to the Sierra and the Cascades.

(4) That in each instance, as w^ould be expected, the percentage of
species common to the two ranges is greater in the case of those rang-
ing down into the Transition zone than in those restricted to the Boreal,
for the obvious reason that geograiihically the Boreal belt is broken by
broad gaps, while the Transition zone is practically continuous.

(5) That of the birds, mammals, and plants of Shasta, birds have by
far the largest jtercentage of species common to the Sierra and the
Cascades, mammals next, and plants least of all. This corresponds
with the relative powers of dispersion possessed by these groups.

Arranged primarily by groups instead of zone limits, it appears that
the percentages of Shasta birds common to the Sierra and the Cascades
are as follows: Of boreal species, including those which range down
into the Transition zone, 87 percent; of species restricted to the Boreal
zones, 85 x^ercent; of species restricted to the Hudsonian and Alpine
zones, GO percent.

The percentages of Shasta mammals common to the Sierra and the
Cascades are: Of boreal species, including those which range down into
the Transition zone, 72 percent; of species restricted to the Boreal
zones, 70 percent; of species restricted to the Hudsonian and Alpine
zones, G7 percent.'

The percentages of Shasta plants common to the Sierra and the Cas-
cades are: Of boreal species, including those which range down into
the Transition zone, 49 percent; of species restricted to the Boreal

' The nmnber of Hudsonian-Alpine species is too small to give this percentage
much value.

OCT., 1899.] BOREAL SPECIES OP SHASTA AND SIERRA-CASCADES.

73

zones, 46 percent; of species restricted to the Hudsouian and Alpine
zones, 44 percent.

For evidence of another kind — that based on the absence from Shasta
of species which occur in tlie Sierra or the Cascades or both — see pages
79-82.

Following are the tables on which the foregoing generalizations are
based :

(1) BOREAL SPECIES COMMON TO SHASTA AND THE SIERRA-CASCADE

SYSTEM.

(Species followed by the letter T range down into or through the Tran.sition zone.)

(a) MAMM.*LS.

Callospermophilns clirysodeirus T.

Canis lestes.

Erethizou epixanthiis T. ?

Eutamias auKt'iius T.

Eiitaniias seuex T.

Evotomys mazama.'

Felis oregonensis T.

LepiiM klaiiKitheiisis T.?

Lutreola visou energuuienos T.

Microtus mordax T.

Mnstela canriua.

Mustela pennanti.

Myotis lucifugiiB lougicrus T.

Myotis yumanensis saturatiis.
Neotoma cinerea T.
Nenrotrichus gibbsi major T.
Ochotona schisticeps.
Odoeoileus colunibiaiiiis T.
Peromysciis ganibcli T.
Phenacomys oropbilus.
Piitorius arizouensis.
Sorex; vagrans amo'mis T.
Sorex (Neosorex) navigator.
Taxidea taxns T.
Ursiis aniericanns T.
Vnlpes macrourus T.

(b) BIRDS

Accipitcr atriciipillns striatnlns.
Accipiter velox.
Aquila chrysai'tos T.
Carpodacus cassini.
Ceophlcvns pileatus abieticohi T.
Certbia familiaris occidentalis T.
Cincliis mexicaniis T. ?
Coccotbraustcs A'espertinns niontauns.
Colaptes cafer T.
Contopus borealis T.
Dendragapus obsciirus fnliglnosns T.
Dendroica aitduboui T.
Dendroica occidentalis.
Drj'obates villosus bj'loscopus T.
Euii)idonax difficilis T.
Eiiipidoiiax liaminoudi T.
Eitipidonax wrigliti T.
Geotblypis tolmiei T.
Helmintbopbibi celata Intescens T.
Helmiiitbopbila rubricapilla gnttiir-
alis T.

Hylocicbla aonalascbka' audtiboiii T.

Mclospiza lincolni T.

Mernla uiigratoria iiropiiKjua T.

Myadestes townsendi.

Nucifraga columbiaua.

Oreortyx pictns plnniiferns T.

Parus gambeli.

Picoides arcticns.

Piranga ludoviciana T.

Regnlus calendula.

Regains satrapa olivaceus.

Salpinctes obsoletns T.

Selaspborus rufus T.

Sidlia arctica.

Sitta canadensis.

Sitta carolinensis aculeata T.

Spbyrapicus tbyroideus.

Spinas pinus.

Stellula calliope T.

AVilsonia pusiila pileolata T. ?

Zouotricbia lencopbrys.

' ICvotomys mazama, a common Cascade species, was obtained by us on Shasta and
Lassen, and doubtless occurs in the Sierra, farther south.

21753— No. 16 10

74

NORTH AMERICAN FAUNA.

[no. 10. i

(o) PLANT.<

Achillea boroalis.

Aconitmn «'<>lumbianuni.

Allium validuni.

Alnu.s sinuata.

Aiitenuaria media.

Aral lis platyspcrma.

Arctostaphylos ne^ adensis.

Bikiikulla imillora.

C'ardamiue bellidifolia pacbyjiLylla.

Cares breweri.

Castanopsis sempervirens T.

Chimapbila meuziesi T.

Cbimaphila nmbellata T.

Chrysothamnus Idoomeri.

Cymopterus terebinthinus.

Drosera rotundifolia.

Epilobinm obcordatiim.

Epilobiiim pringleaiium.

Eriogoiium pyrolypfoliivm.

Gentiana simplex.

Habeuaria leucostacbys.

Habeuaria iinalaschensis.

Heracleum lanatum T.

Holodiseus discolor.

Hypeiicnm auagalloides.

JuiK'us parryi.

Jnnipenis nana.

Kalmia glaiica miciopbylla.

Mimiiliis ]>riiniiloides.
Mitclla ])entandra.
Oreoliroma tripliylla.
Orthocaipus ])ilosus.
Oxyria digyna.
Peutstemon newbeiryl.
Phlox douglasi diffusa.
Pimis albicanlis.
Piuiis moDticola.
Finns miirrayaua.
I'olemoiiiiim pulchellum.
Polygomim shastense.
Potentilla llabellifolia.
Pulsatilla occidentalis.
Pyrola picta T.
Pyrola secunda T.
Ribes cereum T.
Sagina sagiuoidcs.
Salix nuttalli T.
>Salix sitchensis T.
Saxifraga tolmiei.
Sil)baldia procuinbens.
Sorbus sambucifolia T.
Spraguea umbellata.
Tsnga merteiisiana.
Viola blanda.
Viola purpurea.

(2) BOREAL SPECIES COMMON TO SHASTA AND THE
NOT KNOWN FROM THE CASCADES.

SIERRA BUT

(a) MAMMALS.

Aplodoiitia major (represented in Cascades by A. major rainieri).
Ovis canadensis (not known or represented in Cascades).
Peromyscus boyli T. (not known or represented in Cascades).
Sciurus albolimbatus T. (represented in Cascades by S. cascadensis).
Sorex niontereyensis T. (not known i'rom any point north of Shasta).
Thomomys monticola (represented in Cascades by T. mazama).
Za^jus trinotatus aiieni (represented in Cascades by Z. montanus).

(h) HIKDS.

Cyanocitta stelleri frontalis T. (represented in Cascades by C. stelleri).'
.Tunco hyemalis thurberi (rejireseuted in Cascades by J. h. connectens).
Leucosticte tephrocotis- (represented in Cascades by L. t. littoralis).
Loxia curvirostra bendirei (represented in Cascades by L. c. minor).

'The Shasta jay is intermediate between stelleri and frontalis.

-The mention of Leucosticte tephrocotis in this and subsequent lists involves an
assumption. L. tephrocotis is known to be the breeding rosy finch of the High Sierra;
L. t. littoralis that of Mount Rainier in the Cascades of Washington. A rosy
finch was found, but not secured, on Shasta. It is reforred provisionally to iephro-
co/i.'< rather than littoralis, because the great majority of Shasta species which are
not common to both ranges are Sierra and not Cascade forms.

OCT.. 1809.]

BOR15AL SPECIES RESTRICTED TO SHASTA.

75

BOREAL SPECIES OF SHASTA AND SIERRA— Continiu-d.

(f) I'LANTS.

Agoseris monticola.

Arnica meniaiiii.

Castilleja affinis.

Castilleja miniata (alpine form).

Chif uactis nevadensis.

Cheirauthus pereniiis.

Corallorliiza bigdovi.

Crepis intermedia.

Cycladeuiii hnmilis.

Delpliiuinm sonnei.

Dr.iba breweri.

Erlgeron armeri;efolium.

Erigeroii compositns trifidiis.

Erigeron iiiornatus.

Erlogonum polypodnni.

Hieracium albitlonim (alpiue form).

Hieracium horridum.

Hulsea larseni.

Madia bolaaderi.

Parnassia californica.

PentstemoD gracileutus.

Ribes aniictiim T.

Saxilraga bryophora.

Senecio cauiis.

Senecio trigonophyllns.

Stellana crisjia.

Streptanthus orbiculatus.

Vaccininm ra'spitosuni.

Vaguera wtellata T.

Veratrum californi<uui T.

(3) HOREAL SPECIES COMMON TO SHASTA AND THE CASCADES P. IT
NOT KNOWN FROM THE SIERRA.

(a) MAMMALS.

Cer^■Ils occidentalis T.

Aiitlms ])eu8ilvanicu3.

Abies shastensis.
Epilobiiim clavatum.
Hieracii'ui gracile.
Hulsea nana.
Ligusticum grayi.
Ln])inu8 lyalli.
Lnpinns 'ornatus.'
Lutkea pectiuata.
MacbaTauthera ebasteusis.

(b) BIKDS.

Perisorens obscurus.

(C) PLANTS.

< (reastrum alpiginum.
Pentstemon mcnziesi.
Pliyllodoce empetriformis.
Polygonum uewberryi.
Silene suksdorfi.
Tofieldia occidentalis.
Vaccininm arbnscula T.
Veronica cnsicki.

(4) BOREAL

SHASTA SPECIES NOT KNOWN FROM EITHER THE SIERRA
OR THE CASCADES.

Perognathns mollipilosur T.

Arnica longif<dia.
Arnica viscosa. '
Campanula wilkinsiana.
Mimulus implexns.

(a) MAMMALS.

Sorex shastensis.

(b) HIRDS.

[None]

(C) I'LAXTS.

Pentstemon glaber ntahensis.
Pbacelia frigida. '
Scutellaria nana.
Silene grayi. '

So far as known restricted to Shasta.

76

NORTH AMERICAN FAUNA.

[NO. 16.

(5) EXCLUSIVELY BOREAL SPECIES OF SHASTA.

(a) MAMMALS.

Aplodontia major.

Cauis lestes.

Evotomys luazama.

f Lepns klamatbensis.

Microtus mordax.

Mustela cauriua.

Mustela peuuanti.

Myotis yiimaueusis saturatus.

Ochotoua schisticeps.

Ovis canadensis.
Pheuacomys oiophilns.
Putorius arizonensis.
Sorex (Xeosorex) navigator.
Sorex shasten.sis.
Thonioiuys mouticola.
Vnlpes macroiirus.
Zapua tiinotatus alleui.

(6) BIRDS.

Accipiter atricapillu.s striatulus.

Anthus ])ensilvanicii8.

Carpodacus cassini.

Cinclus niexicanus.

Coccothranstes vespertinus luontanus.

Deudroica occideutalis.

Juuco byenialis thurbeii.

Lencosticte tejibrocotis.

Loxia curvirostra bendirei.

Myadestes towuseudi.

Nucifraara Columbiana.

Parus gambeli.
Perisoreus obscurus.
Picoides arcticus.
Reguliis calendula.
Kegulus satrapa olivaceus.
Sialia arctica.
Sitta canadensis.
Spbyrapicus tbyroideus.
Spinus pinus.
Wilsouia pusilla pileolata.
Zonotricbia leueoiibrys.

(c) PLANTS.

Abies sbastensis.
Acbillea borealis.
Aconituni columbianum.
Agoseris mouticola.
Allium validum.

Allium sp. ?

Aluus sinuata.

Antennaria media.

Arabis platysperma.

Arctostapbylos uevadensis.

Arnica longifolia.

Arnica merriami.

Arnica viscosa.

Bikukulla uniHora,

Campanula wilkinsiana.

Cardamine bellidifolia pacbyphylla.

Carex breweri.

Castilleja miniata (alpine form).

Chii'uactis nevadensis.

Cbeirantbus pereunis.

Chrysotbamnus bloomeri.

Corallorbiza bigelovi.

Crepis intermedia.

Cycladeuia bumilis.

Cymopterus tercbintbinus.

Delpbinium sonnei.

Draba breweri.

Drosera rotuudifolia.

Epilobium clavatum.

Epilobium obcordatum.

Epilobium pringleanum.

Erigeron armeria'folium.

Erigeron compositus tritidns.

Erigeron inornatus.

Eriogonum polypodum.

Eriogonum ])yrolie folium.

Gentiana simplex.

Habenaria leucostacliys.

Habenaria unalascbcnsis.

Hieracium albitlorum (aljiine form).

Hieracium graeile.

Hieracium horridum.

Holodiscus discolor.

Hulsoa larseni.

Hulsea nana.

Hypericum anagalloides.

Juncus parryi.

Juniperus nana.

Kalmia glauca micropbylla,

Ligusticnm grayi.

Lupinus albifrons.

Lupinns lyalli.

Lupinus 'oruatus.'

Lutl<ea pectinata.

Macbii'vantbera sbastensis.

Madia bolanderi.

Mimuliis implexus.

Miiiiulns primuloides.

OCT., 1809.] EXCLUSIVELY HUDSONIAN- ALPINE SPECIES OF SHASTA. 77

EXOLUSIVELY BOEEAL species of SHASTA— Coiitiuiied.

(c) PLANTS — continued.

Mitella pentandra.
Mouardella odoratissima.
Oreastrum alpiginuni.
Oreobroma triphylla.
Orthocaipus pilosus.
Oxyria digyna.
Parnassia caliloruita.
Pentstemon denstns.
Peiitsteuion glaber utahensis.
Pentstemon gracilentns.
Pentstemon menziesi.
Pentstemon newberryi.
Pliacelia frigida.
Phlox donglasi diffusa.
Phyllodoce enipetriformis.
Pinus albicaulis.
Pinus monticola.
Piniis murrayana.
Polemonium pulclielhini.
Polygonum newberryi.
Polygonum sbastense.
Potentilla flabellifolia.

Potentilla pseudorupestris.
Pulsatilla occidentalis.
Sagina saginoides.
Saxifraga bryopbora.
Saxifraga tolmit-i.
Senecio cauus.
Senecio trigonopbyllus.
Sibbaldia procumbens.
Silene grayi.
Silene suksdoiti.
Sitauion ciuereum.
Spraguea umbcllata.
Stellaria crispa.
Streptantlius orbiculatus.
Tofieldia occidentalis.
Tsuga niertensiana.
Vaccinium ca'spitosuui.
Vaccinium occidentale.
Veronica cusickl.
Viola blanda.
Viola purpurea.

(6) EXCLUSIVELY HUDSONIAN-ALPINE SPECIES OF SHASTA.

MAMMALS.

Pbenacouiys orophilus.

Ochotona scbisticeps
Ovis canadensis.

Anthus pensilvanicus.
Leucosticte tcpbrocotis.
Nucifraga Columbiana.

Sialia arctica.
Zonotrichia leucophrys.

Achillea borealis.

Agoseris monticola.

Antennaria media.

Arabi.s platysperma.

Arnica merriami.

Arnica viscosa.

Bikukulla unillora.

Cardamine bellidifolia pachyphylla.

Carex breweri.

Cha'uactis uevadensis.

Cheiranthus perennis.

Chrysothamnus bloomeri.

Cycladenia bumilis.

Cymopterus terebinthinus.

Draba breweri.

Epilobium clavatum.

Epilobium obcordatum.

Epilobium pringleanum.

Erigerou armcria^folium.

Erigeron compositus tritidus.

Eriogonum polypodum.

Eriogonum pyrohe folium.

Hieracium albillorum (alpine form).

Hieracium gracile.

Hieracium horridum.

Holodiscus discolor (alpine form).

Hulsea larseni.

Hulsea nana.

Juncusparryi.

Juniperus nana.

Kalmia glauca microphylla.

Lupinus albifrons.

Lupinus lyalli.

Lupinus ' ornatus.'

Lutkea pectinata.

Macha-ranthera shastensis.

Miniulus implexus.

Mitella pentandra.

Oreastrum alpiginum.

Oreobroma triphylla.

78

NORTH AMERICAN FAUNA.

I NO. 16.

EXCLUSITELY HUD.sOX I AX-ALPINE SPECIES OF SHASTA— Continued.

PLANTS — continued.

Orthocarpus iiilosus.
Oxyria digyna.
Parnassia califoruica.
Pentstemou glaber utaheusis.
Pentstemon menziesi.
Pentstemon uewberrvi.
Phacelia frigida.
Phyllodoce empetrifonuis.
Pinus albicaulis.
Polemoniuni pulcbellum.
Polygonum newberryi.
Polygonum sbastense.
Potentilla llabellifolia.
Potent ilia pseudorupestris.

Pulsatilla occideutalib.
Sagiua saginoides.
Saxifraga lnyopbora.
Saxifraga tolmiei.
Scnecio canus.
Sibbaldia procuinhens.
Silene grayi.
Silene suksdorti.
Sitauion cinereum.
Streptantbus orbiculatTig
Tsuga mertensiana.
Aaccinium ciespitosum.
A'eronica cusicki.
Yiola purpurea.

(7) EXCLUSIVELY HUDSOXIAN-ALPINE SPECIES COMMON TO SHASTA
AND THE SIERRA-CASCADE SYSTEM.

Ochotona scbisticeps.

Nucilraga columbiana.
Sialia arctica.

mam:\ials.

Phenacomys orophilus.

BIRD.S.

Zonotricbia leucopbrys.

PLAX1>

Acbillea borealis.

Autennaria media.

Arabis platysperma.

Bikukulla uuillora.

Cardamine bellidifolia paehyjthylla.

Carex lireweri.

Chrysotbamnus bloomeri.

Cymopterus terebintbinu«.

Epilobium obcordatum.

Epilobium pringleanum.

Eriogonum pyrolieiolium.

Holodiscusdiscolor(ty;.icala!piu<' form)

Juncus parry i.

Juniperns nana.

Kalmia glauia micropbylla

Mitrilla jjentandra,
(Jreobroma triplijlla.
Orthocarpu.s pilosus.
Oxyria digyna.
Pentstemon newberryi.
Pinus albicaulis.
Polemonium pulcbellum.
Polygonum sbastense.
Potentilla llabellifolia.
Pulsatilla occideutalis.
Sagina saginoides.
Saxifraga tolmiei.
Sibl)aldia procumbens.
Tsuga mertensiana.
Viola pinpurea.

(8) EXCLUSIVELY HUbSONIAN-ALPINE SPECIES COMMON TO SHASTA
AND THE SIERRA BUT NOT KNOWN FROxM THE CASCADES.

MAMMALS.

( )vi8 canadensis.

BIRDS.

Lencosticte tephrocotis.

PLANTS.

Cycladenia bumilis.
Draba breweri.
Erigeron armeriiBfolium,
Erigeron compositus trifidus.

Agoseris monticola.
Arnica merriami.
Chaenactis nevadensis.
Cbeirantbus perennis.

OCT., 1899.] SIERRA-CASCADE SPECIES NOT KNOWN FROM SHASTA. 79

HUDSONIAX-ALPINE SPECIES OF SHASTA AND SIERRA— Continued.
PLANTS — continued.

Eriogonum polypodnra. Saxifraga bryophora.

Hieracium albiflorntn (alpine form). Seiiecio canus.

Hieracinni horridum. Streptantlms orbiculatns.

Hulsea larseni. Vacciniuni cie8i)itosuni.
Parnassia californica.

(9) EXCLUSIVELY HUDSONIAX-ALPINE SPECIES COMMON TO SHASTA
AND THE CASCADES BUT NOT KNOWN FROM THE SIERRA.

MAM.MALS.

Xone.

lURDS.

Anthns jiensilvanicus.

Epilobinm clavatura. Oreastruni alpiginnm.

Hieracium gracile. Pentstemon nienziesi.

Hulsea nana. Phyllodoce empetriformis.

Lupinus lyalli. Polygonum newberryi.

Lupinus 'ornatus.' Silene suksdorti.

Lutkea pectinata. Veronica cnsicki.
Mach^ranthera shastensis.

BOREAL SPECIES OF THE SIERRA-CASCADES NOT KNOWN FROM

SHASTA.

Turning to another phase of the subject, the absentees, or boreal
species of the Sierra and Cascades which are not known from Shasta,
an equally instructive lesson may be learned.

The boreal species that occur in the Sierra or Cascades, or both, but
which are not known from Shasta, have been grouped in three cate-
gories: (1) species common to the Sierra-Cascades but not known from
Shasta; (2) Sierra sjiecies not known from Shasta or the Cascades;
and (3) Cascade species not known from Shasta or the Sierra.'

(1) Boreal species common to the Sierra and the Cascades

BUT NOT KNOWN FROM ShASTA.

Only three boreal mammals are known to occur in both the Sierra
and the Cascades which have not been found on Suasta. These are
the Sierra marmot {Arctomys Jlaviventer), the wolverine {Gulo luscus),
and the silver-haired bat [Laslonyeteris noctivagans). The marmot,
it maybe stated with confidence, is really absent; the wolverine has
been killed in the near vicinity and probably occurs on Shasta; the
bat is a local species common in the mountains west of Shasta, easily
overlooked and most likely to occur. Hence there is every reason to

' In the accompanying tables and discussion the boreal species are treated col-
lectively, no account being taken of the important distinctions between the Alpine,
Hudsonian, and Canadian species. This course has been rendered necessary by the
absence of discriminative zone lists of Cascade-Sierra species.

80 NORTH AMERICAN FAUNA. [no.ig.

believe that the Sierra marmot is really the only mammal common to
the Sierra and the Cascades which does not occur on Shasta.

Two boreal birds believ-ed to be common to the Sierra-Cascade
system (the western winter wren, Anorthura hiemalis imcifica^ and the
Townsend warbler, Dendrolca townsendi) have not yet been discovered
on Shasta, but are liable to be found there at any time.

With plants the case is (j[uite different, for at least 19 well-known
genera, and a considerable number of si^ecies of other genera, not
known from Shasta are common to the Sierra and the Cascades.

(a) MAMMALS.

Arctomys llaviveuter. Gulo lusciis. Lasionycteris iioctivagans.

(b) BIRDS.

Anorthura bieinalis pacitica. Dendroica towiiseudi.

(c) PLANTS.

Genera nof Jiuowii from Shasta.

Arenaria. Iris. Ranunculus.

Cassiope. Ivesia. Smelowskia.

Clayton ia. Listera. Streptopus.

Clintouia. Mertensia. Thalictruni.

Erytbroniuiu. Pedicularis. Xerophylliini.

Geuni. Primula.

Hencbera. Raillardella.

Ad(1ifio)iaI specieb not Lnoirn from Shasta.

Arnica chamissonis. Lonicera involucrata.

Campanula scouleri. Polygonum bistortoides.

Crepis nana. Populus tremuloides.

Erigerou salsugiuosiis. Salix barclayi.

Gentiaua uewberryi. Saxifraga nivalis.

.Tuncus ortliophyllns. Saxifraga punctata.
T.onicera conjugalis.

(2) Boreal Sierra species not known from Shasta or the

Cascades.

Eight mammals, 1 bird, 3 coniferous trees, several shrubs, and a num-
ber of small plants are known from the High Sierra which do not occur
on Shasta or the Cascades; the majority of them are restricted to tlie
southern part of the Sierra, not reaching as far north as the mountains
about Lake Tahoe, and consequently need not be considered here.
Only three of the Sierra mammals {Spermopliilns beldingi, Euiamias
speciosus frater^ and Sorex obsciirns) which range north to the south
end of Feather River Ga]) fail to reach Shasta, and two if not all three
of these are known to cross this gap and occur on Lassen, showing
that the Feather River Gap of itself is of very little significance. The
details of plant distribution in these mountains are not sufficiently
known to admit of safe generalizations.

OCT., 1899.]

BOREAL CASCADE SPECIES.

81

BOREAL SIERRA SPECIES— Continued.

(a) MAMMALS.

Eutamias alpiuus.^
Eutamias speciosus.'
Eutaniia.s speciosus callipeiilns.'
Eutamias speciosus frater.

Pinicola euucleator californica.

Abies magnifica.
Arenaria compacta.
Arenaria congesta.
Artemisia rothiocki.
Chrysopsis breweri.
Draba lemmoni.
Eulophus parisbi.
Hulaea algida.
? Leiluiii glaudulosum.
Leucotboe davisi;t.
Lycbnis californica.
Montia fontana.

Microtus dutcberi.'
Sorex obscurus.
Spermopbilus beldingi.
Tbomomys alj)inu8. '

{b) BIRDS.

(C) I'LANTS.

Oreobroma uevadensis.
Oreobroma pygnuea.
Pbyllodoce breweri.
Pinus balfouriana.
Piuus flexilis.
Primula suffrutescena.
Queicus vaccinifolia.
Raillardella scaposa.
Ranunculus oxynotus.
Silene bernardina.
Silene californica.

(3) Boreal Cascade species not known from Shasta or the

Sierra.

Eleven mammals, 2 birds, 3 Hr trees, and a number of shrubs aud
other plants which inhabit the Cascade Range in Oregon are not known
to occur on Shasta or in the Sierra Nevada. One of the mammals
[JEutamias townsendi) does not reach as far south as the southern end
of the Cascades, and two others {Aplodontia major rainieri and Sciurus
c«S(?f«^/rjw/.s) are only subspecifically separable from corresponding forms
in the Sierra. The remaining eight are independent specific types not
represented on Shasta or in the Sierra, and all of them i)ush south to
the extreme southern end of the Cascades immediately across Klamath
Gap from Shasta.

(a) MAMMALS.

Aplodontia major rainieri.

Eutamias townsendi.

Lyux canadensis.

Microtus (Arvicola) arvicoloides.

Microtus (Chilotus) bairdi.

Putorius cicognani streatori.

Hylocicbla ustulata.

Scapanus alpinns.
Sciurus douglasi cascadensis.
Sorex (Atophyrax) bendirei.
Tbomomys mazama.
Zapus montanus.

(b) BIRDS.

Leucosticte tephrocotis littoralis.^

' Tbese mammals are restricted to the southern part of the Sierra and none of them
come as far north as the mountains about Lake Tahoe,

-Leucosticte tephrocotis liltoralis breeds in the Cascades of Washington but is not
actually known from the Cascades of Oregon. It is likely to be found among the
glaciers of Mount Hood and The Sisters when the birds of these mountains are
studied.

21753— No. IG 11

82

NORTH AMERICAN FAUNA.

[NO. 16.

(c) PLANTS.

Abies amabilis.
Abies lasiocarpa.
Abies nobilis.
Gaultheria myrsinites.
Menzicsia ferrngiuea.
Rbododemlrou albitJonim.
Ribes erythrocarpum.

Ribes laciistre.
Rubus lasiococcuB.
Sileiie acaulis.
Sorbus occideutalis.
Spiriua arbnscula.
Vacciuiura micropbyllum.
Valeriana sittheusis.

EFFICIENCY OF KLAMATH GAP AS A BARRIER TO BOREAL
SPECIES COMPARED ■WITH THAT OF PITT RIVER AND FEATHER
RIVER GAPS COLLECTIVELY.

In view of the iianowueiss of Klamatli Gap, a break of less than 50
miles, separating- the boreal fanna of Shasta from that of the Cascades,
compared with the breadth of the coinbiiied Pitt Kiver and Feather
River gaps, abont 100 miles, separating Shasta from the boreal fauna
of the Sierra Nevada northwest of Honey Lake, one might expect Shasta
to share more species with the Cascades than with the Sierra. The con-
trary is true. The Feather Eiver Gap, as elsewhere explained (p. 70), is
ineffective compared with the others; the branches of Feather River do
not cut completely through the mountains, and the gap is merely a low
part of the range, with the Honey Lake ridge and small boreal-capped
peaks i)rqjecting here and there as stepping stones between the main
Sierra and Mount Lassen. Pitt River Gap is deeper, cutting completely
through the range between Lassen and Shasta, forming a boreal break
about 00 miles in width, and there is no apparent reason why it should
not be as effective a barrier as Klamath Gap, although from the stand-
point of zone distribution it does not cut so low and therefore has a
slightly cooler summer climate, in consequence of which it is less effect-
ive. But this difference is insufficient to explain the really great dis-
parity in potency of the two, for in checking the extension of boreal
si)ecies Klamath Gap has proved far more effective.

Passing over the species common to Shasta and the Sierra-Cascade
system as a whole (see p. 73), only three of the ten distinctively Sierra
mammals which reach the northern end of the Sierra fail to reach
Shasta, and two if not all of these bridge the Feather River Gap and
reach Mount Lassen, which is separated from Shasta by only the Pitt
River Gap. On the other hand, not one of the ten distinctively Cascade
mammals which occur at the extreme south end of the Cascade Range
has been able to cross the narrow Klamath Gap to Shasta.

If the number of distinctive mammals of the Sierra-Cascade system
be reduced by subtracting those which are represented in the two
ranges by closely related forms ' 8 distinctive specific types will remain

'Close discrimination of species and subspecies is necessary in laying off the
minor subdivisions of faunas; and it is interesting from the zoological standpoint
to know which and how many of tlie specific types common to a given area have
undergone enough change in parts of that area to warrant separate recognition by
name, but from the standpoint of the distribution of specitic types such details are
of little value.

83

84 NORTH AMERICAN FAUNA. [no. 16.

for the southern Cascades aud 7 for the northern Sierra. Of these
distinctive specific types only 3 of tlie 7 Sierra species fail to reach
Shasta, while all of the 8 Cascade species fail.

The significance of these facts appears when the Boreal faunas of
the mountains north and south of Klamath Gap are studied with ref-
erence to their geographic sources of origin, as pointed out iu the next
chapter.

SOURCES OF THE BOREAL FAUNAS OP SHASTA AND OF THE
SIERRA AND THE CASCADES.

The boreal auiinals aud plauts of the Sierra-Cascade system as a
whole are not yet well enough known to admit of positive statements
as to the number of species or the details of their distribution. Hence
a complete study of their distribution with reference to the geographic
source of origin of the various specific types is Jiot possible. Never-
theless, enough has been learned to point to some very interesting
conclusions.

It has been already shown that the boreal fauna and fiora of Shasta
form a part of the fauna and tlora of the Sierra-Cascade system; that
70 percent of the exclusively boreal mammals of Shasta are common
to both ranges; and that of the remainder, 80 percent are common to
the Sierra. It is obvious therefore that, so far as mammals are con-
cerned, Shasta may be considered a part of the Sierra (see p. 71).

Fifty-eight boreal ' species of mammals are known from the Sierra
and the Cascades. Of these, 31 (54 percent) are common to both ranges,
11 (19 percent) are restricted to the Sierra, and 16 (28 percent) to the
Cascades.

Of the 58 boreal species of mammals known to inhabit the Sierra-
Cascade system not a single genus or subgenus is peculiar, though the
genera Aplo(lont'«( and Neuroirlchns and the subgenus Atophi/ra.r are
restricted to the northwest coast region. With species the case is very
different, for 23 of the 58 species (40 percent) are peculiar to the
Sierra-Cascades; but even of these only 10 differ sufticiently from near
relatives elsewhere to be considered distinct specific types. Of the 58
species whose relationships are so obvious that there can be no doubt
as to their afiinities and origin, 5 (9 percent) come from mountains
farther north (in British Columbia, some ranging into southeastern
Alaska), 8 (14 jierceut) are of general transcontinental boreal distri-
bution, 16 (28 percent) are characteristic of the humid west or north-
west coast region, 19 (33 percent) are identical with or closely related
to species living in the Eocky Mountains, and 10 (17 percent) are dis-
tinctive superspecific types restricted to the Sierra-Cascade system.

If, instead of treating the Sierra-Cascade species collectively, we
group them with reference to the particular part of the mountains they
inhabit, putting the Cascade species (those north of Klamath (lap) in

'Our collections from the Cascades have not yet been worked up, aud it is proba-
ble that several species will be added to this number.

85

86 NORTH AMERICAN FAUNA. [xo.ie.

one category, and the Siena species (those south of Klamath Gap) in
another, some additional facts are brought out which emphasize the
widely different sources of origin of the distinctively Cascade species
on the one hand, and the distinctively Sierra species on the other.
Of the distinctively Cascade S})ecies, 25 ])ercent are derived from
mountains farther north, 12 percent are local types, 12 percent belong
to transcontinental boreal types, and 25 percent to northwest-coast
types. Of the distinctively Sierra species, 50 percent are specially
developed local types, and 50 percent belong to types common to the
Sierra and the southern Rocky Mountains.

These facts point not only to the great antiquity and effectiveness of
the Klamath Gap, but also to a former east and west continuity of
range of Boreal species between the Rocky Mountains of Utah and
Colorado and the Sierra Nevada of California, a distance of at least
500 miles.

MAMMALS OF SHASTA.

Sorex shastensis sp. iiov. Shasta Shrew.

Type from Wagon Camp, Mount Shasta (alt. 5,700 ft. in the lower part of the Cana-
dian zone). No. 95450, U. 8. Nat. Mna., Biological Survey Coll. Collected Sept.
26, 1898, by W. H. Osgood. Grig. No. 317.

Characters. — Size small; decidedly smaller than 8. vagrans ; tail
rather short; ears small, but couspicuous. Third uiiicuspid smaller
than fourth. Skull and teeth peculiar.

Color. — Type specimen, in change from summer to winter pelage:
Head and sides of neck to slioulders dull fulvous brown ; rest of upper
parts dark steel gray; underparts ashy brown; tail sharply bicolor,
dusky above, bufiy below, becoming dusky toward tip.

Cranial characters. — Skull small, decidedly smaller than in vagrans
and as small as in calif ornicus ; brain case moderately high — not at all
flattened as in californicns ; rostrum rather small (about as in californi-
eus); constriction swollen. Tooth row, as a whole, somewhat shorter
than in californicns ; unicuspids decidedly narrower, particularly the
first and second; molariform series much as in californicus, but slightly
smaller; large premolar very broad posteriorly.

Measurements. — Type: Total length, 90; tail vertebrse, 35; hind
foot, 12.

Remarlis. — This new species is based on a single specimen caught by
W. H. Osgood in a trap set in a springy place among the Shasta iirs,
immediately above Wagon Camp. In the same trap, and in the iden-
tical spot, he caught also specimens of Neosorex navigator and Neuro-
trichus gibbsi major. Several specimens of Sorex vagraiis amoenus were
caught near by, but no others of this species.

Sorex shastensis is a small shrew of uncertain aflflnities. In several
respects it resembles S. californicus, but differs from this species mark-
edly in color, and still more in the form of the cranium and narrow-
ness of the uuicuspidate teeth.

.Sorex vagrans amoenus Merriam. Sierra Shrew.

Twenty-two specimens of this small shrew were collected on Shasta
and about its base. Two were caught among the tules at Big Spring,
in Shasta Valley, on the north side of the mountain; two at Warm-
castle Soda Springs, in Squaw Creek Valley, on the south side; and
nineteen in the Canadian zone and lower part of the Iludsonian from
Wagon Camp up to upper Squaw Creek, Mud Creek, and Ash Creek.
Most of them were trapped under logs in damp places.

87

88 NORTH AMERICAN FAUNA. tNO.lC

Sorex montereyensis Merriam. Monterey Shrew.

Six specimens of tliis large long-tail shrew were secured — one at the
lower edge of the Hudsouiau zone, near ^lud Creek; four in the Can-
adian /one, in Mud Creek Canyon; and one in the Transition zone, in
Squaw Creek Valley, near Warmcastle Soda Springs.

Sorex (Neosorex) navigator Baird. White-bellied Water-shrew.

Apparently rather scarce, as a large amount of trapping resulted in
the capture of only four specimens. These were obtained at as many
localities, namely, the head of Panther Creek, upper Squaw Creek,
upjjer Ash Creek Canyon, and Wagon Camp, all in the Canadian zone.

Neurotrichus gibbsi major subsp. nov. Large Shrewraole.

Type from Carberry Ranch, Shasta County, Calif, (alt. 4,100 ft., between Mts, Shasta
and Lassen). No. 65321, c? ad., U. S. Nat. Mas., Biological Survey Coll. Collected
May 18, 1894, by C. P. Streator. Orig. No. 3789.

Characters. — Similar to X <jihh,si, but decidedly larger; hind feet
larger; forefeet much broader and longer; tail much longer; under
parts darker; skull larger and broader; fifth upper lateral tooth (the
'large premolar') decidedly .smaller than in X. {/ibhsi, and possessing a
distinct anterior cusp on the cingulum, which is absent in gihhsi; fifth
lower lateral tooth (the 'large premolar') decidedly larger than in gibbsi.

Measurements. — Average of 3 specimens from type locality: Total
length, 120; tail vertebrie, 4(); hind foot, 17. Average of 4 specimens
from Mount Shasta: Total length, 113; tail vertebrte, 11; hind foot,
16.2.

Remarks. — Four specimens of this new form of Gibbs shrewmole
were collected by our party on Shasta — all in the Canadian zone.
Three were caught in Mud Creek canyon near the mouth of Clear
Creek by Walter K. Fisher and W. H. Osgood, and one was taken at
Wagon Camp by Osgood. In May, 1894, my assistant, Clark P. Streator,
obtained 3 specimens (the type and cotypes) at Carberry Ranch, on the
south side of Pitt liiver, between Mounts Shasta and Lassen. Carberry
Ranch is in the upper part of the Transition zone. The Canadian
zone specimens from Shasta are not quite so large, but agree in other
characters.

Scapanus califomicus (Ayres). California Mole.

Vernon Bailey found a dead mole of this species, October 3, in the
road between Wagon Camp and Sisson at an altitude of 4,500 feet.
He reported mole ridges as common in places a little below Wagon
Camp, and also in sandy soil in the Murray pine belt east of the moun-
tain; W. H. Osgood saw mole ridges in Shasta Valley.

Myotis evotis (H. Allen.) Big-eared Bat.

At Sisson, September 5, R. T. Fisher obtained a specimen from a
small boy, who caught it in a kitchen. The species doubtless occurs
in Shasta Valley also.

OCT., 1899.] MAMMALS. 89

Myotis lucifugus longicrus (True),

At Wagon Camp, July 17, Walter K. Fisher shot one of these bats.
Small bats were abundant here, but kept so much in the forest that
they were hard to shoot.

Myotis californicus (And. c^ Bach.), California Bat.

A single specimen of this species, collected on Mount Shasta by C. H.
Towusend, is recorded by Miller in North American Fauna, Ko. 13,
page 71, October, 1S97.

Myotis yumanensis saturatus ]Miller.

Common among- the alpine hemlocks at Squaw Creek Camp, where
they were seen every night, darting* in and out of the flickering light
of the camp fire. Here I shot one the evening of August o, and four
the evening of August 9. Late in July and early in August small bats,
probably the same species, were seen nearly every evening at the tem-
porary camps on or near upper Mud Creek. The species is interesting
as the only bat secured in the Hudsonian z(me.

Vespertilio fuscus Beauvois. Large Brown Bat.

Common at Wagon Camp, where Vernon Bailey shot one July 17,
and I shot three the evening of July 38. Many more could have been
killed if desired. In 1883 C. II. Townsend obtained it at Sheep Rock.
The species is one of the commonest in the foothills and valleys, and is
easily recognized on the wing by its large size and its character of flight.

[Arctomys flaviventer (And. & Bach.). Mountain Marmot.

It may be asserted with confidence that no marmots of any kind live
on Shasta. Our collectors were at work on the mountain from July 15
until October without seeing a single individual. Moreover, when
Vernon Bailey and I made our trip completely around the peak the
latter part of July we kept near timberline all the way and made a
special search for marmots, but were unable to find a trace of their
presence.]

Spermophilus douglasi (Richardson). Oregon Ground Squirrel.

Common at Sisson and in McCloud and Shasta valleys, whence it
ranges up through the manzanita chaparral of the basal slopes nearly
to Wagon Camp. At Sisson R. T. Fisher collected eight during the
first half of September, and says of them : "One of the few really plen-
tiful mammals at Sisson. Hardly an acre in the valley is free from
their burrows. Lender the barns and houses, in the fields, along the
hot slopes east and west of the town, and even in the woods, one con-
stantly sees them. At the time I write of, September 1-15, they seemed
to be feeding chiefl}^ on acorns and chinquapins — acorns in the valley,
chinquapins on the western slopes. In behavior they were wild and
sneaking; at all times diflQcult to approach. None appeared to have
hibernated."

21753— No. 16 12

90 NORTH AMERICAN FAUNA. [no. 16.

At Big Spring, in Shasta Valley, where they abound, W. H. Osgood
saw several climb up on a beam and enter an opening in a granary.

Callospermophilus chrysodeirus Merriani, Golden-mantled Ground
Squirrel.

One of the most abundant and conspicuous mammals of the moun-
tain, where they were seen daily from the manzanita belt up to timber-
line, and where 52 specimens were collected. At Sisson they are rare,
but 2,000 feet higher are fairly common, as they are also in Squaw Creek
Yalley at the south base of the mountain. In the fir forest they make
their homes under logs or about the roots of trees, but in the neighbor-
hood of timberline live in burrows under the rocks, often in slide rock,
associated with small colonies of conies. At low altitudes they are
usually unwary and maybe easily killed with the 'auxiliary' barrel,
but in the neighborhood of timberline they are so exceedingly shy it
is difficult to approach within gunshot. At our camp among the alpine
hemlocks on upper Squaw Creek they first kept at long range, but find-
ing us harmless gradually overcame their fear, and finally, toward
the end of the season, came to be one of the most persistent of camp
robbers, stealing bread and other eatables. At the same time they
never came freely and boldly as did their associates, the chipmunks,
but always stole in silently and if possible kej^t out of sight.

This species goes into winter quarters much later than its relatives in
the Rocky Mountain region. On Shasta it was seen daily near timber-
line until after the middle of September and a few were noticed on
warm days as late as September 24, but all those secured during the
latter part of the month were young of the year. Between Wagon
Camp and Sisson they were seen as late as September 26.

Eutamias amcenus (Allen). Klamath Chipmunk.

Abundant in the chaparral of the lower slopes and thence up through
the forest to timberline. Fifty-three specimens were obtained at vari-
ous points on the mountain. At Wagon Camp they were common
and were seen picking unripe serviceberries the latter part of July.
At the south base of the mountain one was killed as low down as
Warmcastle Soda Springs in Squaw Creek Yalley. In the forest they
live mostly about logs and stumps and are quite fearless, but along
the upper edge of timber, where they live among the bare rocks, they
are tnuch more wary.

Eutamias senex (Allen). Allen Chipmunk.

Abundant in the Shasta fir belt and ranging down to Sisson and
Warmcastle Soda Springs at the base of the mountain and up to the
upper limit of continuous timber, though perhaps not to extreme tim-
berline. Sixty-eight specimens were secured.

At Wagon Camp they were common and were usually associated with
their small cousin, U. ama-iius. They are more arboreal than the other

OCT., 1899.] MAMMALS. 91

chipmunks and we often saw them in the trees 40 or 50 feet above the
ground, moving- about in the branches or chasing one another around
the great trunks of the hemk)cks and firs. At our camp in the alpine
hemlocks on Squaw Creek they were the most abundant and most fear-
less of the diurnal mammals. Here they were constantly associated
with the less abundaut golden-mantled ground sqiiirrelii {Call ospermo-
philus chrysodeirus), compared with which they are bolder, more active,
more graceful, and more interesting.

In camp they made frequent visits to the mess box, which they
clearly regarded as public property, approaching it boldly and without
suspicion and showing no concern at our presence — in marked contrast
to the golden-mantled ground squirrels, which approached silently,
stealthily, and by a circuitous route, in constant fear of detection. If
disturbed while stuffing their cheek pouches with bits of bread, pan-
cake, or other eatables, each chipmunk usually seized a large piece in
its mouth and scampered off, returning as soon as we withdrew. In
fact, they made themselves perfectly at home in camp, and evidently
ranked us with other harmless inhabitants of the forest. They climbed
up the sides of our tent and over towels hung to dry on branches, as if
such things had always been a part of their environment. It should be
added, however, that the most familiar animals were always the young
of the year, which probably had no recollection of the time l)efore our
arrival.

Along the upper border of the timber, where the ground is more
open and is covered with gray rocks and pumice instead of the dark felt-
ing of hemlock and fir needles and cones, the chipmunks are far more
alert and wary.

After the middle of Se^jtember the adults were rarely seen, and after
the 20th the young came out only during the warmest part of the day.

At Sisson, II. T. Fisher found these chipmunks more abundant than
any other mammal. They were common in the woods, in the chapar-
ral, on the hillsides, and in the bottom of the valley. At the time of
his visit — from the end of August to the middle of September — they
were in the molt and very ragged.

Sciurns albolimbatus Allen. Sierra Pine Squirrel.

[ = 8. calif ornicus Allen, preoccupied.]

Common in the Canadian zone forest of Shasta firs, and in the Transi-
tion forest of mixed pine and Douglas spruce. Among the Shasta firs
they were seen on all sides of the mountain and came up as high as the
lower edge of the alpine hemlocks. Among the pines and Douglas firs
they were seen as low as Bear Butte, near Squaw Creek Valley, and
were common at Sisson and thence northward along the base of the
Scott Mountains. Like other pine squirrels they lay up stores of cones
for winter use. At low elevations they rival the large gray tree squirrels
in collecting the seeds of the huge cones of the sugar pines. At higher

92 NORTH AMERICAN FAUNA. [no.16.

elevaiioiis tbey seem to feed largely ou the uiucb smaller seeds of the
Shasta iirs, the cones of which they collect iu large uumbers. These
cones are gathered iu heaps at the bases of trees, where the squirrels
live, and are also stored iu decayed logs, where they are stuffed into
all available openings. As 1898, the year of our visit, was an ' oft" year'
for cones, we were forced, in order to obtain specimens, to take advan-
tage of the stores made by these squirrels the previous year. Iu
them we found innumerable cones, more or less perfect and with the
seeds still untouched, of both Abies shastensi.s and A. loiriana.

Sciurus fossor Peale. Oregon (Tray Squirrel: Large Tree Squirrel.

Fairly common in the pine forest covering* the southern and western
basal slopes of Shasta. At ditt'erent times during the summer these
large squirrels were seen in Squaw Creek Valley and between Sisson
and Edgewobd. On July 13 Vernon Bailey found them common near
Bear Creek, between Fall Eiver Valley and Shasta, where the sugar
pines begin. They were then cutting oft' the scales and eating the green
seeds of the half-grown cones of sugar pines.

Sciuropterus alpinus klamathensis Merriam. Klamath Flying Squirrel.

The only tlying s(juirrel seen by our party was observed by me in
August on a cedar stub near a small stream a couple ol miles below
Wagon Camp, but was not secured. There is therefore some uncertainty
as to the species. At Sisson 1 was informed that a boy had a pair alive
in a cage, but he left town with them before they could be examined.

Castor canadensis Kuhl. Beaver.

Probably not now living in the immediate vicinity of Shasta, although
in 1883, according to C. H. Townsend, "a number of them occui)ied
unmolested a dam, which they had constructed in a corner of a meadow
belonging to Mr. J. H. Sisson.'" They were formerly common in Shasta
Eiver, where Walter K. Fisher was recently told a few were seen in
the winter of 1898-99.

Aplodontia major Merriam. Aplodontia: Sewellel.

In making the circuit of Shasta the latter part of July, Vernon
Bailey an<l 1 discovered a colony of aplodontias in some rank vegeta-
tion covering a springy place in Ash Creek Canyon, in the upper part
of the Canadian zone. A little later W. K. and R. T. Fisher were sent
there and obtained two specimens. About the same time they and
W. H. Osgood caught eight in Mud Creek Canyon near the mouth of
Clear Creek, at an altitude of nearly 7,000 feet.

Aplodontias live in wet or damp places usually overgrown with rank
vegetation, and preferably in springy, sloping ground where some of
their innumerable burrows and sunken runways are kept wet by tlie
cold trickling water. As is well known, they cut various plants, com
monly rank or woody kinds, which they gather and carry iu bundles
to their burrows, or to places near by, where they spread them out to dry.

OCT., 1S99.] MAMMAT.S. 93

In Ash Creek Canyou Walter Fisher found their cuttings to consist
chietiy of ferns and willows— the latter carried from a loug- distance.
In Mud Creek Canyon the cuttings consisted chietiy, according- to
W. H. Osgood, of thiinble-berry bushes, mountain asli, and brake
ferns — tiie latter predominating, and in one place forming a pile as big
as a bushel basket. The animals commonly live in colonies, but Osgood
concluded that in Mud Creek Canyon only one individual, or at mosr,
ft pair, lived in one place, '-though several may be distributed among
the branches of a stream."

Mus museulus Linn. House blouse.

Abundant at Sisson, and running wild like the native s])ecies. R.
T. Fisher reported them as constantly getting into his traps, partic-
ularly in the weeds and sedges in wet places along the banks of Cold
Creek, where he caught a dozen or more.

Reithrodontomys klamathensis sp. nov.

Tiipf from Big Spring ('May ten"). Shasta Valley, Calif. No. !:iri444, ^ ad., U. S.
Nat. Mus., Biological Survey Coll. Collected Sept. 18, 1898, hy W. H. Osgood.
Orig. No. 281.

Characters. — Si/e medium; ears and hind feet large; tail long, only
slightly shorter than in longicauda; color giayish or brownish gray,
decidedly paler than lowjicauda.

Color. — Summer pelage: Upperparts pale grayish brown, washed
with bufty on sides; underparts white, tail bicolor, dusky above,
whitish below.

Cranial characters. — Skull rather large; braiucase and rostrum rel-
atively broad; audita! bulhe small. The skull as a whole agrees better
with that of niet/aJotis than with that of lougicaiida, particularly in the
length of ]>alate and breadth of braiucase; but the rostium is broader
and the audital bulhe are smaller than in either.

Measurements. — Type: Total length, 149; tail vertebra', 71; hind foot,
19. Average of 2 adults from type locality: Total length, 144; tail
vertebr;Te, 66; hind foot, 18.5.

Remarks. — Both in color and cranial characters ReitJirodontomi/s
lJ((mathensis resembles the pale grayish R. megalotis of the desert
region of the southern part of the Great Basin much more closely than
it does the dark brownish R. longicauda of California west of the
Sierra.

This new harvest monse is common in wet places in Shasta and Lit-
tle Shasta valleys, where four specimens were obtained by W. H.
Osgood and 11. T. Fisher. They were caught in little runways in wet
grass near tules. The species doubtless reached Shasta Valley by way
of the open Klamath country. During onr explorations in eastern
Oregon in 1896, numerous specimens of the same species were caught
by my assistants, E. A. Preble and Cleveland Allen, in the tule marshes
bordering the streams connecting Malheur and Harney lakes.

94 NORTH AMERICAN FAUNA.

[no. 16.

Peromyscus gambeli (Baird). Common White-footed Mouse.

Common on all, or nearly all, parts of the mountain from Sisson up
to and possibly a little above extreme timberline. One hundred speci-
mens were collected.

In choosing their homes these mice are easily suited, for they seem
equally contented among the dense vegetation in damp parts of the
bottoms of canyons and among the bare lava rocks and pumice soil ot
the driest timberline slopes. Some were caught also in the heather
meadows bordering the little streams in the Hudsonian zone. At
Sisson E. T. Fisher found them rather rare except in damp woods
along Cold Creek, where he caught a number under roots and stumps
close to the water.

Peromyscus boyli (Baird).

Exceedingly rare, or else of such peculiar habits that it escaped
observation. Only a single specimen was obtained on Shasta. It was
caught at the extreme upper limit of the Canadian zone (alt. 7,800 feet)
on Squaw Creek, August 9, by Walter K. Fisher. Others were secured
at Fall River Lake, in the Transition zone, southeast of the base of the
mountain.

Peromyscus truei (Shf.). Big-eared Mouse.

Collected in Little Shasta Valley by Walter K. Fisher, who found it
living among bushes of Ceanothus cuneatus.

Weotoma fuscipes Baird. Round-tail Wood Rat.

Not found on Shasta, but common in some of the low valleys at its
base. Their characteristic stick houses were seen in the juniper forest
at the southern end of Shasta Valley, in the chaparral near Gazelle,
and in several places in the Scott Mountains (Bailey). In Little Shasta
Valley one was collected September 19 (Osgood).

Neotoma cinerea Ord. Bushy-tail Wood Rat.

Rather scarce. Only four specimens were obtained — two in Mud
Creek Canyon near the mouth of Clear Creek, and two high up on
Squaw Creek (alt. 8,800 feet). Of those caught in Mud Creek Canyon,
one was trapped at the end of an old log, the other at the entrance to
an aplodontia burrow. Shasta abounds in the kinds of ledges and
cliff's usually inhabited by this species, but, except at rare intervals, no
traces of the animals were found-

Microtus californicus (Peale). California Vole.

Not obtained by us except in Shasta Valley, where six specimens
were secured in September by W. H. Osgood and R. T. Fisher. Their
runways were found in very wet places in the tules at Big Spring, in
Shasta Valley, and along Little Shasta Creek. Walter K. Fisher
secured specimens along Shasta River, northeast of Edgewood.

OCT, 1899.] MAMMALS. 95

Micro tus montanus (Peale).

Not obtained on Shasta, but collected at Fall River Lake in August
by W. K. Fisher. In August, 1883, seveu specimens were collected iu
the grassy meadows at Sissou Tavern by C. H. Townsend.

Microtus mordax (Merriam). Mountain Vole.

Common iu suitable moist places througliout the Canadian and Hud-
sonian zones, particularly iu the lieather meadows a little below timber-
line, where, thoug:h chief!}' nocturnal, they were sometimes seen in the
daytime. Their burrows abound in the heather beds, especially along
the overhanging banks of streams, and are so large tlmt some of them
were at first mistaken for those of Microtus ariHcoloides, a species which
does not occur on Shasta. They were not found in the dry forest.
Seventy- four specimens were collected, of which only six were obtained
at Wagon Camp, i^ear Sisson Tavern R. T. Fisher caught fifteen along
"the wet and bushy banks of a long ditch." They probably reach
Sisson not from Shasta, but from Mount Eddy, in the Scott ^Mountains,
near the foot of which Sisson Tavern is situated. Still, it is possible
that the two colonies are connected along some of the few cold streams
that traverse the Transition zone slopes of Shasta.

Evotomys mazama Merriam. Mountain Evotomys.

Fairly common in moist places in the Shasta fir forests of the Cana-
dian zone, where nineteen specimens were obtained. Ten were col-
lected in Mud Creek Canyon near the mouth of Clear Creek (altitude
6,700 feet); seveu along Squaw Creek from 6,700 up to 7,700 feet; one
at the head of Panther Creek at 7,700 feet, and one at Wagon Camp at
5,700 feet. They were usually caught in traps set under logs iu damp
or wet places.

Phenacomys orophilus Merriam. Lemming-Mouse.

Rare and local, judging from the results of our trapping. Only three
specimens were obtained — all in the heather meadows along the upper
part of Squaw Creek, where they were caught August 7, 10, and 12 by
Walter K. Fisher.

Thomomys monticola Allen. Sierra Pocket Gopher.

Abundant throughout the boreal slopes of the mountain, from the lower
part of the Canadian zone to above timberliue. Iu the dark forests of
Shasta firs their mounds were seen wherever there was enough small
vegetation to furnish food, and were commonest along the streams and
about the edges of marshy j)laces, where plant life is abundant and luxu-
riant. Above the line of continuous timber their mounds were noticed
on many of the i^umice sloj^es between the altitudes of 8,000 and 0,000
feet. Above the head of Squaw Creek a small colony was found at an
altitude of 8,300 feet, and another between 8,900 and 9,000 feet. On
the east side of Mud Creek Canyon they were seen at 8,800 feet, and on
the west side at 8,900 feet. On our trip around the mouutain near

96

NORTH .UrERICAN FAUNA,

[no. 16.

tiinberline, the latter part of July, their mounds were found in almost
every ])la(e where the >soil was deep euough for the animals to work;

Fic. 31. — Wiiitir earth pltiys of pocket gopher.

Fig. :!2. — Momul iiiiule by pocket gopher. (Photogiaplitd b.v "W. H. Osgood,)

and on the north side they were abundant nearly up to timberliue, both
on the main peak and on Shastina.

OCT., 1899.] MAMMALS. 97

At Wagon Camp, and thence eastward to Panther Creek, the whole
country is honeycombed with their subterranean jiassages. While we
were at Wagon Camp they were unmitigated pests, throwing up little
mounds of fresh earth in our midst every day and keeping the ground
disturbed the whole time, so that it was impossible to walk in any
direction outside of the marsh without stirring up a cloud of dust. I
shot several in camp in the daytime, as they poked their heads out of
their burrows, pushing little loads of dirt before them. They throw
out the earth so rapidly that it is difficult to observe the process accu-
rately. One appeared to empty it from his pouches, but I shot him in
the act and found his j^ouches free from dirt and full of cut pieces of
roots.

On the higher slopes the winter earth plugs — the cylinders of earth
mixed with heather which in winter are pushed up into the snow from
the underground passages — remain on the ground all summer, a strik-
ing evidence of the absence of rains, for a single hard shower would
disintegrate and wash them away. They usually take the form of
irregular serpentine ridges ; but on Squaw Creek one was found which
formed a complete oval ring with radiating cylinders. A photograph
of this one, taken August 1, 1898, is here reproduced. (See fig. 31.)

Thomomys monticola pinetorum subsp. nov. Pine- woods Gopher.

Type from Sisson, Siskiyou County, Calif. No. 95152, ^ ad., U. S. Nat. Mus., Biological
Survey Coll. Collected Sept. 4, 1898, by R. T. Fisher. Orig. No. 173.

Characters. — Similar in general to T. monticola, but slightly smaller;
skull shorter and broader; color very much paler.

Color. — Upperparts pale fulvous, almost orange fulvous (in striking
contrast to the much darker colors of moiiticola and mazama)', nose
dusky; sides of head in one pelage plumbeous or slaty faintly washed
with buffy; in other pelage strongly washed with ochraceous.

Cranial characters. — Skull, contrasted with that of monticola, short
and broad, with zygomata much more widely spreading.

Measurements. — Type: Total length, 210; tail vertebrte, 70; hind
foot, 28.

Remarhs. — Common at Sisson and thence up to Wagon Camp, grading
gradually into T. monticola.

Dipodomys californicus Merriam. Kangaroo Rat.

Common in the manzanita chaparral on the south side of Shasta
from Squaw Creek Valley, near McCloud Mill, up along the road to
Wagon Camp, as far at least as an altitude of 4,800 feet, where their
unmistakable tracks abounded in the dusty soil. In Shasta Valley
they are exceedingly abundant and destructive to grain, according to
complaints of the ranchmen. Here W. H. Osgood found their little
trails winding about through the sage brush in all directions, and saw
fresh tracks in the road every morning.
21753— No. 10 13

98 NORTH AMERICAN FAUNA. [no. 16.

Perognathus mollipilosus Coues. Mountain I'ocket-Mouse.

Common in tlie mauzanita chaparral, a little below Wagou Camp,
where four were caught in July by E. T. Fisher. But the most extra-
ordinary locality at which the species was found — and for that matter
the most remarkable and abnormal place in which any sj^ecies of the
family has ever been found — is a subalpine pumice basiu near timber-
line at the head of Panther Creek, where Walter K. Fisher discovered
it and caught two the night of July 18. Later, six more were secured
at the same place.

In Shasta Valley Vernon Bailey and W. H. Osgood found abundant
signs of some species of PerognafhuSj but did not obtain specimens.
The species is probably P. ^arr?/.s, which is common in the adjacent
Klamath Basin.

Erethizon epixanthus Brandt. Porcupine.

Apparently common, and yet not a specimen was obtained. Their
characteristic guawings on the trunks of small trees were seen at
many ])oints around the mountain, usually in the Hudsonian or upper
part of the Canadian zone. They were common among the dwarf
timberline white-bark pines on the north sides of both Shasta and
Shastina; and in a small forest of young Shasta firs between Mud
Creek Canyon and Cold Creek. Xear timberline we several times
found small trees whose tops had been gnawed in winter when they
protruded above the snow. In a single instance fresh tracks were
seen in the trail between Wagon Camj) and Squaw Creek Camp ^by
Vernon Bailey). And on August I our fsivorite mule came into camp
with porcupine quills in his nose. C. II. Townsend found porcupines
in surprising abundance in Lassen County, south of Shasta, in 1883
and 1881, and gives an interesting account of their habits.

Zapus trinotatus alleni Elliot. Sierra Jerboa.

Fairly common in damp places on and near the mountain. Twenty
specimens were collected — fifteen in the Canadian zone in Mud Creek
Canyon near the mouth of Clear Creek (alt. 0,700 feet), two near the
uj^per part of Mud Creek (alt. 7,900 feet), and three at Wagon Camp.

One of the most attractive spots near Wagon Camp is a grove of
ponderosa pines in which the ground is carpeted with strawberries
and scarlet painted cups, mixed with ferns and scattered clumps of
serviceberries. The soil, while not wet, receives enough moisture
from the little streams that sink into the ground a few rods above to
enable these plants to grow in such profusion that they form a con-
tinuous meadow — 'Castilleja meadow' we called it, from the abundance
of painted cujis. Here the jerboas abound. We saw several in the
daytime, leaping about like frogs in the dense vegetation, and caught
one or two in our hands.

In Mud Creek Canyon, W. H. Osgood informs me, they were also
frequently seen in the daytime, in wet places under the white hellebore
( Veratrum caUfornicum).

OCT., 1899.

MAMMALS.

99

Zapus pacificus Merriam. Valley Jerboa.

Only two specimens of tbis little-known species were secured and
one of these was destroyed in the trap. They were caught in thickets
on the banks of Little Shasta Creek September 20 by R. T. Fisher.

Ochotona schisticeps (Merriam). Cony; Pika.

Eelatively rare and confined to small and widely separated colonies.
During- our circuit of the mountain, made near timberline the latter
part of July, we saw what we took to be signs of conies among rocks
east of Mud Creek Canyon, but finding no more believed we had been
mistaken, until the evening of July 24, when we camped on some rivu-
lets of snow water on the north side of Shastina. Here we found a
small scattered colony reaching up in the slide rock from about 8,000
to nearly 10,000 feet, and a specimen was secured by Vernon Bailej'.
The next day we found signs in Cascade Gulch a mile or two northwest
of Horse Camp. Later,
when camjDcd in the al-
pine hemlocks ou the
small west branches of
Squaw Creek, we found
a colony in the slide
rock close by. Conies
were afterwards found
on both sides of Red
Butte and on the east
side of Gray Butte, and
Osgood heard one near
the head of Mud Creek
Canyon. In all, 14 speci-
mens were collected.

This species differs in
habits and voice from
those of the Rocky Moui i -
tains; it is less noisy and less often heard in the middle of the day, for
which reason it is more apt to escape detection, and its common note,
instead of the usual 'bleat,' is a loud shrill e/i' e/«,' or eh'' eW eh\ It
seems to be most active in the late afternoon and on moonlight even-
ings, and its voice is heard at all hours of the night.

On most mountains where conies live, their well known accumula-
tions of plants of various kinds, cut and piled on the rocks to dry, are
conspicuous objects. But on Shasta, where I often saw the animals
carrying freshly cut plants to their dens in the slide rock, I failed to
find a single 'haystack.' In one place a few fresh stems of Polygonum
neicherryi, with its large broad leaves, were seen, and in another a large
accumulation of old brown leaves of the same species mixed with a larger
quantity of Phyllodoce empetri/ormis — apparently left over from the
previous year. But the only real 'haystack' found on the mountain by

Fig. 33. — Rock cony (Ochotona nchiaticeps) — Photograplied by
r. Steplieus.

100 NORTH AMERICAN FAUNA, [no. 16.

any of tlie joarty was discovered on tbe east side of Gray Butte Septem-
ber 25 by Veruon Bailey. It contained Upilobinm sincatum, Holodiacus
discolor J Monardella odoratissima, Hieracium horridum, Geanothus velii-
tinni^, and two species of grass. The bulk of the material was Epilo-
hium and jllonardclla.

On the west slope of Goose Nest Mountain, just east of Little Shasta
Valley, Walter K. Fisher found conies common in an area of slide rock
which extends in a practically unbroken stretch from the top to the
bottom of the mountain. I have jiot seen the specimens.

Lepus nuttalli Bachman. Sagebrush Cottontail.

Several seen and two secured by W. H. Osgood in the sagebrusli in
Shasta and Little Shasta valleys, near the north base of the mountain.

Lepus klamathensis sp. nov. Klamath Eabbit.

Tt/jjc from Fort Klamath, Oregon. No. 92248, 9 acL, U. S. Nat. Mus., Biological Purvey
Coll. Collected Jan. 2.5, 1898, by B. L. Cunningham. Orig. No. 86.

Characters. — Similar to L. colunihiensis Rhoads, but color fulvous
instead of yellowish, with a distinct white stripe on hind foot; skull
characters distinctive.

Color. — Summer pelaf/e: Upperparts grizzled fulvous and black, the
fulvous rather pale and dull, but not at all yellowish as in columhiensis;
head, face, and pectoral collar dull fulvous; chin, throat, and belly
white; a white stripe, sometimes irregular, extending along full length
of upper surface of hind foot, on inner side, and usually including toes.
Winter pelage: Either snow white all over, or like summer pelage
but with black hairs much more plentiful.

Cranial characters. — Skull similar to that of columhiensis but some-
,what smaller and narrower; interorbital breadth at anterior notch less;
bullic decidedly smaller (smallest of the americanus-bairdi-icashiugtoni
group); outer face of jugal very deeply grooved anteriorly, and with
upper ridge reaching anteriorly beyond end of groove.

Measurements. — Type: Total length, 432; tail vertebriP, 28 ; hind foot,
127. Average of 3 specimens from type locality: Total length, 410;
tail vertebrae, 39; hind foot, 126.

Bemarl's. — Lepus llamathensis is a member of the americanns-hairdi-
irashinf/toni group. In color it is intermediate between the yellowish
columhiensis and the dark fulvous irashingtoni. In cranial characters
it agrees best with columhiensis^ particularly in the great length of the
postorbital processes, but in the small bulltB and peculiar form of the
jugal it differs from all known members of the group.

This rabbit is common in the alder thickets in marshy places and
along streams near Fort Klamath, Oregon, from which place the late
Major Chas. E. Bendire sent me several specimens in the winter of
1883-84; and from which we have recently obtained additional spec-
imens from B. L. Cunningham.

OCT., 1899.] MAMMALS. 101

A carious feature about Lepus Jdamathensis is the fact that it some-
times does, aud sometimes does not, turn white iu winter. In the
Biological Survey collection we have brown specimens killed late in
January aud white ones killed early in April.

This species was not obtained on Shasta, but rabbit dung, supposed
to belong to it, was found in many places, particularly under the dwarf
Pinus albicauUs on the timberline ridges. Rabbit signs aud tracks
were seen also in the manzanita chaparral, but as no specimens were
secured the species is a matter of conjecture.

Lepus californicus Gray. California Jack Rabbit.

Occurs in Shasta Valley at the north base of the mountain. Several
were seen and one was killed near Edgewood September 30 by W. H.
Osgood.

Odocoileus columbianus (Richardson). Columbia Black-tail Deer.

Abundant on Shasta and throughout the surrounding region. Even
at Wagon Camp, which probably is visited by more hunting parties
than any other part of the mountain, deer were numerous, and their
well-beaten trails were in constant use during our stay. At first the
animals were commonest in the lower part of the Shasta fir forest,
where for a long time they were not driven away even by the frequent
shooting of our bird collectors. When we had been at Wagon Camp a
week they were still common within an eighth of a mile. Later, how-
ever, they became less numerous in the open forest and more abundant
in the dense chaparral of manzanita aud buck-brush a little lower
down. They were common also on Red Butte, and along all of the
streams and canyons on the west, south, and southeast sides of the
mountain. On the west side, where water is scarce, numbers used
to visit the pools in Cascade Gulch, northwest of Horse Camp. In Mud
Creek Canyon their trails were so abundant as to form almost a mesh-
work. When we visited this canyon first, July 22, Vernon Bailey saw
eight deer; and several of us, resting on the west rim of the canyon,
watched a doe and fawn on one of the trails on the opposite side.
They were so plentiful in a canyon about a mile east of Squaw Creek
that I named the place Deer Canyon. Several times during the season
does with spotted fawns were seen in the Shasta fir forest. A yearling
'spike-buck' killed on Squaw Creek by Vernon Bailey August 7 wasiu
the velvet, aud his worn summer coat was scant and faded. Another
'spike-buck,' killed in the mountains west of Scott Valley September
15, was in the fresh gray winter coat, with only a few red hairs of the
summer coat left.

In September the old bucks, which had not been observed earlier,
climbed the mountain and began to appear on the higher ridges, where
they travel extensively in the timberline tongues of dwarf white-bark
pines. On September 18 I followed the tracks of two large bucks along
the upper part of Panther Creek and found where they had bedded

102 NORTH AMERICAN FAUNA. [no. 16.

close togetlier under a low Shasta tir on a steep sloi:>e, from which they
could overlook the country below.

O. H. Townsend, in his notes on the mammals of northern California,
gives an interesting account of this deer.

Odocoileus hemionus (Rafinesque). Mule Deer.

In the region east of Shasta, where the Columbia black-tail is the
prevailing species, C. H. Townsend occasionally found the mule deer.
" But in Lassen County, a hundred miles farther south, the reverse was
found to be the case," and he saw nothing of the Columbia black-tail.
This was in 1883 and 1884. The mule deer was not observed on Shasta
by our party, although the tracks of an immense buck, seen by me
early in August on the rim of Mud Creek Canyon, may have been made
by it. Sherman Powell, in a recent article in Forest and Stream (April
27, 1899), states that mule deer are plentifnl a little east of Shasta " on
and around Glass Mountain, and also on the northeast slopes of Black
Fox Mountain."

Cervus occidentalis Ham. Smith. Elk.

One of our party, E. T. Fisher, was informed by George B. Mitchell,
county surveyor of Siskiyou County, that elk were shot in the neigh-
borhood of Sisson as late as the early seventies. They were formerly
abundant on and about Shasta, particularly in Squaw Creek Valley
and Elk Flat, and used to range along the Scott Mountains, and thence
westerly to the coast, where a few still exist.

Antilocapra americana Ord. Prong-horn Antelope.

Antelope, we were told, still inhabit the open pine forest east and
northeast of Shasta. Formerly they were common in Shasta Valley and
ranged west into the foothills of the Scott and Siskiyou mountains.

The following information regarding their distribution was obtained
by Walter K. Fisher: In winter they ranged in the country between the
Edgewood divide and the foothills of the Siskiyou mountains north of
Hornbrook, extending into the low valleys west of Shasta Eiver. They
were most plentiful in the region between Little Shasta and Gazelle.
Mr. Masgrave, one of the tirst settlers in Little Shasta Valley, is
authority for the statement that formerly they frequently herded with
his cattle. In Scarface Valley, west of Gazelle, he once saw a large
herd which contained not less than two thousand animals.

In summer the antelope ranged extensively through Goose Nest
Mountain and wooded valleys in Butte Creek region, as well as in Shasta
Valley, Big Valley, Fall River Valley, and about Tule Lake, Klamath
Falls, and Goose Lake. At present only a small herd remains. They
stay in the remoter valleys east of the mountains and rarely come to
Shasta Valley. In the summer of 1898 three were seen on the road
between Little Shasta post-office and Butte Creek.

OCT., ]899.]

MAMMALS. 103

Ovis canadensis Sliaw. Bighorn; Moautaiu Slieep.

The bighorn no longer inhabits Shasta, but its bleaching bones still
remain. In early days, and as late as the seventies, many were killed
here by J. H. Sisson, of Sisson Tavern. Sheep Rock, at the northeast
base of the mountain, was one of their favorite and latest resorts, but
probably was not nsed during the breeding season. In 1SG8 George
B. Mitchell saw a band of twenty near the head of Mud Creek Canyon.
In 1883 C. H. Townsend found numbers of their horns and bones scat-
tered about everywhere on Sheep Eock, and saw the complete skeleton
of a bighorn at the foot of Mud Creek glacier, high np on Shasta. An
old skull was found on Red Butte by Vernon Bailey during our stay.

Canis lestes Merriam. Mountain Coyote.

Common high up on the mountain, frequenting the pumice slopes
above timberline, where their tracks were often seen. Coyotes were
especially common east of Mud Creek Canyon, where R. T. Fisher
saw two early in August. As no specimens were secured, the species is
uncertain, but since C. lestes is the animal inhabiting the Sierra farther
south and (in a somewhat less typical form) the Klamath country farther
north, it is assumed to be the mountain coyote of Shasta.

Canis ochropus Esch. Yalley Coyote.

Common in Shasta valley and also at Sisson. R. T. Fisher heard them
nearly every night during his stay at Sisson, from the end of August
until the middle of September; and about the end of September Ver-
non Bailey and W. H. Osgood heard them howl at night in the south
end of Shasta valley. A little farther north, in the flat valley near
Montague, I saw a highly colored one at close quarters in the day time
August 31. In July, 1899, Walter K. Fisher secured one in Shasta
Valley.

Vulpes macrourus Baird. Mountain Red Fox.

Common on the upper slopes above timberline, Avhere fresh tracks
were seen nearly every day ; but the animals were very wary and refused
to enter our traps.

Urocyon californicus townsendi subsj). nov. Townsend Gray Fox.

Type from Baird, Shasta County, Calif. No. 14130, U. S. Nat. Museuiu. Collected
November 11, 1893, by C. H. Townsend. Orig. No. 49.

Characters. — Similar to IT. californicus Mearns, but ears decidedly
smaller; fulvous tints everywhere darker and richer; rostrum broader;
zygomata broader anteriorly; carnassial teeth above and below, larger
and thicker. The color of the upperparts and tail is grizzled gray and
black, as in californicus, but the fulvous of the ears, legs, and under-
parts is very much darker and somewhat more extensive. The sides of
neck and posterior part of throat are rufous, in sharj) contrast with the
restricted white of the chin and anterior part of throat; the ears are

104 NORTH AMEKICAN FAUNA. [no.16.

dark grizzled fulvous; the white stripe on the hind foot of cali/or ulcus
has disappeared and is represented by a pale streak.

Bemarls. — Gray foxes are abundant in tlie Transition zone about
the base of Shasta, and although no specimens were secured by our
party a number were obtained by 0. II. Townsend in 1883 along the
McCloud Eiver.

Felis oregonensis Rafinesque. Oregon Puma; Mountain Lion.

Rather rare, but formerly common on Shasta. In July Bailey saw
fresh tracks among the Shasta firs a little below Red Butte. C. H.
Townsend obtained seven specimens in Shasta County in 1883 and
1884, several of which were killed near the fish hatchery on the McCloud.
He states: "It is i)ractica]ly impossible to raise colts in the Shasta
County hills on account of these pests. They destroy many hogs and
young cattle also, but do not present so serious an impediment to the
keeping of these animals as in the case of horses. Mr. J. B. Campbell,
who trapped two panthers for me in 1883, told me that he had actually
never seen more than two or three of the numerous colts born on his
stock range, as they had been killed and devoured by i)anthers soon
after birth."

Ljmx fasciatus pallescens subsp. nov. Wild-cat.

Ty])e from sontli base of Mount Adams, near Trout Lake, Washington. No. 76585, <?
ad.j U. S. Nat. Mus., Biological Survey Coll. Collected January 10, 1895, by
D. N. Kaegi. Orig. No. 23.

Characters. — Similar to L. fasciatus, but slightly smaller and every-
where very much paler, ^particularly the head and face f basal black
ear i)atch (in winter pelage) indistinct or absent; gray ear patch larger;
general color hoary gray, contrasted with the dark rich rufous of
fasciatus. Skull like that of fasciatus, but slightly smaller; carnassial
teeth and second upper molar less swollen.

Remarlcs. — Specimens from the region about Shasta (Klamath country
on the north and Pitt and McCloud rivers on the south) differ slightly
from Trout Lake specimens; they show less tendency to spotting and
have slightly larger carnassial teeth and audital bulla'.

This wild-cat is fairly common in the chaparral of the basal slopes
of Shasta. One was seen a little below Wagon Camp, and tracks were
several times noted by Yernon Bailey in the trail a little above Wagon
Camp. Tracks were seen also in Shasta Valley by W. H. Osgood. In
1883-81 C. H. Townsend trapped many wild-cats along McCloud River
and the lower part of Squaw Creek, where they were "apparently as
numerous as [gray] foxes, and as easily secured." Most of these belong
to the present species, but one belongs to the following:

Lynx californicus (Mcarns). California Wild-cat.

A specimen which I provisionally refer to this species was obtained
by C. H. Townsend at Baird, on McCloud River, in 1883. It is a
much smaller animal than L. fasciatus paUescens and has conspicu-
ously smaller teeth.

OCT., 1899.1

MAMMALS.

105

Gulo luscus (Liim.). Wolverine.

We liave no positive record of the wolverine from Shasta, hut it is
known to occur on both the Cascades and tlie Sierra, and one was killed
near Carberry Eanch, between Mounts Shasta and Lassen, about the
year 1893 (reported by C. P. Streator).

Wolverines are notorious wanderers and it is most i^robable that they
occur on Shasta at intervals, even if they do not x>ernianently live
there — which is by no means certain.
Taxidea taxus (Schreber). Badger.

Very rare on Shasta. In the trail between Wagon Camp and Squaw
Creek Camp fresh tracks were seen August 4, by Yernon Bailey, and
in Shasta Valley, at the north base of the mountain, badger holes were
reported by W. H. Osgood, who also found a dead badger in the road
a few miles east of McCloud Sawmill.
Mephitis occidentalis Baird. Large Skunk.

Common in the low country around Shasta. On 'Horse Trail,' lead-
ing up the mountain from Sisson, Bailey saw their tracks as high as
an altitude of 6,500 feet. At Sisson they are particularly abundant in
the wet meadows and thickets, where in September B. T. Fisher caught
two along Cold Creek. In Shasta Valley they are also common and
several were caught by W. K. Osgood and E. T. Fisher.
Spilogale latifrons Merriam. liittle Spotted Skunk.

l^Tot obtained by us, but known to occur in the general neighborhood
of Shasta, both north and south of the mountain. In March, 1SS4,
C. H. Townsend caught one on McCloud Kiver, about midway between
the mouth of this stream and Mount Shasta.

Fig. 34.— Mink. (Drawn by Ernest S. Thompson.)

Lutreola vison energumenos (Bangs). Pacific Mink.

Common along the streams at Sisson, where R. T.Fisher caught two,
one on Cold Creek, the other ou the upper Sacramento River.
21753— No. 1(5 11

106

NORTH AMERICAN FAUNA.

[no. 16.

Mustela caurina Merriam. Pacific ]\raitcii.

Common in parts of the dense forests of Shasta, i)articnlarly on the
east side, where many are caught every winter by trappers. We did
not succeed in obtaining- a specimen, although a line of 'meat trails'
was kept out fully a month. Still, fresh tracks were seen iu the mud
along Sijuaw Creek a short distance below camp by Vernon Bailey.

Fig. 35. — Marten. (Drawn by Ei uest b, iliompson.)

Mustela pennanti Erxleben. Fisher.

Said to occur in the dense forest on the east side of Shasta, whence
skins are brought to Sisson for sale nearly every winter. C. H. Town-
send obtained two ou McCloud Eiver, about halfway between the
mouth of the river and JNIount Shasta, in February, 1884.

Lutra hudsonica Lacepede. Otter.

Occurs along the upper Sacramento and McCloud rivers; not noted
by our j^arty.

Putorius arizonensis Mearns. ^Mountain Weasel.

Api^arently not common. Early in August three were caught in
Mud Creek Canyon, near the mouth of Clear Creek, in traps set in
Aplodoniia runways. It is quite possible that another species, P.
xantlioijenyx oyc<jonMs^ inhabits the low country about the base of the
mountain.

Bassariscus astutus raptor (Baird). Cacomistle; liing-tail Cat.

Kot observed by us, but doubtless occurs about the basal slopes of
Shasta, as it has been captured both north and south of the mountain.

OCT., 1899.1 MAMMALS. 107

In February, 1884, C. H. Townsend caught oue on McCloud Iliver,
about halfway between the mouth of the river and Mount Shasta; he
also mentions a Pitt River miner who had a pair alive ; and C. P. Streator
reports it from old Fort Crook and Carberry Ranch. In September,
189G, one of our collectors, E. A. Preble, caught one in upper Rogue
River Valley, near Prospect, Oregon.

Procyon psora pacifica subsp. nov. Pacific Raccoon.

Type from Kechelus Lake, Cascade Moiautaiu.s, Washiuston. No. 93137, aiL, U. S.
Nat. Mus., Biological Survey Coll. Collected Jan. 15, 1898, by C. Hansen.

Characters. — Coloration dark; ground color dark gray instead of
buffy; tail rings continuous (not interrui)ted along median line below);
last premolar, first molar, and audital bulla* larger than m psora.

Color. — Upperparts dark gray everywhere profusely mixed with
and obscured by black hairs; underparts sootyidumbeous sparingly
sprinkled with long buffy whitish hairs; sooty collar under throat con-
tinuous and much darker than in psora ; black rings on tail continuous ;
pale rings, particularly the last one, obscured above by black hairs.

Measurements. — Type (from dry skin): Total length, 940; tail verte-
bra^, 310; hind foot, 115.

Remarls. — This new raccoon is most typical in the northwest coast
region, particularly about Puget Sound and along the basal slopes of
the northern Cascades. Specimens from Pitt River and Little Shasta
Valley are much nesbrer 2)acijica. than 2)sora.

Raccoons do not occur on the higher slopes of Shasta, but are common
about its base. In Little Shasta Valley W. H. Osgood and R. T. Fisher
found them particularly abundant about the middle of September, and
caught several in traps set under j>rune trees. The animals were caus-
ing much annoyance to the fruit-raisers by nightly visits to the
orchards.

Ursus americanus Pallas. Black Bear.

Abundant. When we reached Shasta the middle of July, black
bears were exceedingly common throughout the Shasta fir forest. On
July 17, in going from Wagon Camp to timberline, we saw fresh tracks
of a dozen along Panther Creek, and on the way down, three hours
later, saw where four had crossed our trail after we had gone up. Their
well-worn trails abounded along Panther Creek and were traversed
daily until about the end of July, when, after our collectors had been
shooting for nearly two weeks, the bears became alarmed and moved
down into the numzanita chaparral below the Shasta firs, Avhere they
remained the rest of the season.

Ilrsus horribilis Ord. Grizzly or Grisly Bear.

Formerly abundant in the Shasta region; now exceedingly rare or
al)sent. W. H. Osgood was told when in Shasta Valley that a few
years ago a huge grizzly known as 'Old Clubfoot,' which had been shot
at repeatedly, was killed near (ioose Nest Mountain, just north of
Shasta.

BIRDS OF SHASTA AND VICINITY.

(All dates except where the year is stated are for th.e year 1808.)

1. Podilymbus podiceps. Pied-billed (Irebe; Hell Diver.

Several seen in the pond at Big Spring, in Shasta Valley, September
17, by W. H. Osgood. C. H. Townsend obtained it " on ponds near the
base of Mount Shasta in summer" in 1883 or 1884.

2. Anas boschas. Mallard.

W. H. Osgood shot a mallard in Lower Squaw Creek, near Warm-
castle Soda Springs, September 13, and found the species common at
Big Spring, in Shasta Valley, September 17. In 1883 O. H. Townsend
"found mallards August 1, at the base of Mount Shasta, in certain wet
meadows where, in all probability, they had nests."

3. ftuerquedula cyanoptera. Cinnamon Teal.

The commonest duck on the small ponds and streams in Shasta Val-
ley September 17-20 (Osgood).

4. Spatula clypeata. Shoveller.

Several seen in Shasta Valley September 17 by W. H. Osgood.

5. Dafila acuta. Pintail.

A flock of six seen in Shasta Valley September 18 by W. H. Osgood.

6. Branta canadensis. Canada Goose.

Three seen at Big Spring, in Shasta Valley, September 18 (Osgood).

7. Ardea herodias. Great Blue Heron.

Seen at Sisson July 13 by W. H. Osgood, and repeatedly observed
during the first half of September by R. T. Fisher; several seen in
Shasta Valley September 17-20 by W. H. Osgood; common along
Shasta River (W. K. Fisher).

8. Porzana Carolina. Sora Rail.

One seen in Shasta Valley September 18 by W. H. Osgood.

9. Fulica americana. Coot; Mud Hen.

One seen among the numerous ducks in the pond at Big Spring, in
Shasta Valley, September 18 (Osgood).

10. Gallinago delicata. Wilson Snipe.

In Shasta Valley one was seen by W. H. Osgood September 18; and
I saw one at Edgewood August 2, 1899. On a small tributary of
MeCloud River, flowing from the east base of Mount Shasta, one was
shot by C. H. Townsend July 25, 1883.

109

110 NORTH AMERICAN FAUNA. [no. 16.

11 . Ereunetes occidentalis ( ?) Western Sandpiper.

A Hock of small sandpipers snpi)osed to be this species was seen by
W. H, Osgood in Shasta A'alley September 19.

12. Totanus flavipes. Yellow-legs.

A wader believed to be this species was heard in Shasta Yalley Sep-
tember ID by E T. Fisher.

13. Helodromas solitarius cinnamomeus. Western Solitary Sandpiper.
I^ot observed by us, but in 1883 a male was seen at Sisson on August

3 by C. H. Towusend.

14. Actitis macularia. Spotted Sandpiper.

C. H. Towusend secured a spotted sandpiper at Sisson August 2,
1883. Late in July, 1899, Walter K. Fisher found it rather common on
Shasta River.

15. JEgialitis vocifera. Killdeer.

Common at Sisson, where from July to early September a small flock
was always crying and feeding in the meadows between the town and
Sisson Tavern. Abundant and noisy in Shasta Valley September
17-20 (W. H. Osgood).

16. Oreortyx pictus plumifems. Plumed Mountain Quail.

Fairly common, but not often seen, in the Shasta fir belt and the
manzanita chaparral along its lower edge. Two or three broods were
found near Wagon Camp the latter part of July, when the young were
hardly a third grown. Several were seen and killed high up on Mud
Creek Canyon August 8 by R. T. Fisher and W". H. Osgood, who also
saw several flocks in Shasta Valley September 17-20. At Sisson, the
first half of September, R. T. Fisher found many small flocks in the dry
woods and chai)arral west of the village.

17. Lophortyx californicus vallicola. California Valley Quail.

Not found on the mountain, but common in Little Shasta A'alley,
where W. H. Osgood secured a specimen September 19. In August,
1883, C. H. Towusend saw theui "in considerable numbers at the base
of Mount Shasta," near Sisson.

18. Dendragapus obscurus fuliginosus. Sooty Grouse.

Fairly common in the Shasta fir forest, and less so at higher alti-
tudes. At Wagon Camp, during the last half of July, we often saw a
hen grouse with half a dozen young; and in a rocky place among the
alpine hemlocks near Stjuaw Creek, in September, we several times
ran across a small flock, all full grown, feeding among patches of mouu
tain chinquapin and the dwarf mountain manzanita {Arctostajyhi/Ios
nevadensis). Berries of the latter formed the principal contents of the
crop of one killed by Vernon Bailey August 21. Mixed with these
berries were seed capsules of Fentstemon (/racilenfxs and a few large

OCT., 1899.] BIRDS. Ill

ants. Usually tlie grouse Avere found singly or in bunches of two or
three in the forest. When disturbed they generally flew up into the
tall Shasta firs, where, instead of remaining motionless like many
grouse, they walked about among the branches, stepping deliberately
from limb to limb, but keeping on the opposite side from the enemy, so
that it was almost impossible to see them.

The old males were sometimes found high up on the lava ridges
among dwarf i*/w?r9 albie((idi>t^ 1,000 feet or more above the continuous
forest. When flushed they usually spread their wings and soared down
the steep mountain side until lost from sight in the forest below. The
'hooting' or 'booming' of the old males, so often heard in the northern
Cascades, was not heard by any of our part}^ on Shasta.

19. Zenaidura macroura. Mourning Dove.

Fairly common at Sisson. In Shasta Valley W. H. Osgood found
them abundant September 17-20.

[The band-tail pigeon {Columha fasciata) was not observed in the
region by my party, but in the fall and winter of 18S3 C. H. Townsend
found it abundant in the foothills of the lower jNIcCloud River, "gather-
ing in the i^ine trees on the higher ridges in immense flocks."]

20. Cathartes aura. Turkey Vulture.

One or two seen at Wagon Camp and between Wagon Camp and
Sisson in July; one seen by Walter K. Fisher at timberline, east of
Mud Creek Canyon, about the end of July. At Sisson and in Shasta
Valley they were more common.

21. Circus hudsonius. Marsh Hawk.

Seen at Sisson the middle of July (F. A. M.) and end of August
(C. H. M.). Seen in Shasta Valley September 17-20 (W. 11. Osgood).

22. Accipiter velox. Sharp-shinned Hawk.

Rather rare. One shot near Squaw Creek Camp August 10 l)y Ver-
non Bailey; one seen on Mud Creek in August, and several in Shasta
Valley by W. H. Osgood September 17-20. C. H. Townsend shot one
at timberline on Shasta September 7, 1S83.

23. Accipiter cooperi. Cooper Hawk.

Rather rare. One killed above timberline (alt. 9,400 feet) above the
head of Squaw Creek August 21 by R. T. Fisher, In September 1883,
C. H. Townsend shot one near timberline on Shasta.

24. Accipiter atricapillns striatulus. Western Goshawlc,

One visited our camp at Squaw Creek August 28. It was shot at
but not secured. July 28, 1883, C. H. Townsend shot two young gos-
hawks near timberline on Shasta.

112

NORTH AMERICAN FAUNA.

[NO. 16.

25. Buteo borealis calurus. Western Eed-tail.

Rather common. Seen from time to time at Wagon Camp in July
and early August, frequently at Squaw Creek Camp in August and
Sei^tember, and several times above timberline. One was seen flying
over the summit of Shasta July 31 by W. H. Osgood and R. T. Fisher.
Common during fall migration in Shasta A'alley, where Osgood saw
many September 17-20, and I saw several September 29 — chiefly about
the narrow meadows bordering Shasta River. At Sisson Miss Florence
A. Merriam saw them September 3.

Fig. 36.— Ked-tail (Buteo borealis). Drawn by J. L. Kidgway.

26. Buteo swainsoni. Swainson Hawk.

Rare. July 28 I saw a nearly black Swainson hawk near an aban-
doned sawmill (alt. 4,800 feet) on the road from Wagon (amp to
McOloud Mill. In July, 1899, Walter K. Fisher saw one in Shasta
Valley.

27. Aquila chrysaetos. Golden Eagle,

Rather rare, but seen from time to time flying over the mountain,
usually at high altitudes. C. H. Townsend shot one near Sheep Rock,
at the nortli base of Shasta, August 21, 1883.

28. Haliaeetus leucocephalus. Bald Eagle.

In speaking of the bald eagle, C. H. Townsend states that when he
had climbed to the extreme peak of Shasta (alt. 14,140 feet), on July 27,

OCT., 1>!!)9.]

BIRDS.

113

1883, ''an eagle caiiie up thiougli the fog- tliat had gathered imme-
diately below us and shared with ns our rooky pinnacle above the
clouds."

29. Falco mexicanus. ]*rairie Falcon.

Several seen and one shot in the south end of Shasta Valley, at the
north base of the mountain, September 30, by W. H. Osgood, who
thinks it probable that they breed on the cliffs at Sheej) Rock.

30. Falco columbarius suckleyi. Black Merlin.

At Wagon Camp, August 8, John H. Sage and I saw a small dark
falcon, supposed to be Suckley's merlin, fly into a Shasta fir, but we
were unable to shoot it.

31. Falco sparverius. Sparrow Hawk.

Common at timberline throughout the summer. TTsually seen in the
heather patches and among the white bark pines, where they were

Fig. 37. — Sparrow Hawk {Fulro spari^erius). — Ilrawn by .J. L. Ridgway.

feeding on grasshoppers. Several were killed near timberline, east of
Mud Creek Canyon, the latter j^art of July and early August, and one
was recorded as high as 13,000 feet by Walter K. Fisher. On the west
side of Mud Creek Canyon two were killed August 11 at an altitude of
10,000 feet by Vernon Bailey. Their stomachs were distended Avith
21753— Ko. IG 15

114 NORTH AMP:RICAN fauna. (no. 16.

grasshoppers. At the north base of the iiiountaiu one was seen by me
on the narrow meadows bordering- Shasta River in Sliasta Valley Sep-
tember 29, and several were observed farther north in the valley Sep-
tember 17-20 by W. H. Osgood and E. T. Fisher, At Sisson they were
seen July 15 by Miss Florence A. ^lerriam.

32. Bubo virginianus. Great Horned Owl.

Rare. Feathers were found at two or three places on the mountain,
and at Sisson birds were heard hooting.

33. Speotyto cunicularia hypogaea. Burrowing Owl.

Not observed by us, but reported by C. H. Townsend as common ''on
the sage-covered districts north of Mount Shasta," about 15 miles from
the mountain.

34. Glaucidium gnoma californicum. California Pigmy Owl.

0. H. Townsend obtained two specimens of the pigmy owl August 7,
1883, at the big spring, a mile or two north of Sisson Meadows.

35. Megascops asio bendirei ( ?). California Screech Owl.

A screech owl was heard at Wagon Camp at different times during
the season, particularly the latter part of September, but as no speci-
men was secured there is some doubt as to the subspecies.

36. Coccyzus americanus occidentalis. California Cuckoo.

One shot and others seen in July, 1S99, by Walter K. Fisher, among
the alders and birches on Shasta River east of Edgewood.

37. Ceryle alcyon. Kingfisher.

A common resident in the neighborhood of Sisson, where it was seen
repeatedly along Cold Creek, at tlie fish hatchery, and along the upper
Sacramento ; seen also along Shasta River north of Sisson.

38. Dryobates villosus hyloscopus. Cabanis Hairy Woodpecker.
Rather rare. Two or three were seen near Wagon Camp in July;

two were collected where the Hudsonian and Canadian zones meet,
near S<|uaw Creek Camp, in August (August 9 and 17); one was seen
there Sej)tember 1, and two were shot in the Transition zone in Sep-
tember (one in Squaw Creek Valley September 13, the other at Sisson
September 30). At Sisson R. T. Fisher saw three or four, during the
first half of September, in the big firs west of Sisson Tavern.

39. Dryobates pubescens gairdneri. Gairduer Woodpecker.
Obtained at Sisson by C. H. Townsend August 3, 1883.

40. Xenopicus albolarvatus. White-headed Woodpecker.

A common breeder in the upper i^art of the pine belt, and also among
the Shasta and white firs near Wagon Camp, where several specimens
were collected, and where a nest-containing noisy young was found the
latter part of July. The nest was in a stub in a burn, with the

OCT., issiy.

BIRDS.

115

entrance about G feet from the ground. At Sissou tlie white-headed
woodpecker was seen, the middle of July, by Miss Merriani, and early
in September by R. T. Fisher. In the pine woods bordering the south
end of Shasta Valley I saw half a dozen September 29.

41. Picoides arcticus. Arctic Three-toed Woodpecker.

Kare, but evidently breeding in the Shasta fir forest of the Canadian
zone, where a male was shot near Wagon Camp by John H. Sage July

Fig. 38.— Arctic Three-toed AVoodpecker (Picoiil

— I>ru\vii hy L. A. Fuertes.

20. Another was shot by Vernon Bailey July 30 in the gap between
Red Cone and the lower end of Gray Butte, where a third individual
was seen.

42. Sphyrapicus ruber. Red-breasted Sapsucker.

Fairly common at Wagon Camp, where specimens were collected in
July by John H. Sage and Vernon Bailey. One was shot in Mud
Creek Canyon about the end of July by II. T. Fisher, and one was seen
at Sisson July 15 by Miss Merriam. September IS, one was seen in
the brush along Little Shasta Creek by W. H. Osgood. In 1883 C. H.
Townsend found the si)ecies "in midsummer iu limited numbers on the
heavily pine-timbered slopes of Mounts Shasta and Lassen.''

116 NORTH AMERICAN FAUNA, [no. 16.

43. Sphyrapicus thyroideus. Williamson Sapsucker.

A uiale was killed August 4 by R. T. Fisher at tiraberline on the
east side of Mud Creek Canyon. This was the only specimen collected,
though others were seen near the same place. August 25, 1883, C. H.
Townsend secured one at timberline on Shasta.

44. Ceophloeus pileatus abieticola. Pileated Woodfiecker.

Fairly common about Wagon Camp, where their loud hammering
and unmistakable cries were often lieard. Several times in July one
visited a tree on the edge of camj), and one was heard there the day
we left, September 25. Sei)teniber 16, W. H. Osgood and li. T. Fisher
saw three between Sisson and Edgewood.

45. Melanerpes formicivorus bairdi. California Woodpecker.

Seen from time to time among the oaks at Sisson, where one was shot
by R. T. Fisher early in September.

46. Melan'^rpes torquatus. Lewis Woodpecker.

Common at Sisson and in Squaw Creek Valley near McCloud Mill.
July 25 Miss Merriam saw one at Wagon Camp; August 2 W. H.
Osgood killed one and saw others high up on Mud Creek Canyon;
August 3 I saw a small flock in the Shasta fir forest between Mud and
Squaw creeks; August 10 Walter K, Fisher shot one near Squaw Creek;
and September 17-20 W. H. Osgood saw several in Shasta Valley.

47. Colaptes cafer. Red shafted Flicker.

Fairly common on the mountain; seen or heard nearly every day,
during the latter half of July and early August, near Wagon Cam[).
July 28 several were seen on the lower slope between Wagon Camp and
McCloud Mill; late in July and early in August several were seen high
up on Squaw and Mud creeks; August 16 one was killed by R. T.
Fisher at S(]uaw Creek Camp; and during migration, the latter half of
Sei>lember, a few were seen nearly every day a little below timberline
near Squaw and Panther creeks. At the time of our visits to the
juniper forest in Shasta Valley, September 19 and 2i), flickers were
common and were probably feeding on the Juniper berries. At Sisson,
Florence A. Merriam found them common, the middle of July, and
R. T. Fisher, the first half of September.

48. Phalaenoptilus nuttalli. Poor-will.

The unmistakable note of the poor- will was heard in July at Wagon
Camp, where the species doubtless breeds. One of the birds was seen
on Lower Squaw Creek near VVarmcastle Soda Springs August 13 by
W. H. Osgood. Others were heard near Edgewood by Walter K.
Fisher.

49. Chordeiles virginianus. Nighthawk.

Often seen flying at Wagon Camp, on the lower edge of the Shasta
fir forest, the latter part of July. Several flocks were seen at Sisson,
the first half of September, by R. T. Fisher.

OCT., 1899.] BIRDS. 117

50. Chaetura vauxi (?). Vaux Swift.

A couple of miles below Wagon Camp a small black swift was seen
by Vernon Bailey about July 1*4. At Sisson numbers of swifts were
seen September 1 and September 10 by E. T. Fisher, and on Septem-
ber 3 by Florence A. Merriam, but they flew too high to be shot.

51. Aeronautes melanoleucus. White-throated Swift.

Several were seen high up on ^lud Creek Canyon early in August
by W. H. Osgood.

52. Calypte anna. Anna Hummingbird.

Apparently breeding at Wagon Camp, where one was shot Julj^ 22
by John H. Sage. Breeds on the lower McCloud lliver (Townsend).

53. Selasphorus rufus. Rufous Hummingbird.

The commonest hummingbird of Shasta, breeding, apparently, from
the lower edge of the Shasta firs to timberliue, though it is possible
that those seen at high altitudes had moved up to feed from the painted
cups in the heather meadows after the breeding season was over. At
Wagon Camp, where they were abundant in July and early August,
they seemed to feed chiefly from the scarlet painted cup [CasUUeja
mimata).

54. Stellula calliope. Calliope Hummingbird.

Nearly as abundant as SehisphovKs rnfus, and the commoner of the
two at high altitudes. In early August both species were constantly
hovering over the superb flowers of the scarlet paint brush {GastUleJa
mlnlata) in the heather meadows near timberliue. They were seen also
visiting the large yellow blossoms of MimulHs implexus, which singular
species forms mucilaginous beds in the little streams at and below tim-
berline. Walter K. and R. T. Fisher found them abundant in Ash
Creek Canyon about the first of August, particularly among the painted
cups and delphiniums.

[Trochilufi ale.vandrl is recorded by C. H. Townsend as breeding along
the lower McCloud, but was not obtained by us. In this connection
it should be remembered that we did not collect hummingbirds in the
low country, either at Sisson or in Scjuaw Creek Valley, j

55. Tyrannus verticalis. Arkansas Kingbird.

At Gazelle, on the west side of Shasta Valley, Vernon Bailey saw
three on telegrai)h wires October 31. C. H. Townsend records the
species as breeding on the lower McCloud River. In July, 1899, Wal-
ter K. Fisher found this kingbird common in Shasta and Little Shasta
valleys.

50. Sayornis saya. Say Phoebe.

One was seen about an old windmill in Shasta Valley September 19
by W. H. Osgood.

118 NORTH AMERICAN FAUNA. [No.ir,.

57. Contopus borealis. Olive-sided Flycatcher.

Coiniuon at Wagon Camp, where several pairs reared families early
in the seasou, aud where, on July 22, Miss Merriam saw a parent l)ird
feeding young in the nest, whicli was in a fir tree 30 to 40 feet from the
ground. In July the species was also seen at Sisson, and in Mud
Creek Canyon near the mouth of Clear Creek, on the north side of the
mountain.

58. Contopus richardsoni. Western Wood Pewee.

A fairly common breeder at Sisson, where Florence A. Merriam saw
old birds feeding young in the nest (in a fir tree) July 15.

59. Empidonax difficilis. Western Yellow-bellied Flycatcher.

On 'July 29 I shot one in a thicket of cherry bushes {CerasKs emar-
ginata) on the boundary between the Canadian and Transition zones, a
little west of Wagon Camp, where it was doubtless breeding.

60. Empidonax hammondi. Hammond Flycatcher.

A common breeder near Wagon Camj), in the lower edge of the
Shasta firs, where several were collected late in July. Near Squaw
Creek Camp, at the upper edge of the Shasta firs, one was shot August
21, and in Mud Creek Canyon at the mouth of Clear Creek, one was
shot August 7.

01. Empidonax wrighti. Wright Flycatcher.

One was shot at AVagon Camp July 24 by John H. Sage.

02. Otocoris alpestris merrilli. Dusky Horned Lark.

Common in places in Shasta Valley, particularly about Montague,
where a number were secured by Walter K. Fisher. This is doubtless
the form reported by C. H. Townsend, under the name ruhea, as
found ''in limited numbers on the sage-covered districts north of Mount
Shasta in midsummer." The locality referred to, he tells me, is in
Shasta Valley, about 15 miles from the mountain.

63. Pica pica hudsonica. Magpie.

One was seen tlying over the east side of Shasta Valley, near Sheep
Rock, September 29 by Vernon Bailey and W. H. Osgood.

64. Cyanocitta stelleri. Steller Jay.

The form of Steller jay of the Shasta region is intermediate between
true stelleri and the Sierra subspecies, //v>/*/rtZ/'A;.

It is one of the commonest, noisiest, and best-known birds of the
region. Early in the season it was rarely seen above the lower part of
the Canadian zone, and was most numerous in the Transition; but on
August 2 one came all the way up to Squaw Creek Camp, in the alpine
hemlocks, and a few days later a small flock was encountered, scream-
ing, in Mud Creek Canyon at the mouth of Clear Creek. The latter
half of September they were common at high altitudes and paid daily
visits to our camp on upper Squaw Creek. At Sisson, apparently, they
are always common.

OCT., ] 899.1 BIRDS. 119

65. Aphelocoma californica. California 'lay.

Fairly commou at Sisson, and more abundant in the lower country
farther north. Seen by Vernon Bailey near Ciazelle August 31, by
W. H. Osgood among the junipers in Shasta Valley Sej^tember 17 to
20, and by me in the undergrowth along the edge of the open pine
and oak forest bordering tlie south end of Shasta Valley, where it
was fairly common, September -J9.

66. Perisoreus obscurus. Oregon Jay.

Unaccountably rare on Shasta during our stay. On August 6, when
in a deiise part of the forest east of the lower end of Gray Butte, I saw
a flock of Steller jays, and with them several birds I took to be Oregon
jays in the dark plumage of the young. August 20 Vernon Bailey shot
one on Horse Cami) Trail at an altitude of 6,600 feet, and two days later
saw ten in the fir forest between Squaw Creek and Mud Creek Canyon.
September 28, on his way around the mountain, he saw three above the
point where the wagon road crosses Ash Creek, at an altitude of about
5,000 feet, by far the lowest point at which the species was seen. On
July 29, IS99, Walter K. Fisher saw about 15 Oregon jays on Horse
Camp trail.

In 188.'> C. H. Townsend saw four small flocks (July 30 to September 7)
in the heavily timbered forests of Mount Shasta.

67. CorvTis americanus. Crow,

A few seen at Sisson. In July, 1889, Walter K. Fisher found them
common in Shasta Valley.

68. Nucifraga columbiana. Clark Crow; Nutcracker.

Clark crows are among the most common, most characteristic, and
most interesting birds of the higher slopes of Shasta. In summer they
are closely restricted to the Hudsonian zone and adjacent rocky slopes
immediately above timberline, but in fall they wander far and wide in
search of food and are liable to be seen almost anywhere. Two or
three, api^arently young of the year, visited Wagon Camp, at the lower
edge of the Shasta firs, as early as August 8; and in September it was
not unusual to see small flocks or single individuals iiying over the
chaparral belt between Wagon Camp and Sisson.

The usual food of the Clark crow — the large nut-like seeds of Pin us
albicaulis — having failed in 1898, the birds were feeding mainly on
insects. The stomachs of specimens killed at extreme timberline con-
tained in some cases grasshoppers only, in others chiefly beetles (Cole-
optera) ; of those killed lower down, in the alpine hemlocks and Shasta
firs, chiefly small hairless greenish caterpillars. They sometimes flew
uj) to masses of yellow lichen, where they seemed to be picking out
something to eat. On upper Squaw Creek, August 30, two were seen
eating blueberries (F. A. M.). During hot afternoons the latter part
of July they were often seen soaring and performing aerial antics above
the forest, and also chasing insects in the air, launching out from the
tree tojis after them like flycatchers. As a rule, they are silent when

120

NOKTH AMERICAX FAUNA.

[no. 16.

feeding and noisy when flying about the wliite bark pines. When on
the ground they are very deliberate, and their broad heads and general
form suggest gulls, particularly when the birds are moving away from
the observer.

AVhen searching for insects in the young hemlocks they souietimes
began at the bottom and worked up, sometimes at the top and worked
down. One day in early August a young-of-the-year, showing the true
nutcracker spots on the breast, spent some time in camp, feeding in a
small tree in our midst without showing the least annoyance at our
presence. He began at the top aiul worked slowly downward, drop-
ping from branch to branch and ])eering searchingly over the foliage
and into the tufts of hemlock needles, often lianging almost bottom

Fig. 39. — Clark Crow (Xi(ci/ra[ia cohiwbmna). Drawn by L. A. Fueites.

side up to pick off the small green caterpillars which infested both the
hemlocks and the Shasta firs. We could plainly see him grasp the lit-
tle caterpillars crosswise and give a big gulp in swallowing them, as if
bolting something several times as large. He went over a branch at a
time, examining the whole of it carefully before moving to the one
below, and sometimes went out so far toward the tip that the slender
branch bent down with his weight. Another bird reversed this order
of procedure, and after finally reaching the toji of the tree gave a
jump, aided by a slight flap of the wings, and perched on the very top-
most sprig, when, gaining his balance, he opened his bill and uttered
a little cry of exultation.

Clark crows were almost daily visitors to our camp among the alpine
hemlocks on upper Squaw Creek until near the end of August, when

OCT., 1899.1

BIRDS.

121

. 4U. — Clark Urow (Nucifraya columbiana)
(Photograpbed by Walter K. Fisher.)

they moved up to tiraberline to feed on the large wingless grasshoppers
then abundant along the upper edge of the tongues of dwarf white-
bark pines and on the lava-strewn pumice slopes at still higher eleva-
tions. Some were seen along the edge of the snow at an altitude of
11,000 feet, where dragon-flies, grasshoppers, and other insects were
common.

Clark crow is a little larger than a blue jay, and his colors are put
on in blocks. The body is gray; the wings and tail are black and wliite,
in conspicuous contrast. Still, singular as it may seem, this colora-
tion is both directive and protective.
When in motion the bird is most con
spicuous, the black and wliite i)atches
flashing with great effect; but when
quietly feeding on the ground among
the gray lava rocks of the higher
slopes it is not easily seen, the gray
of the body resembling the gray
rocks, the black markings the dark
shadows. The coloration, however,
is doubtless most protective at night
when the bird is at roost in the trees
and exposed to its worst enemies,
presumably owls and martens. Con-
trasts of gray or white with black are among the most effective of
disappearing colors at night — the black resembling patches of night
shadow, the gray the interspaces.

The true home of the Clark crow is among the white-bark i)ines of
the rocky wind-swept ridges not far from the region of perpetual snow.
Here, from the thaws of early spring till the storms of approaching
winter, not a day passes without his presence. He is a bold, powerful
bird, a lit tenant for such a home, where his loud cry wakes the echoes
of glacier cliffs a thousand times oftener than it reaches a human ear.

69. Cyanocephalus cyanocephalus. Piiion Jay.

Not an inhabitant of Shasta, but occurs in migration about its base,
and may breed in the junipers in Shasta Valley.

September 28, Vernon Bailey saw six in the chaparral and yellow
pines at an altitude of J,000 feet on the wagon road between Elk Creek
and Ash Creek, and the next day found a few near Sheei) Kock. At
Fort Crook, a little southeast of Shasta, a number were collected some
years ago by Captain Feiluer.

70. Xanthocephalus xanthocephalus. Yellow-headed Blackbird.

Not observed by us, but in 1883 C. H. Townsend often saw it "among
the flocks of Brewer blackbirds that fre(j[ueiited the timothy meadows
of Berryvale, at the western base of Mount Shasta, 3,500 feet altitude."
Berryvale is the old name for the meadows near Sisson Tavern.
21753— No. IG ] G

122

NORTH AMERICAN FAUNA.

[NO. 16.

71. Agelaius phoeniceus. lied-winged Blackbird.

Common in Shasta Valley September 17 to 20 (Osgood). C. H. Town-
send gives it as "an abundant summer resident of the cultivated coun-
try north of Mount Shasta."

72. Sturnella magna neglecta. Western Meadowlark.

Common in the meadowland at Sisson and Edgewood, and in the
narrow meadows bordering Shasta IJiver, in Shasta Valley. At Wagon
Camp I saw one in a grassy opening August 13, and a few days earlier
Walter K. Fisher saw one in a similar place a little higher on the
mountain. In the Sisson Meadows ('Strawberry Valley'), during the
first half of September, It. T. Fisher found meadowlarks "gathered in
flocks of as many as a hundred birds."

73. Icterus bullocki. Bullock Oriole.

Common along streams in Shasta and Little Shasta valleys (W. K.
Fisher). Vernon Bailey tells me that at Ager, in the north end of
Shasta Valley, some silver j)oplars in a door yard are literally full of
nests of this oriole, and that when he examined them June 20, 1899, the
new nests contained young orioles and the old nests young house
finches {Carpodacus m. ob.scurus).

74. Scolecophagus cyanocephalus. Brewer Blackbird.

Fairly common at Sisson and in Shasta Valley, but not observed on
the mountain except in one instance, when several were seen at extreme

timberline on the east side of
Mud Creek Canyon August
24 by Vernon Bailey and
Florence A. Merriam.
75. Coccothraustes vespertinus
montanus. Western

Evening Grosbeak.
One of the commonest and
most characteristic birds of
the Shasta fir belt (Canadian
zone), and much less common
in the alpine hemlocks (Hud-
sonian zone). At Wagon
Camp, near the lower border
of the Canadian zone, small
flocks were seen or heard
daily whenever the camp was
occupied, from the time of our
arrival, the middle of July,
until our departure, Septem-
ber 25. At Squaw Creek
Camp,iutheHudsonian zone,
they were much less common, but still by no means rare, and during
early August their distinctive call, a short whistle, was heard every day.

Fig. 41. — Western Eveuinj: Grosbeak {Cuccotlnaustes
vespertinus montanus.) Drawn by L. A. Fuertes.

OCT., 1899.1 BIRDS. 123

In September they were rarely observed on upper Squaw Creek, but
September 18 a flock was seen near the head of Panther Creek. They
were common in INIud Creek Canyon about the end of July and begin-
ning of August. On September 29 I visited the juniper forest in Shasta
Valley, and was surprised to find there dozens of evening grosbeaks,
and still larger numbers of Townsend solitaires, feeding on the fat berries
of the junipers {Juniperus oecidentaJis). At Sisson in September R. T,
Fisher found flocks flying restlessly over the valley.

When in the molt the extraordinary color combinations of the males
surpass even those of the crossbills.

76. Carpodaciis cassini. Cassin Purple Finch.

A common summer resident in the Canadian and lludsouian zones.
Specimens were collected in July and August at various points from
Wagon Camp, in the lower edge of the Shasta flrs, to timberline, at
the upper limit o{ Pinits alhicauUs; and during our circuit of the moun-
tain near timberline July 22 to 25 the species was heard or seen every
day. At our camp in the alpine hendocks on upper Squaw Creek, it
was common and tame, and in early August usually began singing
about daylight. On August 5 I saw one singing in a small white-bark
pine on a ridge east of Mud Creek Canyon at an altitude of 8,800 feet,
far above continuous timberline. A flock, probably migrating, was ob-
served at the head of Panther Creek September 18.

77. Carpodacus mexicanus obscurus (McCall). House Finch.
[= Carpodacus m. frontalis Auct].'

Common at Sissou and in Shasta Valley, but not seen on the moun-
tain, where it is replaced by the preceding species. At Ager Vernon
Bailey found it breeding abundantly in old nests of Bullock orioles.

78. Loxia curvirostra bendirei Eidgway. Sierra Eed Crossbill.

Red crossbills are common on Shasta. Small flocks were heard
nearly every day throughout the season and were fre(|uently seen to
light in the tops of the high flrs, both at Wagon Camp and Squaw
Creek Camp, but they never came into camp to feed, and were rarely
seen near by. One was killed at Sisson September 12 by R. T. Fisher,
who found the species common there at that time, feeding in the tall
pines and also on tlie garden sunflowers. Several killed by us near
timberline lodged in the dense foliage of the alpine hendocks. C. H.
Townsend obtained three specimens on Shasta. The crossbill of Shasta
is intermediate between the northern subspecies minor and the southern
subspecies stricldandi. It is the same as the Fort Klamath form named
by Ridgway as subsi)ecies bendirei; and specimens collected near
Mount Whitney in the southern High Sierra by the Death Valley
expedition, though jjublished as Strickland i,'^ are practically indistin-

I For change of name see Obeiholser, Auk, XVI, p. 186, April, 18i>9.
-North Am. Fauna, No. 7, p. 81, 1893.

124 NORTH AMERICAN FAUNA. [no. 16.

guisliable. It is customary amoug" ornithologists to name recognizable
intergrad.es wliicli have definite and clearly defined breeding ranges.
On this basis bendirei seems entitled to stand, since it occupies the
isolated crest of the Sierra Nevada of California from the southern end
north to Shasta, and pushes on a short distance into southern Oregon
(to Fort Klamath, its type locality). At Diamond Lake, in the south-
ern Cascades, only a few miles north of Fort Klamath, subspecies minor
occurs. The Diamond Lake specimen ' was shot by my assistant, E.
A. Preble, August 13, 1806, and is an adult male. Owing to the late
date and to the well-known erratic habits of crossbills, this bird may
have been a wanderer from farther north. Hence the actual northern
breeding limit of bendirei and the southern breeding limit of minor
remain to be established.

It is a singular fact that two of the specimens obtained by C. H.
Townsend on Lassen, between Shasta and the High Sierra, are as large
and have as large bills as stricldandi. They were killed in summer and
may have been stragglers from some of the mountains to the southeast.

79. Leucosticte tephrocotis (?). Gray-crowned Leucosticte.

At an altitude of 10,000 to 11,000 feet, on the south side of Shasta
Peak, ^'ernon Bailey saw half a dozen leucostictes August 17, and
again the next day. They were feeding among the rocks and on the
glacier which occupies the deep cross gulch just below Konwokitan
glacier. He tells me they ilew down into the crevasses just as we had
seen them do on Mount Kainier the previous year. Since none were
collected there may be some doubt as to the species. The Sierra bird
is ieplirocotis ; the Eainier form Hitorulis.

80. Astragalinus tristis salicamans. Willow Goldfinch.

Xo specimens obtained, but I am sure I saw the species at Sisson in
July and August with .1 . pHultria. Walter K. Fisher reports it as com-
mon in Little Shasta Valley.

81. Astragalinus psaltria. Arkansas Goldfinch.

Common at Sisson and in Shasta Valley; breeds at Sisson.

82. Spinus pinus. Pine Siskin.

Fairly common throughout the Shasta fir forest, and rargiug thence
upward into the alpine hemlocks and white-bark pines. Seen or heard
at frequent intervals from the time of our arrival, the middle of July,
until late in September.

83. Ammodramus sandwichensis alaudinus. Western Savanna Sparrow.
In Shasta Valley W. H. Osgood found this species common Septem-
ber 17 to 20, and just below timberline on Panther Creek I saw several
September 18.

'This specimen, an adult male, agrees with specimens of minor from Wraugel,
Alaska, and Neali Bay and Lapush, Wasliiugton.

OCT., 1899.]

BIRDS.

125

84. Chondestes grammacus strigatus. Western Lark Sparrow.

Not obvserved ou the mountain, but seen at Edge wood August 31,
and in the bushes borilering the narrow meadows on Shasta Kiver, in
the southwestern part of Shasta Valley, September 29. Ten days ear-
lier W. H. Osgood reported them as common a little farther north in
Shasta Valley. At Sisson B,. T. Fisher saw two flocks the first half of
September. In August, 1883, C. H. Townseud found the species abun-
dant "on the high-lying plains northward of Mount Shasta." In July,
1899, Walter K. Fisher reported it as the commonest bird in Shasta
Valley.

85. Zonotrichia leucophrys. White-crowned Sparrow.

Probably breeds near timberline in the Hudsonian zone, where, near
the head of Mud Creek, one
was shot August 4 by W. H.
Osgood. About the middle of
September numbers were seen
just below timberline ou Pan-
ther C r e e k — doubtless mi-
grants— and a little later they
were common in the bushes
from Wagon Camp to Sisson
and along Shasta River in
Shasta Valley.

86. Spizella socialis arizonse.
Western Chipping Spar-
row.

As tlie chipping sparrow is
not an inhabitant of dense I'or-
ests it was not seen in the
Shasta fir belt; but in the open-
ings along the lower edge of
this belt it breeds, and was
feediugyoungat Wagon Camp
in July. After the breeding season it was seen from time to time near
timberline. It is a common breeder at Sisson and in Shasta Valley.

87. Junco hyemalis thurberi. Sierra Junco.

Common from timberline down to the lower edge of the Shasta firs.
The commonest 'ground bird' of the forest. Several nests with eggs or
young were found near Wagon Camp the latter part of July.

88. Melospiza melodia montana. Mountain Song Sparrow.

Song sparrows supposed to be this subspecies were found breeding
at Wagon Camp and Sisson the middle of July. Others were seen in
the tules at Big Spring, in Shasta Valley, September 17 to 20 by W. H.
Osgood. One was secured at Edgewood July 19, 1899, by Walter K.
Fisher, who states that it was the only one seen during his stay.

-White-crowned Sparrow (Zonotrichia
phrys). Drawn by L. A. Fuertes.

126 NORTH AMERICAN FAUNA. [no.16.

89. Melospiza lincolni. Lincoln Sparrow.

Breeds alonji' the edfje of the forest at Wagou Camp, where I shot
cue at the spring July 1^*0. Another was obtained by 11. T. Fisher on
Squaw Creek (alt. about 0,700 feet) August 17. Others were seen
among the alders in Mud and Ash Creek canyons about the end of July
by Walter K. Fisher. At Sisson, the first half of September, II. T.
Fisher found the species fairly common, and shot one among the alders
bordering Cold Creek. W. H. Osgood reported it as common in Shasta
Valley September 17 to 20.

90. Passerella iliaca unalaschcensis. Townsend Sparrow,

Common during nugration (September) throughout the chaparral of
the lower slopes from Sisson up to Wagon Camp. At Wagon Camp
one was shot by W. H. Osgood September 22. Common also late in
September in bushes in the more open parts of the forest along the east
base of Scott Mountains from Sisson north to Edgewood, and in the
bushes bordering Shasta Eiver in the southern part of Shasta Valley.

Passerellas supposed to be this species were seen repeatedly at
Sisson, the tirst half of September, by R. T. Fisher.

91. Passerella iliaca megarhyncha. Thick-billed Sparrow.

Breeds near Wagon Camp, apparently in the chaparral, where Jolin
H. Sage shot one July 20 and another July 22. At Sisson W, II.
Osgood and Miss Merriam saw this species July 13 and 15. In the
summer of 1883 C. H. Townsend found it a common breeder in the
chaparral tracts of Shasta.

92. Pipilo maculatus megalonyx. Spurred Towhee.

Common in the undergrowth along the edges of openings in the i)on-
derosa pine forest from Sisson to Shasta Valley. One was shot at
Upton September 15 by R. T, Fisher. C. H. Townsend found it breed-
ing commonly on the lower slopes of Shasta in 1883.

93. Pipilo fuscus crissalis. California Towhee.

Several seen in the chaparral near Gazelle August 31 by Vernon
Bailey, and near Edgewood by Walter K. Fisher.

94. Oreospiza chlorura. Green-tailed Towhee.

Apparently a fairly common breeder on the lower slopes of Shasta,
where in the chaparral near Wagon Camp several were seen and one
was killed the latter part of July and early in August, and one was
found feeding full-grown young July 28. At Sisson Miss Merriam
found them the middle of July. The last one noted by me was seen
in the manzanita at an altitude of 4,900 feet, on the road from Wagon
Camp to Sisson, August 20.

95. Zamelodia melanocephala. Black-headed Grosbeak.

Seen at Sisson by W. H. Osgood July 13. At the McCloud River
fish hatchery C. H. Townsend found it a common breeder in 1S83.

OCT., 1899.] BIRDS. 127

IKi. Cyanospiza amoena. Lazuli Buuting-.

Common at Sissoii (where it was seen feeding young July 15 by
Miss Merriam), and ranging thence up through the chaparral of the
Transition zone all the way to Wagon Camp^ where a nest was found
and the parent secured July 17 by W. H. Osgood, who obtained
another in Mud Creek Canyon August 3.

97. Piranga ludoviciana. Mountain Tanager.

One of the commonest, as well as handsomest, birds of the Shasta
lir forest. At Wagon Camp, at the lower edge of the Canadian zone,
these tanagers were seen every day from the middle of July until
August 10, when that camp was temporarily abandoned. They were
very tame, and spent much time in camp, drinking from our little spring
and peering at us from the lower branches of the firs. On July 21 I
saw several picking caterpillars from the leaves of tlie buck brush
{Ceanothus velHiinuH) in an opening in the firs. Walter K. Fisher
states that one; sang continuously at his camp in Mud Creek Canyon
at the mouth of Clear Creek about the end of July. At Sisson this
tanager was seen July 15 by Miss Merriam.

98. Progne subis hesperia. Western Martin.

Not observed by us, but recorded by C. H. Townsend, who, in 1883,
noticed a few "about some buildings at the west base of Mount Shasta
in midsummer." One seen about G miles northwest of Edgewood by
Walter K. Fisher.

99. Petrochelidon lunifrons. Clifif Swallow.

Common at Sisson, where it was noted by Miss Merriam the middle
of July, and by E. T. Fisher the first half of September. Abundant in
Shasta Valley (W. K. Fisher).

100. Hirundo erythrogastra. Barn Swallow.

Common at Sisson, and seen in Shasta Valley September 17 to 20 by
W. H. Osgood. Common in Shasta A' alley in July (W. K. Fisher).

101. Tachycineta thalassina. Violet-green Swallow.

Common at Sisson, and seen once or twice tlying over the chaparral,
nearly up to Wagon Camp.

102. Tachycineta bicolor. Tree Swallow: White-bellied Swallow.
Swallows believed to be this species were seen several times between

Wagon Camp and Sisson, and C. H. Townsend records them as abun-
dant breeders on the lower McCloud.

103. Stelgidopteryx serripennis. Rough-winged Swallow.

Walter K. Fisher saw some swallows at Edgewood in July, 1899,
which he believed to be this species.

128 NORTH AMERICAN FAUNA. [no. 16.

104. Lanius ludovicianus excubitorides. White-riiniped Shrike.

One seeu at Gazelle, on the west side of Shasta Valley, August 31,
by Vernon Bailey, and one in Shasta Valley September 18 by W. H.
Osgood. Rather common in Shasta Valley in July, 181)9 (W. K. Fisher).

105. Vireo solitarius cassini. Cassin Vireo.

Fairly common along the lower border of the Shasta fir forest at
Wagon Camp, where two or three were obtained the latter part of July,
and where a nest containing one egg and one young was found by John
H. Sage July M. The nest was in a clump of buck-brush [Ceanothiis
velut'inus) ouly 3 feet above the ground, in open chaparral, near the edge
of the Shasta fir woods.

106. Vireo gilvus swainsoni. Western Warbling Vireo.

Recorded by C. H. Townsend as "found at rare intervals in midsum-
mer about the base of Shasta." A vireo heard by us several times at
Sisson was probably this species. Walter K. Fisher found it at Edge-
wood, among the bushes along Shasta River, where he secured a
sijecimen July 19, 1899.

107. HelminthopMla rubricapilla gutturalis. Calaveras Warbler.

Common in the chaparral at Wagon Camp, where several specimens
were collected the last half of July. Later the species was seen with
other warblers in the trees higher \\\) on the mountain.

108. Helminthophila celata lutescens, Lutescent Warbler.

Fairly common and apparently breeding at Wagon Camp, in the
lower edge of the Shasta firs, where two were collected July 20. One
was shot in the alpine hemlocks on Squaw Creek August 6 by Vernon
Bailey.

109. Dendroica sestiva. Yellow Warbler.

Seen at Sisson, the middle of July, by Miss Merriam. The only one
seen on the mountain was shot in the chaparral near the edge of the fir
forest at Wagon Camp July 29 by John H. Sage. Common in bushes
along Shasta River in Shasta Valley (W. K. Fisher).

110. Dendroica auduboni. Audubon Warbler.

Abundant throughout the forest region of Shasta, from timberline
down to the lower edge of the Canadian zone. At Wagon Camp Miss
INIerriam saw them feeding fullgrown young July 18 and August 1. In
August and September they were seen daily at our camp in the alpine
hemlocks, often in mixed flocks of chickadees and other birds. When
we left Wagon Camp, September 25, they were still common there.

BIRDS.

129

August 9 and 10, spotted young were collected, one with the brown
sides coming in and one with the yellow just beginning to show. At

Fig. 43.— Audubon Warbler (Dendi-oica aiiduboni). Drawn by L. A. Fuertes.

Sisson, Miss Merriam saw them the middle of July, and R. T. Fisher
found them common the first half of September.

111. Dendroica occidentalis. Hermit Warbler.

FBjirly common in the Hudsonian and upper part of the Canadian
forest during August (collected at Sfiuaw Creek Camp at intervals from
August 3 to August 28)5 i^ot seen in July.

112. Geothlypis tolmiei. Macgillivray Warbler.

Seen at Sisson and Wagon Camp about the middle of July, and again
at the latter locality August 1 and August 3. One was shot in Mud
Creek Canyon at an altitude of G,7(>0 feet by W. H. Osgood August 7.
In July and August, 1883, C. H. Townsend found it not uncommon on
the mountain.

113. Geothlypis trichas occidentalis. Western Yellow-throat.

At Big Spriug, in Shasta Valley, W. H. Osgood found this warbler
common in the tules September 17 to 20; and August G, 1883, C. H.
Townsend secured an immature specimen at the base of Mount Shasta.
21753— No. 10 17

130 NORTH AMERICAN FAUNA. [no. 16.

114. Icteria virens longicauda. Long-tailed Chat.

Fairly comnion at Sisson, and common in the lower country farther
north — at Gazelle and on toward Yreka. Common along Shasta River
in Shasta Valley (W. K. Fisher).

115. Wilsonia pusilla pileolata. Pileolated Warbler.

Common in Mud Creek Canyon the latter part of July and early
August; seen at Sisson July 13 by W. H. Osgood, and at Wagon Camp
August 3 by Miss Merriam; several seen at Squaw Creek in flocks of
chickadees and warblers during August and September; seen in wild
cherry bushes at Wagon Camp September 25. In the summer of 1883
C. H. Townsend found it rather common on Shasta.

116. Anthus pensilvanicus. Pipit; Titlark.

In a barren rocky basin above timberline, near the head of Panther
Creek, on July 17 I heard titlarks and saw Arctic bluebirds. At the
same place two months later (September 18) titlarks were common.
In Shasta Valley September 17 to 120 W. H. Osgood saw flocks along
the road.

117. Cinclus mexicanus. Dijiper ; Water Ouzel.

Common on the upper jiart of Squaw Creek as far as the timber
extends, and sometimes seen above timberline. Seen also on Panther
Creek, near its head, and on Mud and Ash creeks. On Squaw Creek
we often watched the ouzels feeding among the cascades and clear
cold pools between the upper heather meadow and the main fall. One
afternoon just before dark (I'l o'clock) I was sur[)rised to see an ouzel
fly up into the dead top of a tree, light on a branch, and climb up
several feet on the trunk with his short tail hanging straight down,
after the manner of a woodpecker.

118. Salpinctes obsoletus. Rock Wren.

Rather rare on Shasta and confined apparently to the bare rocky
slopes near and above timberline, August 2, at an altitude of 8,000
feet, on one of the ridges above the head of Squiiw Creek, I found a
pair — the first noted on the mountain. August 11, on the west rim of
Mud Creek Canyon, I first heard them at 9,000 feet, and Vernon Bailey
followed them up and shot one at 9,800 feet. A week later he saw one
at an altitude of 10,400 feet. August 27, Miss Merriam saw one in
'The Gate,' between the heads of Panther Creek and Squaw Creek. In
1883 C. H. Townsend found them common above timberline on Shasta,
and observed them also on Sheep Rock. In July, 1899, Walter K.
Fisher found them among lava ledges in Shasta Valley.

119. Catherpes mexicanus punctulatus. Canyon Wren.

Not observed by us, but recorded by Townsend, who in 1883 saw
one — doubtless a straggler — above timberline on Shasta, and others at
Sheep Rock, at the northeast base of the mountain, where he heard
the bird's wonderful song and was deeply impressed by it.

OCT., 1809.] BIRDS. 131

120. Thryomanes bewicki spilurus. Vigors Wren.

Seen at Sisson, bnt not on the mountain. Kecorded by C. H. Town-
send from "the base of Shasta."

121. Troglodytes aedon parkmani. Parkman Wren.

Rare on tlie mountain. One collected near Squaw Creek Camp (alt.
G,750 feet) August 18 by R. T. Fisher, and one in Mud Creek Canyon
(alt. 6,780 feet) August 9 by W. H. Osgood. More common at Sisson,
where it was observed by Miss Merriam the middle of July. C. H.
Townseiid records it as abundant in midsummer " among the piled-up
logs of a certain clearing in the forest at the base of Mount Shasta."
The bird seems to be intermediate between parlinanl and aztecus.

122. Cistothorus palustris paludicola. Tule Wren.

At Big Spring, in Shasta Valley, W. II, Osgood found these wrens
common September 17 to 20.

[The western winter wren {Anorthura hiemalis imcific<<) was not
observed by us, but was found byC. H. Townseud in the mossy canyons
of the lower McCloud. It doubtless occurs in the upper canyon of the
Sacramento near Sisson, and quite possibly in other suitable places
about the mountain.]

123. Certhia familiaris occidentalis. Western Brown Creeper.

Not common, but seen now and then in the alpine hemlocks on Squaw
Creek and in the Shasta firs, lower down. The earliest record is July
24 (J. H. S.); the latest. September 15 (C. H. M.). At Wagon Camp
John H. Sage shot one and saw another July 27; and Miss Merriam
saw one in the same place August 1, one at Sijuaw Creek August 30,
and two at Sisson September 3.

124. Sitta carolinensis aculeata. Slender-billed Nuthatch.

Not common on the mountain, and only moderately so at Sisson. A
few were seen at Squaw Creek Canq) from time to time in August, once
or twice in the mixed flocks of chickadees and warblers.

125. Sitta canadensis. Red-breasted Nuthatch.

Common from the lower edge of the Shasta fir belt up to timberline;
and from the time of our arrival, the middle of July, until our depart-
ure, the latter part of September. In September it was often seen in
mixed flocks with m<mntain chickadees and Audubon warblers.

120. Sitta pygmaea. Pigmy Nuthatch.

In the ponderosa pines near Sisson, W. H. Osgood saw pigmy nut-
hatches July 13, and I saw a small flock September 30. At the south-
west end of Shasta Valley, not far from Edgewood, Osgood saw several
in the pines September 20.

132 NORTH AMERICAN FAUNA. Ino.16.

127. Parus gambeli. Mountain Chickadee.

Common and widely distributed. In the firs at Wagon Camp they
were among the commonest birds, and at higher altitudes were seen or
heard nearly evoy day. Early on the morning of July 24, after camp-
ing for the night in the shelter of a narrow fringe of dwarf white-bark
pines at timberline, on one of the desolate torrent-swept beds of Incon-
stance Creek, high up on the north side of Shasta, we were saluted by
a small tiock of these cheerful little birds. In September they helped
form the mixed tlocks, along with Audubon and other warblers, Canada
nuthatches, and other small fry so often seen among the Shasta firs and
alpine hemlocks. At Sisson Miss Merriam saw them about the mid-
dle of July; they were common there in September, and were noted in
Shasta Valley by W. H. Osgood September 17 to 20.

128. Parus rufescens. Chestnut- backed Chickadee.

Not observed by us, but recorded by C. H, Townseud, who " obtained
a single individual at the western base of Mount Shasta on July 14,

1883."

129. Psaltriparus minimus californicus. California Bush-Tit.
Common along Little Shasta Creek Sei)tember 18 (VV. H. Osgood).

Seen in the chaparral west of Gazelle, on the west side of Shasta Val-
ley, August 31 (Vernon Bailey). Townsend found it a common breeder
near the fish hatchery on the Lower McCIoud River.

130. Regulus satrapa olivaceus. Western Golden-crowned Kinglet.
Both kinglets are common on Shasta and doubtless breed in the

Shasta firs of the Canadian zone. The golden-crown was often seen
and heard at Wagon Cam]), where it was collected July 18. It was
also common just within the lower edge of the fir forest in Mud Creek
Canyon July 23, and a couple of miles higher up about the end of the
month. O. H. Townsend obtained specimens at timberline in July, 1883.

131. Regulus calendula. Ruby- crowned Kinglet.

The ruby-crown was frequently heard by various members of the
expedition, and at different altitudes, from Wagon Camp in the lower
edge of the Shasta firs, to timberline, where one was collected by W. H.
Osgood August 4. In 1883 C. H. Townsend obtained two specimens,
one at timberline August 15, the other at an elevation of about (!,0()0
feet September 2.

132. Myadestes townsendi. Townsend Solitaire.

Not an uncommon breeder on the higher slopes of Shasta, where I
saw SIX during our stay. The lowest of these was at Wagon Camp, in
the lower edge of (he Shasta firs, where one drank at our spring July 31.
The others were in the Hudsonian zone, and most of them in the alpine
hemlocks at or near our camp on Squaw Creek, Just below timberline.

OCT., 1899. J BIRDS. 133

Tbey were usually seen in early evening when they came to drink from
the little streams in the forest, and were always silent and rather shy.
Late in July and early in August they were seen with arctic bluebirds
by Walter K. Fisher at and below timberline near Mud Creek Canyon.
They probably winter in the junipers in Shasta Valley, where W. H.
Osgood saw them September 17 to 20, and where I saw dozens feeding
on the juniper berries and singing freely September 139. Late in July,
1883, C. H. Townsend found one of these birds "frozen in the snow and
ice which filled the crater of the extinct volcano of Shasta."

133. Hylocichla aonalaschkae auduboni. Dwarf Hermit Thrush,
Evidently breeds in the dark Shasta fir forest of the mountain and

in the damj) forest of white and Douglas firs near Sisson Tavern, where
Miss Merriam heard them in full song the middle of July. At Wagon
Camp W. H. Osgood shot one July 17; and near Squaw Creek Camp
I shot another September '20. C. H. Townsend records it from the
southern slope of Shasta under date of July 25.

134. Merula migratoria propinqua. Western Robin.

Kobins were fairly common at Wagon Camp, where we heard their
evening song the day of our arrival, July 15, and where they were seen
carrying food to young July 26 (F. A. M.). Higher up they were seen
from time to time, but were not common. Along the upper part of
Mud Creek Canyon Walter K. Fisher found them "fairly common but
erratic" the latter part of July and tirst few days of August. At Squaw
Creek Camp, in the alpine hemlocks, we saw a flock on the morning
of August 12, and others September 15. Just below timberline on
Panther Creek a few were seen migrating September 18. In Shasta
Valley they were common among the junipers September 20. At Sis-
son they are common breeders, and a few were seen whenever any of
our party were there, from July 14 until the end of September. A nest
containing three eggs was found July 13 by W. H. Osgood.

135. Sialia mexicana occidentalis. Western Bluebird.

A common breeder at Sisson, where families, including lately fledged
yonng, haunted the fences the first half of September (R. T. Fisher).
In Shasta Valley W. H. Osgood found them common September 17 to
20, and a few small flocks were seen by me among the junipers Septem-
ber 29. At Sisson the species was common the latter part of Septem-
ber. In 1883 a nest containing nearly grown young was found by C.
H. Townsend in a post hole in the ground at Sisson Tavern late in
July.

13(5. Sialia arctica. Mountain Bluebird; Arctic Bluebird.

A common breeder on the higher slopes of Shasta, where they were
constantly seen in July and early August, but rarely afterwards. On
our first visit to timberline, July 17, they were very abundant on the

134 NORTH AMERICAN FAUNA. [no. 16.

bare rocks above timberline near the head of Pauther Greek, and a
week later were seen on the rim of Mud Creek Canyon. On the rocky
slopes at and above timberline on the east side of Mud Creek Canyon
early in August they were the most characteristic birds ( W. K. Fisher) ;
and they were seen near the same place August 18 (V. Bailey). Sep-
tember 20 a small flock visited our camp on upper Squaw Creek.

In July, 1883, C. H. Townseud found the species abundant at timber-
line, where full fledged young accompanied their parents.

NOTES ON THE DISTRIBUTION OF SHASTA PLANTS.

The study of the geographic distribution of animals and [)hints in the
Sierra-Cascades can not be completed until authentic lists of species
are brought together from several important localities from which at
present no data are available. The need of such lists, with detailed
altitudes and zone positions, is particularly urgent in the case of
detached mountains, as Shasta and Lassen, which are separated from
each other and from the continuous ranges on either side by gaps
low enough to be broadly filled by Transition zone species. The Boreal
species of these mountains, being thus completely cut off from the
nearest corresponding colonies, form islands in the long Boreal chain
that stretches southward from British Columbia to southern California.

The present imperfect list of the plants of Mount Shasta is offered
as an humble contribution toward the needed material. It is based
almost wholly on my own personal observations and is known to be
far from complete. More pressing work along other lines made it
impracticable to give nuich time to plants, and the date of arrival at
the mountain (the nnddle of July) was so late that many of the early
flowering species had disappeared. The Boreal species, owing to their
greater importance, have received most attention ; the Transition zone
species least. In the case of Alpine and Hudsonian species it is believed
that few remain to be added.

I am indebted to Miss Lewanna Wilkins for collecting and pressing
most of the plants preserved during the first six weeks of our stay on
the mountain; and to John H. Sage for the use of a collection made
by him during the same period. The plants obtained subsequent to
August 8 were collected by Vernon Bailey and myself

Although two seasons' field work in the Cascade Bauge had given
me a certain acquaintance with the more conspicuous jDlants common
to these mountains and Mount Shasta, I was still much handicapped
in the determination of the species, and not being a botanist myself
was obliged to appeal to professional botanists for assistance. I am
indebted most of all to Miss Alice Eastwood, curator of the herba-
rium of the California Academy of Sciences, whose courtesy and
promptness in identifying plants, sent her from time to time while I
was still in the field, were of the utmost assistance. I am also particu-
larly indebted to Prof. E. L. (ireene, of the Catholic University at
Washington, who has taken the trouble to examine a large number of
species, and to describe several which proved to be new. Other bota-

135

136 NORTH AMERICAN FAUNA. [no. i6.

nists who kindly determiued special species or special genera are Mrs.
N. L. I5ritton, Mr. Frederick V. Coville, Prof. W. U. Dudley, Mr. M. L.
Fernald, Mr. John B. Leiberg, Mr. C. L. Pollard, Dr. B. L. Robinson,
Dr. J. IS^. Rose, Prof. P. A. Rydberg, Mr. J. K. Small, and Prof. Wil-
liam Trelease. In order to properly place the credit for this assistance,
the botanist responsible for the identification is in each instance men-
tioned. Where no one is named I am personally responsible.

A few mosses and ferns were found in the timberline region of
Shasta. The mosses, kindly determined by Mrs. Britton, are Har-
pidium (? t'xamiulafum Gumb.), Aulacomnium andro//i/nitm Sell., and a
species of PhilonoUs. They are contined rather closely to the borders
of the alpine streams, where the Aulacomnium forms compact mats of
a yellowish color.

The ferns have been determined by Mr. William R. Mason through
the courtesy of Mr. Frederick V. Coville, curator of the National Her-
barium. The species which grow sparingly "u the heather meadows and
under the edges of rocks near timberline are : Dryoptcris aculeata scopn-
Una (Eaton), Cystopteris fragiUs (L.) Bernh., Cheilanthes gracilUma
D. C. Eaton, Cryptogramme acrosUchoides R. Br., and Fliegopteris alpe^i-
tris (Iloppe) Mett. Much lower down, along the boundary between
the Canadian and Transition zones. Tier is aquilina lanuginosa, (Bory)
Hooker, and Asplenium filixf(emina (L.) Bernh. occur. The brake
{Pteris) is very abundant on the pumice sand at AVagon Camp and
Sisson and wherever there is sufficient moisture in the soil in spring
and early summer. The black-beard lichen {Alecioriafremonti) and the
handsome yellow tree lichen [Erernia vulpina) abound in the dark forest
of Shasta firs.

In the chapter on 'Life Zones' the more distinctive species have
been groui)ed according to their vertical distribution (see pp. 52-68), but
for convenience in finding the notes relating to particular species,
tliey are here arranged in systematic order. In the case of certain
plants found by us only on the borderland between adjacent zones, the
zone position is in doubt and nuist be determined by future study.

The most important kinds of plants from the standpoint of geo-
graphic distribution are naturally those that remain longest in a par-
ticular spot. Hence, as pointed out by Coville,' trees, shrubs, and
perennials are the kinds most useful in determining zone boundaries.
For this reason little attention is here given to annuals.

Pinus monticola Douglas. Silver Pine; Mountain White Pine.

Common in places, chiefly in the upper half of the Canadian zone,
but local and by no means generally distributed. Wherever it occurs
it is mixed with Shasta firs, and in places it reaches up high enough to
overlap the lower edge of the black alpine hemlocks and white-bark
timberline pines. (See ]>. 3.S.)

Botany of Death Valley Expedition, pp. 17-18, 1893.

OCT., 1899.] PLANTS. 137

Pinus lambertiana Douglas. Sugar Pine.

Fairly common in most parts of the Transition zone forest of ])onderosa
pines, and occurring here and there in the iiiiniense areas of chaparral
that cover the lower sh)pes on the south and west sides of the mountain.
(See pp. 32-3:5.)

Pinus albicaulis Engelmann. White-bark Pine.

The timberline tree of Shasta, which it encircles at altitudes ranging,
according to slope, from about 7,0))Q up to 8,000 feet, and i)ushing up
on the warmest ridges to an extreme elevation of 9,800 feet. In its
distribution therefore it tills the Hudsonian zone except in places
uusuited to tree growth. The only tree competing witli it on Shasta
is the black alpine hemlock, which, requiring more moisture, is at a
disadvantage and is confined to special localities, as exj)lained in full
under that species. (See pp. 39-42.)

Pinus ponderosa Laws. Pomlerosa or Yellow Pine.

The most abundant and characteristic tree of the lower slopes and
basal i)lane of Shasta, where, tilling the Transition zone, it forms a con-
tinuous o[)eii forest of wide extent. (See pp. 30-32.)

Pinus murrayana Balf.iur. Lodge-pole Pine.

Confined to the northeast quadrant of Shasta, where it occupies the
lower part of the Canadian zone. (See pp. 38-39.)

Pinus attenuata Lemmon. Knobcone Pine.

Common in a limited area in the Transition zone on the south side of
Shasta, where it is confined to the lower slopes (from about 4,000 to
5,000 feet altitude) from Panther Creek easterly to between the
branches of Mud Creek. (See pp. 33-34.)

Tsuga mertensiana (Bong.) Carr.' Black Alpine Hemlock.

A characteristic tree of the Hudsonian zone, where, however, it is
not generally distributed for the reason that the upper slopes of Shasta
are in most places too dry for it. It occurs in the same belt with Pinus
albicaulis, but does not reach so high, and, requiring more moisture, is
confined to disconnected localities, usually in canyons and gulches or
along the shady sides of buttes or ridges. (See pp. 42-40.)

Pseudotsuga mucronata (Raf.). Sudw. Douglas Fir or Spruce.

Common, scattered through the, less arid parts of the forest of pon
derosa pines which clothes the lower Transition zone slopes of Shasta
and extends away in all directions (see p. 32). A subspecies pendula
(Engelm.) Sud worth, with "branches, at least the lower ones, very
slender and long-pendent," has been descinbed from Sisson (Bot. Calif,
II, 483, 1880; Sud worth. Check List Forest Trees of U. S., 24, 1898),
and is common thence westerly to the coast. (See pp. 34-35).

' For change of name from Tsitgajxitloni to T. inertinsiana, see footnote p. 42.
21753— No. 16 18

138 NORTH AMKRICAN FAUNA. [no. 16.

Abies shastensis Leiiunoii. Shasta Fir.

The dominaut tree of the Canadian zone, covering the middle slojjes
and completely encircling the mountain in a solid belt about 2,000 feet
in vertical breadth. Its upper border overlaps the lower edge of the
Hudsonian ; its lower border, the up]>er edge of the Transition. Along
Panther and Squaw creeks, on tlie south slope, it ranges from about
0,500 up to 7,500 feet, and on steep southwesterly slopes considerably
higher. (See pp. 36-38.)

Abies concolor lowiana (Murray) Lemmon. White Fir.

The common and only true iir of the basal slopes of Shasta, where
it occurs in moist places from the altitude of Wagon Camp (5,700 feet)
down to the very bottom of Sissou Valley at the base of Mount Eddy
(alt. 3,400 feet). It thus fills the Transition zone, except in places that
are too dry for it. (See p. 34.)

Libocedrus decurrens Torrey. Incense Cedar.

Common throughout the Transition zone forest of ponderosa pines,
except in the dryest places. (See p. 35.)

Chamaecyparis lawsoniana (Murray) Parlat. Lawson Cypress.

Attributed to "the Shasta Mountains^' (Bot. Calif , II, p. 115, 1880),
but not found by us.'

Juniperus nana Willd. Dwarf Aljjine Juni])er.

Fairly common in i)laces close to timberline, usually associated with
Pinns aJbicaulu in the upper i)art of the Hudsonian zone. On Shasta
it usually grows in small patches less than a foot high and, as a rule,
only a few feet in diameterj in the Olympics, on Mount Eainier, and on
numerous other mountains it forms much larger patches. Between
Mud Creek Canyon and the high ridges above Squaw Creek, it pushes
up to extreme timberline at 9,800 feet, along with Pinus alhicaulis.
On the north side of Shasta and Shastina it was found in crevices
among the sharp lava rocks at altitudes varying from 8,.'>00 to 9,000
feet, and was common on the curious (ilhicauli.s plain stretching west-
erly from 'North Gate' to Bolam Creek.

Juniperus occidentalis Hooker. Western rluniper.

Abundant in the south end of Shasta Valley, where it forms a forest
many miles in extent. Scattered trees begin 4 or 4i miles easterly from
Edgewood, and become more and m(jre plentiful to the north until, at
a distance of about 5 miles south from Big Spring, they suddenly
become abundant and cover the whole country east of the main mass
of lava buttes, forming a continuous Juniper forest which fills the south-
ern r)art of Shasta Valley and r-eaches northward, I am told, into Little

'Another cypress attributed to Shasta but not seen by us is Cupressus macnabiana
Murr., " originally reported by Jeffrey from Mount Shasta at 5,000 feet altitude"
(Bot. Calif., II, p. Ill, 1880). In both iustauces, i)robably, the term 'Shasta' was used
in a rather loose sense, as foveriug adjacent niouutaius uot then named.

OCT., 1899.J PLANTS. 139

Shasta Valley. When visited about the end of September, the trees
were full of their large berries, and many birds, including evening-
grosbeaks and Townsend solitaires, were there in numbers feeding
on them. The zone position of this tree, which appears to be the
type form, is high Upper Sonoran and low Transition. The boreal
form common in the Sierra, but not found on Shasta, should be
diiferent.

Sitanion cinereum J. G. Smith. Alpine Grass.

The common grass of the glacial meadows, but nowhere sufficiently
abundant to form anything like a sod. It is closely related to S. ely-
moidcs llaf., from which it has been recently separated by Mr. Jared G.
Smith.

Carex breweri Boott. Alpine Carex.

Common in the glacial meadows and scattered sparingly over the
moist slopes. (Identified by F. V. Coville.)

Juncus parryi Eugelm. Parry Juiicus.

Common in the heather patches and other moist places from slightly
below timber] ine up through the Alpine zone. Xoted by Vernon Bailey
as high as 11,300 feet. (Identified by F. V. Coville.)

Allium validum Watson. Large Wild Onion.

Abundant in the Canadian zone along the streams of the Shasta fir
belt, and often growing in the lowei- heather beds along the lower
border of the Hudsoniau zone. (Identified by Professor Greene.)

Allmm sp. — !

A very small species is common in the glacial meadows at the head
of Squaw Creek, but matures so early that we were uiuible to obtain
anything but the bulbs.

Calochortus nudus Watson.

Fairly common along the lower edge of the Canadian zone neap
Wagon Camp, flowering late in July. (Identified by Miss Eastwood
and Professor Greene.)

Calochortus maweanus Leichtlin.

Not rare at Wagon Camp, but not seen elsewhere. An elegant little
species with white hair}' flowers. (Identified by Professor Greene.)
Fritillaria atropurpurea Nutt.

Common in the chaparral of the Transition zone basal slopes from
Sisson up to Wagon Camp. (Identified by Miss Eastwood.)
Hastingsia alba (Durand) Wats.

Common near Wagon Camp, where its long and slender cylindrical
spikes were in flower the latter ])art of July. (Identified by Miss East-
wood and Professor Greene.)
Lilium parvum Kellogg. Tiger Lily.

Common in marshy places in the lower part of the Shasta fir belt,
^particularly at Wagon Camp, where it was flowering abundantly about

14(1

NORTH AMERICAN FAUNA.

[NO. 16.

the middle of July, and was still in fruit sis late as tlie end of September.
(Identified by Miss Eastwood.)

Lilium washingtonianum Kellogg. Washington Lily.

This superb lily, with large white and very fragrant flowers, is com-
mon in the manzanita in the upper part of the Transition zone, and
occurs sparingly at lower altitudes. At Sisson it was in fruit the end
of September. The flowers had passed their prime by the middle of
July, but occasional i>lants were found in blossom as late as the end
of the month.

Tofieldia occidentalis Watson.

Common near the bottom of the Canadian zone at Wagon Camp.
(Identified by Professor Greene.)

Tritelia ixioides (Ait.) Greene {-^Brodkva).

Common at Wagon Camp on the border between the Canadian and
Transition zones, where its yellow star flowers were in bloom the latter
half of July. (Identified by Professor Greene.)

Vagnera stellata (Linn.) Morong.

Common at Wagon Camp, particularly in open grassy places along
the edges of the fir forest, where it was flowering abundantly when
we reached the mountain, the middle of July. Plants still holding

their fruit were ob-
served when we left
Wagon Camj), Septem-
ber 25.

Veratrum californicum
Dnrand. White
Hellebore.
Abundant at numer-
ous localities a long
the streams and in
marshy spots in the
Canadian and upper
part of the Transition
zones. Particularly
common at Wagon
Camp, and also on
Squaw Creek Just
above the middle
meadow.

Fig. 44.— White Hellebore (Veratrum californicum).
Photographed by W. K. Fisher.

Smilax californica Gray. California Smilax.

Occurs in places in the lower part of the Transition zone along Shasta
River between Edgewood and Sisson, but nearer Sisson. Not observed
elsewhere.

OCT.. 1899.] PLANTS. 141

Sisyrinchium bellum Watson. Blue-eyed Grass.

Fairly conuuon in the Transition zone near Wagon Camp.
Corallorhiza bigelovi Watson.

Xot uncommon in the woods near Wagon Camp. (Identifled by Pro-
fessor Greene.)
Habenaria leucostachys (Lindl.) Watson.

Common in the marsh at Wagon Camp. (Identiued by Professor
Greene.)

Habenaria unalaschensis Watson.

A boreal species, fairly common in the marsh near Wagon Camp,
growing with the last. (Identified by Professor Greene.)
Populus tricbocarpa T. .S: G. Western Balsam Poplar.

Common in the upper Sacramento Canyon near Sisson, and less so
along Shasta River at the south end of Shasta Valley.
Salix lasiandra Bentham. Black Willow.

Abundant in cool moist jdaces about Sisson. (Identified by Miss
Eastwood.)

Salix nuttalli Sargent. Xuttall Willow.

Common in moist places in canyons of the Canadian zone and near
Wagon Camp. (Identified by F. V. Coville.)
Salix sitchensis Sanson. Sitka Willow.

Common in canyons in the Canadian zone. Found in Mud Creek
Canyon near the mouth of Clear Creek. (Identified by F. Y. Coville.)
Alnus rhombifolia Xutt. White Tree Alder.

Observed only on Shasta River in the southern part of Shasta
Valley, where its zone position appears to be Upper Sonoran, Shasta
Valley is one belt lower than the rest of the region about Shasta and
contains a dilute tongue of Upper Sonoran species that come in from
the north by way of Klamath River Valley.
Alnus sinuata (Regel) Rydb. Alder.

Common in moist i>laces in the canyons of the Canadian zone. In
Mud Creek Canyon noted as high as 6,700 feet. Found also near
Wagon Camp. (Identified by F. V. Coville.)
Alnus tenuifolia Xutt. Paperleaf Alder.

[ = A. incana var. rirescens Wats].

A Transition zone species common along streams in the neighbor-
hood of Sisson Tavern and along the east base of Scott jMountains,
(Identified by P. V. Coville.)
Betula occidentalis Hooker. Birch.

Fairly common along Shasta River at the south end of Shasta
Valley. Not observed elsewhere.
Corylus rostrata californica A. DC.

Fairly common in the Transition zone in Squaw Creek Valley near
McCloud Mill, and probably elsewhere at the base of the mor.ntain.

142 NORTH AMERICAN FAUNA. [no. lo.

Castanopsis sempervirens (Kellogg) Dudley. Sierra Chinquapin.

The distribution of the mountain chin<iuapin on Shasta is discon-
tinuous. It is possible that two forms exist, one apparently restricted
to the manzanita chaparral of the Transition zone from Sisson up to
the lower edge of the Shasta fir Delt; the other to the scattered
stretches of Pinus albicanlis of the Hudsonian zone, where it reaches
timberline on the rocky slopes and ridges.

Prof. William K. Dudley, of Stanford University, California, has, at
my request, kindly looked up the proper name for the boreal Sierra
chinquai)in, which he linds to be senipervirens of Kellogg.' The type
locality of semperrirens is the west slope of the Sierra near Marii>osa.
I have found the species abundant on a ridge near a stage station
called Chinquapin, between Mariposa and the Yosemite, where it
occurs with Pimis Jeffrey i, P. lambertiana^ Abies in agnifica, Pseudotsuga
mucronafa, and Prumis emarginaia at and above an altitude of 6,200
feet. The locality, therefore, is along the overlapping borderland
between the Transition and Canadian zones.

Professor Dudley tells nie that the ' var. minor'' Bentham is the small
southern coast range form of the true coast chinquapin, Castanopsis
chrysophyJla, and that the type locality is the Santa Cruz Mountains.
C. chrysophylla is a handsome tree 75 to 125 feet in height, with large
leaves, ending in long, slender attenuate points; C. sempervirens is a
bush with small and relatively bluntly rounded leaves. I found both
species common on the Trinity Mountains : G. chrysophylla on the sunny
lower slopes in the Transition zone; C. sempervirens on the cold summit
in the lower edge of the Canadian zone, where it is ^associated with
Arctostaphylos nevadensis, Cerasus emarginata, Ceanothus velutinus, the
dwarf mountain form of Quercus chrysolepis, and the very distinct Q.
vaccinifolia.-

' Proc. Calif. Acad. Sci., I, p. 75, 1855 (reprint).

2 Quercus vaccinifolia Kellogg is another excellent species, usually confounded
with the dwarf mountain form of Q. chrysolepis, with which it has nothing to do.
Their zone relations are much the same as those of the two species of Castanoijsis, for
Q. raccinifoJia occurs along the lower edge of the Boreal, and ranges up through the
Canadian zone, always in rocky places, while <J. clirysolepis belongs to the Transition
zone. Their rauges join where these zones meet, anil I have found both growing
side by side ou the Trinity Mountains, and also on the Sierra. Quercus clirysolepis
is a Transition zone tree which at the upper limit of its range is always dwarfed
and often reduced to a shrub; but irrespective of size it always retains its charac-
teristic leaves and acorn cups. (^>uercus vaccinifolia is always a small bush — rarely
much over a meter in height — and, whether in f/'uit or not, is distinguishable at a
glance by tlie character of its leaves and cups. The leaves are smaller, narrower,
thinner, and blunter (commonly narrowly oval with an obtuse point instead of
sharply lanceolate) and lack the yellow totoentum underneath; furthermore, their
margins, although somewhat thickened, are not distinctly revolute. The acorn cups
are smaller and thinner, and lack the beautiful yellow 'turban' so characteristic of
clirysolepis ; the acorns average shorter nuA thicker and the basal scar is smaller. The
branchlets are much more slender, and glabrous or nearly so, instead of tomentose.

OCT.. 1899.] PLANTS. 143

It is remarkable that a shrub of such wide distribution, and one dif-
fering-so conspicuously from the tree chinquapin [Castiinop.sis chryso-
phi/Ua), should so lon^ escape recognition as a distinct species.

ftuercus californica (Torr.) Cooper. Bla(;k Oak.

[=Q. helloggi Newb.]

The only oak of Shasta, where it is confined to the basal slopes of
the Transition zone, reaching up on the south and west sides to an
altitude of 4,500 feet. It is fairly common in McCloud Valley, more
common at Sisson, and increases in abundance to the northward.
Between Black Butte and Edgewood it is mixed with iwnderosa pines
and is one of the most conspicuous trees. It does not attain such large
size in tliis region as nearer the coast and farther south in the Sierra.

IQuercus garrijana was not found immediately about Shasta, and
Quercns icislizeni, which pushes up the canyon of the Sacramento along
distance, does not fairly enter the region.]

Asarum hartwegi Watson. Wild Ginger.

Occurs here and there throughout the Transition zone, but is com-
monest in dam J) places on the lower slopes. It does not reach quite up
to Wagon Camp, but in a warm lava basin on the west side of the
mountain was found at an altitude a little higher than Wagon Camp.

Eriogonum marifolium T. & G. Yellow Eriogonum.

Common at Wagon Cami? and other points along the lower edge ol
the Canadian zone and in the upper part of the Transition. The species
seems to belong to the Transition rather than the Canadian zone, and
Shasta is its type locality. (Identified by J. K. Small.)

Eriogonum polypodum Small. Small-leaf Ali)ine Eriogonum.

The commonest and most widely distributed Eriogonum of the higher
slopes, where it ranges from the lower edge of the Hudsonian zone up to
and far above timberline. The highest altitude at which it was obtained
is 10,000 feet. Its leaves are small and densely covered with a white
woolly or hairy material, and its tortuous j)rostrate branches are so
intertwined as to form little mats several inches in diameter on the stony
pumice slopes; these whitish mats are vastly more comi)act and dense
above timberline than below. The roots are strong but rather slender,
and, like those of many other plants that live on the barren, wind-swept
pumice slopes, are of extreme length. The main root usually slopes
obliquely for 80 or 90 millimeters, and then divides into four or five
slender rootlets 900 to 1,000 millimeters in length. The whitish tomen-
tose leaves rarely rise more than 25 or 30 millimeters above the ground;
the fruit stems 100 to 150 millimeters.

This plant and Polygonum shastenae^kxe probably tlie most abundant,
conspicuous, and widespread plants of the Hudsonian and Alpine
zones. They thrive in very dry soils and theretbre are not confined

144 NORTH AMERICAN FAUNA. [no. 16.

to the moist basins and strips borderinj.^' the streams, as are Phyl-
lodoce, Lntlea, and n)aiiy others. (Ideutitied by J. K. ymall.)

Eriogonum pyrolsefolium Hooker. Larg^e-leaf Alpine Eriogonum.

Abundant on the pumice slopes of the Hudsonian and Arctic-Aljiine
zones, where it is associated with the foregoing' species and with Poly
gonum shasteu.se, 1\ neivberryi, Pentstenwn <larl(honi, and others. It is
easily recognized by its large roundish deep-green leaves, in striking
contrast to the much smaller whitish-tomentose leaves of its congener
and i\ss,oc\'dte, Erigonum polf/poduDi. Both species are common all the
way around the mountain. The present species {pyroIa'foUum) is rarely
found above an altitude of 9,500 feet, but on a warm slope east of Mud
Creek Canyon Vernon Bailey found it as high as 10,000 feet. It flow-
ered the second time above the head of Squaw Creek the latter part of
September and was then in flower and fruit simultaneously. Its root
is large, thick, tapering, and moderately strong; it subdivides into
about half a dozen rootlets which penetrate so deeply into the soil
that it is difficult to obtain specimens without breaking them. The
longest root measured was 750 millimeters to the broken end. The
leaves reach about 30 millimeters above the ground; the fruit stems
70 millimeters. The old imbricating leaf stems remain attached for
several years and form a series of scales around the upper part of
the perennial rootstalk.

Eriogonum nudum Dougl. Xaked-stem Eriogonum.

Common in the lower part of the Transition zone, i)articularly about
Sisson, where it was flowering plentifully as late as the end of Sep-
tember. It is easily recognized by its tall, green naked stems, which
rise from a bunch of large tomentose leaves.

Oxyria digyna (Linn.) Alpine Sorrel.

One of the characteristic alpine species, growing in cold spots among
the rocks at high altitudes on all sides of the mountain; found by
Vernon Bailey as high as 11,200 feet. On the north side of Shastina
we collected it at 8,900 feet, and on the northeast side of Red Butte as
low as 7,000 feet. This is the lowest station at which it was found, and
since Pinus alhicavUs occurs above Red Butte the locality might be
mistaken to be below timberline, but the sorrel grows only among the
rocks on the cold northeast sloj^es, where there are no trees and where
the temperature is truly alpine.

Polygonum shastense Brewer. Shasta Polygonum.

One of the commonest and most characteristic plants of the stony
pumice slopes of the Hudsonian and Ali)ine zones. A singular and
attractive plant, particularly in September, Avhen it is heavily laden with
white and red flowers and fruit. As a rule the flowers are whitish, turn-
ing red as the fruit begins to develop. No two plants could well be more
different in general ai)pearauce than this species and its congener and

OCT., 1899.] PLANTS. 145

associate Polyqamnn newberryi, and few species bear a closer resem-
blance than P. shastense and its geographically remote relative P. paro-
nyclila. The resemblance is not only most striking, but is exceedingly
interesting from the standpoint of geographic distribution. Poli/gonum
shastense lives at high altitudes in the High Sierra and Cascade Range,
while P. paronychia inhabits the outer sea beach in northern California
and Oregon, where it is bathed in the chilly fogs of the Pacific Ocean.
The root of Polygonum shastense is of moderate size and slopes very
obliquely into the soil. It divides into half a dozen long slender rootlets,
which penetrate to a depth of 550 millimeters or more. One specimen
examined divided into three main roots of rather large size, which
tapered very gradually and reached a length of 750 millimeters. The
branches are prostrate and usually form loosely intertwined mats 300
to 500 millimeters in diameter, rising in some cases 40 to 50 millimeters
above the surface, but usually flattened on the ground.

Polygonum newberryi Small. Broad leaf Polygonum.

Abundant from the lower edge of the Hudsonian zone up to a little
above timberline, where its big green leaves are very conspicuous on
the i)ale pumice soil and among the broken fragments of gray lava rock.
About the middle of September the leaves turn red — often a deep
handsome red — and begin to fall, so that by the end of the month the
plant has practically disappeared. Its buckwheat-like fruit is a favorite
food of the mice inhabiting the higher slopes.

Oreobroma triphylla (Wats.) Howell. Dwarf Alpine Spring Beauty.

This tiny inconspicuous plant was found near timberline north and
northwest of Red Butte. (Identified by Professor Greene.)

Spraguea umbellata Torr. Pussy-paws.

Abundant on the pumice slopes of the Hudsonian zone, beyond which
it pushes both upward and downward on suitable soils. The highest
altitude at which it was noted is 9,400 feet on the east side of Mud
Creek Canyon, but it was rarely seen above 9,000 feet. On the other
hand, a form of it descends in suitable spots to the lower edge of the
Canadian zone near Wagon Camp (altitude 5,600 feet), and to the same
altitude in the lower part of Mud Creek Canyon.

Stellaria crispa C. &, S.

Collected by Vernon Bailey and Miss Wilkins in Mud Creek Canyon
near the junction of Clear Creek. (Identified by Professor Greene.)

Sagina saginoides (L.) Brit.

Occurs above timberline; collected by Miss Wilkins.
Silene grayi Watson. Hudsonian Catch fly.

Common in stony places and along streams below timberline in the
Hudsonian zone. Found both in the stony pumice soil and in the
heather beds. (Identified by Miss Eastwood and Dr. B. L. Robinson.)
21753— No. 10 19

146 NORTH AMERICAN FAUNA. [no. 16.

Silene suksdorfi Robinson.' Alpine Catclitiy.

An alpine species coninion in scattered tufts, which form small com-
pact mats under the edges of rocks, well above timberline. Often
mixed with Erigeroii conipositus. (Identitied by Dr. B. L. Robinson.)

Aconitum columbianum Nutt. Monkshood.

Common in a marshy place in the Shasta fir forest at Wagon Camp,
where its tall wands of handsome blue tlowers were conspicuous in July
and its fruit in September.

Aquilegia truncata Fisch. & Mey. Red Columbine.

Common near Wagon Camp, in the upper part of the Transition
zone. (Identitied by Professor Creene.)

Delphinium sonnei Greene. Larkspur.

Common in moist spots in the Shasta fir forest near Wagon Camp,
and also in the canyon of Ash Creek. (Identified by Professor Greene.)

Pseonia browni Dougl. Wild Pa'ony.

Occurs sparingly in the u[)per part of the i)onderosa pine forest
(Transition zone), particularly in the neighborhood of Wagon Camp.

Pulsatilla occidentalis Watson.

Common in places on the higher slopes (Hudsonian zone), particu-
larly where the snow lies late. It blooms as the snow recedes, leaving
a handsome globular head of feathery plumes which waves in the
breeze long after the season of flowering.

Bikukulla uniflora ( Kellogg) Howell.

Reported by Miss Eastwood from above tindterline on Horse Camp
Trail; not fouiul by us.

Arabis platysperraa Gray. Flat- pod Arabis.

The commonest and most widely distributed crucifer of the higher
slopes of Shasta, where it occurs on stony slopes and along the edges of
the heather beds from the lower edge of the Hudsonian zone upward
on warm slopes to 10,200 feet. Its extreme vertical range appears to be
about 3,000 feet. (Identified by Miss Eastwood.)

Cardamine bellidifolia pachyphylla Coville.

This small alpine crucifer, with white flowers and rather broad dark
green leaves, is nowhere abundant, but was observed here and there
above timberline, both on the main peak of Shasta and on Shastina.
(Identified by F. V. Coville.)

Cheiranthus perennis (Coville) Greene. [ = Erysimum, asperum 2)erenne
Coville.]
Yernon Bailey and I found this coarse yellow-flowered crucifer at
timberline on the north slope of Shastina July 124, but did not observe
it elsewhere. (Identified by Professor Greene.)

i Botanical Gazette, vol. 16, p. 44, pi. 6, 1891.

OCT, 1899.] PLANTS. 147

Draba breweri Wats. Sierra Alpine Draba.

Collected east of Mud Creek Canyon at an altitude of 13,000 feet by-
Miss Wilkin s; not observed elsewhere. This Draba av.d rolemonium
pulchelhim are the only plants found at so great an elevation. (Identi-
fied by F. Y. Coville.)'

Streptanthus orbiculatus Greene.

A Hudsonian species common on pumice soil in stony places at and
below timberline all the way around the mountain. The plant is easily
recognized by its long curved slender pods and its rather large domed
leaves. The length of the large tapering root only slightly exceeds the
height of the plant. (Identified by Professor Greene.)

Chrysamphora californica (Torr.) Greene. California Pitcher Plant.
[^=l)arHngtoma Auct.)
This interesting pitcher plant is exceedingly local in distribution.
Mr. Elmer Aj)plegate tells me tliat it is conunou in the upper part of
the Sacramento Canyon, a short distance from Sisson Tavern. It has
been reported as occurring in the 'marshes' of Shasta, but we did not
find it on the mountain, nor did we find any marshy areas more than a
few rods in extent.

Drosera rotundifolia Linn. Sundew.

Collected by Miss Wilkins in the springy bog just above Wagon
Camp in the Canadian zone. Not observed elsewhere.

Mitella pentandra Hook.

Common in the heather patches along the overhanging banks of
streams in the Hudsonian zone. Abundant at Squaw Creek Camp.
(Identified by Professor Greene.)

Parnassia californica (Gray) Greene. Grass-of-Parnassus.

This elegant flower is common along the water courses of the Hud-
sonian zone, usually growing along the overhanging banks of the tiny
rivulets. It blossoms late, and a few plants were still in flower when
snow fell, the last week in September.

Saxifraga bryophora Gray.

Re])orted by ^Nliss Eastwood from the timberline region on Horse
Camp Trail; not found by us.

Saxifraga tolmiei' T. & G. Alpine Eock Saxifrage.

Common above timberline, where it usually forms small dense mats
in moist places among rocks. On Inconstance Creek, on the north side
of Shasta, it was found as low as 7,600 feet. On the south slope it
ranges up to 11,000 feet, or possibly higher. On the cold northeast
side of Eed Butte it occurs among the rocks with Oxyria digyna. (See
p. 50.)

' The spelling tolmcei should be regarded as an oIin ions typographical error.

148 NORTH AMERICAN FAUNA, [no. 16.

Peltiphyllum peltatum (Torr.) Engler. Giant Water Saxifrage.

One of the most cons])icuous i>lants in the bottom of the Sacramento
Canyon, where its clusters of huge incised roundish leaves along the
water's edge suggest the devil's club. It ranges along the Sacramento
Kiver from Just below Sisson nearly to the Sacramento Valley.

Ribes amictum Greene.

Occurs at Wagon Camp, along Squaw Creek, and in the bottom of Mud
Creek Canyon at an altitude of 6,700 feet. (Identified by Professor
Greene.)

Ribes cereum Dougi.

Found near timberlitie near the head of Panther Creek, and also
much lower down.

Ribes viscosissimum Pursh.

Common near Wagon Camp and observed at other points within the
Transition zone, particularly along Squaw Creek.

Ribes klamathense Coville.

Occurs in cool moist places in the Transition zone at Sisson. (Iden-
tified by F. V. Coville.)

Cercocarpus ledifolius Nutt. Mountain Mahogany.

Common on Sheep Rock and thence to the southeastern corner of
Shasta Valley (V. Bailey), and on warm, dry slopes of the Scott
Mountains, where C. jyarvifolius also occurs.

Fragaria bracteata Heller. Small Strawberry.

Common on the lower slopes. Just below Wagon Camp strawberries
and i)ainted cups are so abundant as to form an almost continuous car-
pet under the uppermost grove of ponderosa pines. (Identified pro-
visionally by P. A. Eydberg.)

Fragaria chile 5nsis Duchesne. Large Strawberry.

Occurs with the last a little below Wagon Camp, but is much com-
moner lower down, particularly near Sisson. Sisson Tavern was
formerly called 'Berryvale' and is located in 'Strawberry' Valley.
Both names were derived from the abundance of this wild fruit there
in early days. (Identified provisionally by P. A. Eydberg.)

Holodiscus discolor (Pursh) Maxim. Alpine Spirtea.

Common on rocks at and a little below timberline, usually associated
with Finus albicauUs. It is a small, fragrant bush, usually less than a
foot in height, and always grows among rocks at high altitudes. In
the Iludsonian zone it was found all the way around the mountain, and
was in blossom from about July L*0 until September. A larger form,
which the botanists do not appear to have named, occurs lower down,
in the Canadian zone. It has larger, broader, and thinner leaves and
should be separated.

OCT., 1899.] PLANTS. 149

Spiraea douglasi Hook. Red Spira?a.

Common in moist places in the Canadiau and Transition zones, but
most abundant in the latter. It is common at Sisson, at the west base
of the mountain, and in Squaw Creek Valley on the south side, and
thence upward, in suitable moist spots, to Wagon Camp at 5,700 feet,
along Squaw Creek at 6,000 feet, and in Mud Creek Canyon as high
as 6,700 feet.

Horkelia pseudocapitata Rydberg.

Abundant in the Transition zone near Wagon Camp, particularly
about the upper limit of Pinus ponder osa. (Identified by P. A. Rydberg.)

Kunzia tridentata (Pursh.) Spreng.

Irregularly distributed in the Transition zone; commonest in the
manzauita chaparral of the lower slopes. On the north side of the
mountain it is exceedingly abundant, and in the open pine forest bor-
dering the south end of Shasta Valley attains unusually large size.
On the west side, north of Sisson, it occurs si^aringly on most of the
warmer and drier knolls, and more i)lentifully between Black Butte
and Shasta Valley. It is almost always associated with Arctostapltylos
patula, with which it ascends some of the warmar slopes to points far
above the npper limit of its usual distribution. Tlius it was found on
a southwest slope in Mud Creek Canyon between the altitudes of 6,700
and 7,400 feet; on similar slopes near Horse Trail and in Diller Canyon
as high as 7,800 to 7,900 feet, and on a warm ijumice ridge north of
Shastina at 7,500 feet.' In Squaw Creek Valley, near McCloud Mill, a
form occurs which has exceedingly narrow leaves.

Lutkea pectinata (Hook.) Kuntze.

Abundant in the Hudsonian zone, chiefly in the neighborhood of
timberline, where it is common along the little streams in the upper
edge of the forest, and in the glacial basins which are wet from melting-
snows in the early part of the season, but may be dry at the time the
plant blossoms. In damp spots, particularly along the borders of cool
springs, the individual plants often stand so near together as to form
extensive beds.

Potentilla flabellifolia Hook.

Occurs here and there, a little below timberline, in the Hudsonian
zone. (Identifled by Miss Eastwood and P. A. Rydberg.)

Potentilla pseudorupestris Rydberg. Dwarf Alpine Potentilla.

A dwarf Alpine or high Hudsonian Potentilla of the (/landulosaty])e,
collected on the north side of Shastina at an altitude of about 8,800
feet, is x)i'ovisionally referred to this species by Mr. Rydberg.

' For an explanation of this .seemingly abnormal range, see p. 49.

150 NORTH AMERICAN FAUNA. [xo.l6.

Potentilla glandulosa Liiidl. Large Yellow Potentilla.

A large Potentilla from the Transition zone, near Wagon Cara]), is
identified by Mr. Ilydberg as P. glandulosa.

Rosa californica C. & S. California Kose.

Abundant in moist places in the Transition zone near Sisson Tavern
and along the east base of Mount Eddy.

Rosa gymnocarpa Nutt. Wild Hose.

Common in places in the Transition zone, particularly a little below
Wagon Camp.

Rubus parviflorus Nutt. [ = R. nutJcanus Auct. 1 Western Thimbleberry.
Common in cool moist places in McCloud Valley and at Sisson, and
thence up through the Transition zone to an altitude of 5,200 feet on
the road to Wagon Camp, and G,0()0 feet along Squaw Creek and in
Mud Creek Canyon. Most of the Transition slopes of Shasta are too
dry for the thimbleberry.

Rubus vitifolius C. & S. Blackberry.

Common in cool moist places in the Transition zone near Sisson and
along the east base of the Scott Mountains.

Sibbaldia procumbens Linn. Alpine Sibbaldia.

Common in the Hudsonian zone near timberline, particularly in
springy places and in the heather beds. Usually occurs in small
patches below extreme timberline.
Amelanchier alnifolia >utt. Serviceberry.

Abundant in moderately moist parts of the Transition zone. On the
west and southwest sides of the mountain it reaches from Sisson to
Wagon Camp. On a warm southwest slope on the steep ridge between
Mud Creek Canyon and Clear Creek it occurs, with several other
Transition zone species, at the unusual altitude of about 7,000 fpet.
(See p. 49.)

Crataegus rivularis Kutt. Black Haw.

Common in the Transition zone about the west base of Shasta, par-
ticularly along streams at the east base of Mount Eddy, from the head
of the Sacramento northward, usually in cool moist soil. Common near
Sisson Tavern.

Sorbus sambucifolia (C. & S.). Mountain Ash.

Rather scarce and confined chietly to the relatively moist Transition
zone sloi)es of the canyons. In Mud Creek Canyon it was found along
the bottom from 5,000 feet to 0,700 feet. Along Squaw Creek it was
found at about 5,500 feet.

Cerasus demissa (Nutt.). Western Chokecherry.

Common in places in the lower part of the Transition zone. Observed
on the south slope above jNlcCloud Mill, mainly in the gulches; also ia
the neighborhood of Sisson, and near the south end of Shasta Valley.

OCT., 1899] PLANTS. 151

Cerasus emarginata Doagl. Bush Cherry; Bitter Ked Cherry.

xVbuiuhiut ill phices in the chaparral of tlie Transition zone. In
Sacramento Canyon, south of Shasta, it begins at 'The Loop," and
ranges up to tlie lower edge of the Canadian zone. It is profusely
abundant at Wagon Cami), where it is a straggling bush a little higher
than a man's head, and usually grows in thickets. It ascends Mud
Creek Canyon to an altitude of o,(i()0 or 5,700 feet.

Cerasus glandulosa Kellogg.

A sai^ling or small tree having large broad leaves; collected by me
a short distance south of Sisson Tavern. (Identified by Professor
Greene.)

Prunus subcordata Benth. Wild Plum.

Common in places along the southern and western borders of Shasta
Valley, usually in the edge of the open Pinus ponderosa forest and
often growing with Rhus and Kunzia. We found it also near Etna,
on the west side of Scott Valley.

Cercis occidentalis Torr. Eed-bud.

lieported from Mount Shasta in the Botany of Califorina, but not
found by us except in the Sacramento Canyon, where it is common.

Lotus americanus (Nutt.) Bisch. [= Hosackia americana.]

Common in the Transition zone at Wagon Camp, and thence along
the road to Sisson, growing chiefly in open places in the chaparral.

Lupinus elmeri Greene [= Ij. «lbicai(lis si/lvestris Auct.]

Common in plactr-s througliout the ui)per i)art of the Transition zone,
and ranging into the Canadian. Common at Wagon Camp and a little
above. Found also in Mud Creek ( 'anyon at mouth of Clear Creek.
(Identitied by J. B. Leiberg.)

Lupinus 'ornatus' Auct. [not of Douglas.] Silvery Lupine.

This beautiful species is common in spots near and a little below
tiniberline, sometimes covering extensive areas, but not uniformly dis-
tributed. Where it occurs it is usually sufficiently abundant to give
the mountain side the effect of a distinct silvery covering. It always
grows on stony or pumice slopes and usually among or near white bark
pines, although in a few instances it was found outside of the pine
areas. It is common near the head of Squaw Creek and on both sides
of Mud Creek Canyon. On the east side of this canyon it is particu-
larly abundant just above the trees, and stops abruptly at an altitude
of 8,200 to 8,100 feet, to be replaced by the dwarf Alpine L. hjalli. On
the north side of Shasta it is common in an open forest of white-bark

'In this and subsequent references to 'The Loop' the statement means that in
goiny north along the railroad track we first saw the i)lant at * The Loop.' The
species may occur farther south in the canyon on suitable slopes above the level of
the railroad.

152 NORTH AMERICAN FAUNA. [no.ig.

pines in a shallow gulch at the east base of the lava buttes just below
'North Gate.' The upper limit of the silvery lu[)ine usually coincides
with the lower limit of the dwarf lupine [L. h/alli), which species gen-
erally i^ushes from this point upward through the Alpine zone.

The root of the silvery lupine is slender and tough, and soon divides
into two or three very long wire-like rootlets which run a rather shallow
course in the sand. Some of them measure 750 millimeters. The plant
at timberline averages about 00 millimeters in height.

Mr. Lei berg tells me that this species, although commonly referred
to '•ornatus\ is not or nut us of Douglas. It has also been called L.
argenteus decumhens Watson.

Lupinus albifrons Benth.

Collected near Horse Camp August 20 by Vernon Bailey and Miss
Wilkins. (Identified by J. B. Leiberg.)

Lupinus lyalli Gray. Dwarf Alpine Lupine.

Abundant and widely distributed over the higher rocky pumice
slopes from timberline or a little above up to an altitude of slightly
more than 10,000 feet. (Identified by J. B. Leiberg.)

Lupinus minimus Dougl. Dwarf Lowland Lupine.

Common in the Transition zone at Sisson. (Identified by J. B. Lei-
berg.)

Vicia americana Muhl.

Bather common at and below Wagon Camp, and still in fiower when
we left, September 25.

Linum lewisi Pursh. Wild Hemp.

Abundant at Wagon Camp, where its delicate blue llowers were con-
spicuous in July, and its large subglobular seed capsules in September.

Polygala cornuta Kellogg.

Occurs plentifully in the dry pine woods of the Transition zone near
Sisson Tavern, but was not observed on the mountain proper. (Iden-
tified by Miss Eastwood.)

Rhus trilobata Nutt.

In going north from Sisson we first observed this species a mile or
two south of Edgewood, to the north and east of which it became more
and more common. Its zone position here is along the borderland
between the Transition and Upijer Sonoran zones.

Pachystima myrsinites Raf. Oregon Boxwood.

Common in the Transition zone at the west base of Shasta, from Sis-
son up to an altitude of about 4,700 feet, usually in manzanita chapar-
ral. Its absence from the higher slopes within the proper zone limits
of the species is probably due to heat and dryness, as explained else-
where (p. 5(1), but it is possible that the Sisson plant is a Transition
zone subspecies of the true Boreal P. myrsinites.

OCT., 1809] PLANTS. 153

Acer macrophy Hum Pursh. Oregon Maple; Big-leaf Maple.

The tree maple is rare in the region about Shasta, where it was
observed only in moist places in tlie lower part of the Transition zone
near Sisson and in the upper part of the Sacramento Canyon.

Acer glabrum Torr. Bush Maple.

Fairly common in McOloud Valley and in moist places along streams
and canyons in the Transition zone. In Mud Creek Canyon it was
found up to an altitude a little above 5,600 feet, and along Squaw Creek
to nearly 6,000 feet.

Acer circinatum Pursh. Vine Maple.

This characteristic west-coast species, with nearly circular 7 -point
leaves, occurs sparingly in moist places near Sisson Tavern, but is not
common. It is one of the most distinctive plants of the humid Pacific
coast division of the Transition zone.

Ceanothus cordulatus Kellogg. Snow Bush.

Common in the lower part of the Transition zone on the southern
and western basal slopes of Shasta. In the upper part of Sacramento
Canyon it first occurs at 'The Loop,' whence it is abundant northerly
to within a mile or two of Edgewood. At Sisson and in Squaw Creek
and McCloud valleys it is very abundant, but for some reason not
apparent it does not reach on Shasta its usual upper limit, and was
not observed anywhere above 4,900 feet. Possibly it is choked out by
the other chaparral, which is made up mainly of manzanita {Arcto-
staphylos patula) and buck-brush [Ceanothus velutinus).

Ceanothus cuneatus Xutt. Wedge-leaf Ceanothus.

This is one of the most characteristic shrubs of tlie Upper Sonoran
zone in California, where it is generally associated with the curious
digger pines. The region about Shasta is too high for it. It occurs,
however, in the lower valleys, both north and south of Sisson. On the
north it reaches from Yreka to a little beyond Edgewood, appearing a
mile or two south of the latter point on the road to Sisson, and about
4 miles southwest of it on a road farther west. Skipping the broad
Transition belt between Edgewood and the Sacramento, it occurs next
at Delta, in the Sacramento Canyon, and ranges thence southerly. Its
upper border meets the lower border of another species, C. cordulatus,
which is common at Sisson, as well as on the southern and western
basal slopes of Shasta.

Ceanothus velutinus Dougl. Buck-brush.

Profusely abundant on all the manzanita-covered slopes of Shasta,

from the lower part of the Canadian zone down nearly to the lower

edge of the Transition. Mixed with Arctostaphylos patula in nearly

equal proportion it forms the dominant chaparral of the mountain.

21753— No. 16 20

154 NORTH AMERICAN FAUNA. [no. 16.

Tongues of it push up on warm southerly slopes to about 6,600 feet'
altitude, aud it descends on the west slope to about 4,200 feet (half a
mile east of Sisson), and on the south slope to 3,600 feet (near McCloud
Mill). Along the east base of Mount Eddy, where it is sheltered from
the afternoon sun, it appears about 2 miles north of Sisson (altitude
3,700 feet), and continues northerly for many miles.

Ceanothus integerrimus H. & A. California Lilac.

Common in parts of the Transition zone, notably in Squaw Creek
Valley near McCloud Mill, and in the upper Sacramento Valley a little
below Sisson, but very scarce or absent on the slopes of Shasta proper.
Like G. cordulatus it seems to be kept out by some unknown cause,
possibly choked out by other species. Perhaps the soil is too dry for it.

Ceanothus (Cerastes) prostratus Benth. Squaw Carpet.

In following the Sacramento Canyon northward, Sc^uaw Carpet was
first seen at the ' The Loop,' a few miles south of Sisson, whence it
occurs in greater or less abundance throughout the ponderosa ])ine
forest and manzanita chaparral to the lower slopes of Shasta, where on
the southwest side it reaches up to 5,200 feet.

Rhamnus californica Esch. (This form may be B. rubra Greene.)

Scarce and confined to low altitudes in the Transition zone. Found
by Vernon Bailey in Squaw Creek Valley below 4,000 teet. Specimen
lost.

Hypericum anagalloides C. & S. Dwarf Hypericum.

Common in spots in wet places in the Hudsonian and Canadian zones.
It was most common in the second meadow on Squaw Creek, aud by
no means rare near Wagon Camp, and was in flower in late July aud
throughout August.

Viola blanda Willd. White Violet.

Collected by Miss Wilkins in the middle meadow on Squaw Creek,
at an altitude of 7,500 feet. Not observed elsewhere.

Viola purpurea Kellogg. Alpine Yellow Violet.

Common on dry rocky slopes on pumice soil from some distance
below timberline upward, on warm southerly exposures, to 9,300 feet.
When we reached the mountain, the middle of July, its yellow flowers
were conspicuous, although even at that time the species had nearly
passed flowering. Its fruit and dark green leaves remained as late as
the latter part of September, sharply outlined against the whitish
stones and pumice of the bleak and barren upper slopes, where a violet
seemed singularly out of place. (Ideutilied by Miss Eastwood and
Professor Greene.)

' For its extreme upper limit, on hot canyon slopes, see remarks under Arctoata-
pht/loa patula, p. 158.

OCT., 1899.] PLANTS. 155

Mentzelia laevicaulis (Dougl.) T. & G.

Rather common in Shasta Valley, but not found elsewhere about the
mountain. This species seems to belong to the Upper Sonoran zone,
and therefore has no place in the flora of Mount Shasta.

Epilobium spicatum Lamarck. Willow Herb ; Fire- weed.

Abundant in places, chiefly on old burns in the Canadian and Tran-
sition zones. At Wagon Camp it did not begin to flower until near the
middle of August; at Sisson it was still in flower in places as late as
the middle of September.

Epilobium brevistylum Barbey.

Collected at Wagon Camp, where it is tolerably common in moist
places; the largest species next to spicatum. (Identified by Professor
Trelease.)

Epilobium oregonense Hausskn.

Common at Wagon Camp. A large form occurring also at Wagon
Camp Professor Trelease considers as probably E. glaberrinium Barbey.

? Epilobium hornemanni Eeichenb.

Some poor specimens, with exceedingly long seed capsules, collected
by me a little below timberliue about the end of the season, are doubt-
fully referred to this species by Professor Trelease.

Epilobium clavatum Trelease.

Abundant in the heather meadows and along the edges of the cool
streams of the Hudsonian zone at and a little below timberline.
(Identified by Professor Trelease.)

Epilobium pringleanum Hausskn.

A tiny alpine species, rarely as tall as one's thumb, occurring here
and there on the borders of the highest streamlets above timberline.
(Identified by Professor Trelease.)

Epilobium obcordatum Gray. Rose Epilobium.

Local — common near timberline on the steep west side (east slope) of
Mud Creek Canyon. A remarkable plant, strikingly unlike the others
of its genus. It has a woody base and is really a dwarf bush. Its
large red flowers are showy and very handsome. (Identified by Miss
Eastwood and Professor Trelease.)

Gayophytum ramosissimum T. «& G.

Abundant throughout the Transition zone, where it was equally com-
mon at Sisson and at Wagon Camp and was in flower as late as the end
of September. (Identified by Miss Eastwood.)

Carum gairdneri (Hook. & Arn.) Gray.

Rather common in damp soil at Wagon Camp where the Canadian
and Transition zones meet. (Identified by Dr. J. N. Rose.)

156 NORTH AMERICAN FAUNA. [no. 16.

Cymopterus terebinthinus T. & G.

Commou iu the neighborhood of timberline, usually in stony places
on dry j^umice slopes; found also on top of lied Cone, a little east of
Wagon Cami). This species has a large and woody root and a remark-
ably rank and lasting odor. Its zone position is Hudsonian; a larger
unrecognized form occurs in the Canadian zone. (Identified by Miss
Eastwood and Dr. Rose.)

Ligusticum grayi Coulter & Rose.

Common throughout the Hudsonian zone, i)articularly a little below
timberline, where it occurs most abundantly iu the beds of heather
along the little streams. In places it pushes down into the Canadian
zone. (Identified by Miss Eastwood and Dr. Rose.)

Heracleum lanatum Michx. Cow Parsnii*.

Occurs in damp places in the Canadian zone. In Mud Creek Canyon
near the mouth of Clear Creek it was found as high as 6,700 feet.
(Identified by Dr. Rose.)

Osmorrhiza nuda Torr.

Found sparingly at Wagon Camp and at Sisson.
Cornus nuttalli Audubon. Oregon Dogwood.

Rather common in moist places in the lower half of the Transition
zone. It occurs in Squaw Creek Valley near McCloud Mill, and is
common along the streams near Sisson Tavern; on the road from
Wagon Camp to Sisson it was seen up to an altitude of 4,500 feet.

Cornus pubescens Kutt.

Common in cool damp soil near Sisson and along the upj)er Sacra-
mento.

ChimapMla umbellata (L.) ISTutt.

Occurs here and there throughout the drier parts of the forest, but
is less common than Ghimaphila menziesi.

Chimaphila menziesi Spreng.

Fairly common throughout the Shasta fir forest, and also in the
mixed forest of pines and firs from the upper Sacramento Canyon
northward (Canadian and Transition).

Pyrola picta Smith.

Occurs here and there throughout the forest of Shasta firs, where
its ornamental light-marbled leaves are pleasingly conspicuous against
the dark soil.

Pyrola pallida Greene. '

Found sparingly on the dry summit of Red Cone, about a mile east
of Wagon Camj). (Identified by Professor Greene.)

1 Pittonia, IV, p. 39, March 17, 1899.

PLANTS.

157

Pyrola secunda Liiui.

Decidedly less couiniou than P. picta, but like it found in the Shasta
fir forest.

Pterospora andromedae i^utt. Pinedrops

Found in the dry woods along the border between the Canadian and
Transition zones. (Identified by Professor Greene.)

Pleuricospora fimbriolata Gray.

Collected at Wagon Camp by Miss Wilkins. (Identified by F. V.

Coville.)

Sarcodes sanguinea Torr. Snow Plant.

This handsome plant is rei)orted as common on the forested slopes of
Shasta at the time of melting snow in spring. It jjrobably occurs in
both the Canadian and Transition zones.

Arctostaphylos nevadensis Gray. Dwarf Mountain Manzanita.

Common in the Canadian and Hudsonian zones, growing in extensive
beds a foot or less in height. On the high ridges, among the timber-
line Pmus alhicaulis, these beds of green cover the pale gray lava
rocks, and in the dark forests of Shasta fir they form the only con-
spicuous surface vegetation.

Arctostaphylos patula Greene. Green Manzanita.

The most abundant and troublesome chaparral of Shasta. It is a
characteristic Transition zone species and covers the lower slopes all
the way around the mountain except a belt about 10 miles wide on the

Maii/.aiiita chaparral.

northeast base, reaching from Ash Creek to about 3 miles northwest of
Inconstance Creek, which is too cold for it and is occupied by Canadian
zone species. On the north, west, and south it covers practically the
whole of a broad belt several miles in width, reaching from base level

158 NORTH AMERICAN FAUNA. [no. 16.

to the lower edge of the Canadian zone and formerly occupied in the
main by a forest of ponderosa pines, some of which still remain scattered
over it. Except at its extreme lower limit, it is usually mixed with
buck-brush {CeanotJms velutinus).

Seen from a distance, the extensive areas of manzanita on the lower
slopes of Shasta are very deceptive. They look like meadows of green
grass, but to cross them is in most cases impossible, owing to the den-
sity of the growth and rigidity of the branches. For this reason they
form secure retreats for black bears, deer, wild-cats and other animals.

At two places on the west side of the mountain Arctostaphylos patula
reaches the extraordinary altitude of 7,800 to 7,900 feet. One of these
is on the north side of Diller Canyon, the other on the north side of
Horse Camp Trail. Both stations are on long and steep southwest
pumice slopes which receive the hot rays of the afternoon sun almost
at a right angle, carrying up numerous Transition zone si)ecies nearly
2,000 feet above their normal limit. (See p. 51.) From the southwest
base of Shasta the green manzanita reaches down the canyon of the
Sacramento Kiver to 'The Loop.'

Phyllodoce empetriformis (Gray), lied Heather.

\=Bryant}ms empetriformis Auct.]

Abundant along the cold streams of the Hudsonian zone and in the
bottoms of the glacial basins that are kept moist by melting snows;
commonest between the altitudes of 7,500 and 8,500 feet, and not seen
above 9,100 feet. This is the only 'heather' found on Shasta. It
usually forms extensive beds or carpets in which numerous other
plants find a congenial home. These beds are practically the only
flower gardens on the mountain, and the only areas where small plants
are plentiful enough to give the effect of continuous green; hence they
are commonly spoken of as 'heather meadows.'

Kalmia glauca microphylla Hook. Alpine Laurel.

Abundant in most of the heather meadows just below timberline,
particularly along Squaw Creek. In general size and aspect the alpine
laurel resembles the red heather so closely that at a little distance it is
difficult to tell them apart.

Vaccinium caespitosum Mich. Dwarf Alpine Blueberry.

Abundant on the higher slopes within the Hudsonian zone, reaching
a little above timberline. Common here and there in the stony
pumice flats and basins, but commonest near the streams, where it is
scattered through the heather and forms beds of its own along the
outer edges of the heather beds. Its leaves turn dark red or garnet dur-
ing the latter half of September, contrasting handsomely with the dark
green of the heather. In the neighboihood of timberline the dwarf
blueberry is rarely more than three or four inches in height. It was in
full bloom the middle of July.

OCT., 1899.] PLANTS. 159

The form here referred to is the dwarf High Sierra one, given under
ca'spitosum by Coville, in liis 'Botany of the Death Valley Expedition'
(p. 145, 1893).

Vaccinium arbuscula (Gray) | = Yaccinium ccespitosum var. arb}iscnla
Gray].
Common in moist places in the Transition zone from Wagon Camp
down to Warmcastle Soda Springs at the south base of the mountain.
This species seems to be distinct from the dwarf boreal V. vwspitosum.
It averages about 2 feet (60 centimeters) in height and has red branches
and decidedly larger and thicker leaves than the alpine ca'spitosum.
Whether or not the latter is the typical form is anotlier question. The
zone position of V. arbuscula is probably Canadian and Transition.

Vaccinium occidentale Gray. Blueberry.

Common in the Canadian zone in some of the canyons, and in the
marsh at Wagon Camp, where it fruited early.

Fraxinus oregana Nutt. Oregon Ash.

Occurs sparingly in the upper Sacramento Canyon, but was not
observed about the actual base of the mountain.

Gentiana simplex Gray. Blue Gentian.

Common in the Canadian zone swamp at Wagon Camj) the first half
of August. (Identified by Professor Greene.)

Apocynum pumilum (Gray) Greene.

Common in suitable places throughout the Transition zone, from
which it pushes up a short distance into the lower edge of the Cana-
dian. With other Transition zone species it was found on a warm
slope in Mud Creek Canyon at 6,700 feet. (Identified by Professor
Greene.)

Cycladenia humilis Benth.

Common in places in the Hudsonian zone, l>ut very local. It occurs
plentifully on the top of Eed Cone, a short distance northeast of Wagon
Camp, at an elevation of about 6,600 feet; on a red lapilli hill which
forms a part of Eed Butte, just east of ' The [South] Gate,' and on both
sides of Mud Creek Canyon a little below true timberline. The highest
altitude at which it was observed was 8,700 feet on a southwesterly slope
on the west side of Mud Creek Canyon. Cycladenia JtumiUs is a curious
and rather striking plant. It usually has four large entire leaves, sug-
gesting those of Pohigonum neicherryi, and bears a pair of conspicuous
red tubular flowers followed by two curious seed pods, which are very
long and lie side by side, one above the other, like the barrels of a Win-
chester rifle. (Identified by Miss Eastwood.)

Gilia aggregata (Pursh) Spreng.

Common throughout the ponderosa pine forests of the Transition zone
and sometimes seen in the lower part of the Shasta fir forest, where its

160 NORTH AMERICAN FAUNA. [no. 16.

liandsome scarlet flowers are ^'ery attractive. On warm southwest
slopes near Panther Creek it was found as high as 6,600 feet, and
between Mud and Clear creeks as high as 6,700 feet, but these are
abnormal altitudes due to unusually warm exposures and soil.

CoUomia grandiflora Dougl.

Common in the upper part of the Transition zone a little below Wagon
Camp, where it was flowering abundantly the last half of July.

Phlox doug-lasi diffusa (Benth.) Gray. Alpine Phlox.

A common, widely distributed, and conspicuous plant of the higher
slopes within the Hudsonian zone, sometimes straggling down into the
Canadian zone. It occurs in scattered tufts on the dry rocky slopes
and ridges, usually on i)umice soil, and is commonest in the neighbor-
hood of timberline. It blossoms early, and flowers were rarely seen
as late as the latter part of July. After flowering the whole plant
withers and is disintegrated by the wind, so that little more than the
woody base remains.

Polemonium pulchellum Bunge.

A characteristic but not abundant Alpine species, occurring here
and there among rocks far above timberline. On the north side of
Shastina it was in full bloom July 24 at an altitude of 8,900 feet. On
the south side of Shasta, above Squaw Creek, and on both sides of
Mud Creek Canyon, it was not found below 9,500 feet, whence it ranges
up to 13,000 feet. This species and Draba hreiceri were both observed
at 13,000 feet, and are the highest plants found on Shasta. (Identified
by Professor Greene.)

Phacelia frigida Greene.' Dwarf Alpine Phacelia.

This new species, which Professor Greene has kindly described at
my request from specimens collected by us above the head of Squaw
Creek, is common and widely distributed on the higher and more bar-
ren rocky slopes, beginning above timberline and reaching, on south-
erly slopes, as high at least as 10,200 feet. The lowest altitude at
which it was f<mnd is 8,700 feet, on a cold slope.

Phacelia magellanica (Lam.) Coville.

A i)lant which, in the present unsatisfactory state of the group, it
seems necessary to refer here, is abundant on the lower slopes, par-
ticularly in the Transition zone, where it was in flower throughout the
summer and as late as the end of September.

Cryptanthe geminata Greene.

Rather common in the Transition zone below Wagon Camp. (Identi-
fied by Professor Greene.)

' Pittoiiia, IV, pp. 39-40, March 17, 1899.

OCT., 1899.] PLANTS. 161

Lappula nervosa (Kellogg) Greene.

Abundant in the oj)ening.s near and a little below Wagon Camp,
where it is the most troublesome 'stick-tight' of the region, filling the
forelocks and manes of the horses and binding them together in a
dense mat.

Monardella odoratissima Benth.

Abundant in the Canadian zone and much less common in the
Hudsonian. Abundant in the chaparral at Wagon Camp and found
in several places on warm sunny slopes as high as 7,800 or 7,900 feet,
and in one place near the head of Squaw Creek at 8,300 feet. In the
latter locality only a few bunches occur among the rocks and no others
were observed for a long distance below.

Scutellaria nana Gray. Dwarf Skullcap.

This interesting little yellow-flowered skullcap was found at one
place only — the north sloi)e of Shastina, at an altitude of 8,800 feet,
where it was flowering July 24. If this alpine plant is the same as the
type of S. nana, which came from the hot desert region near Pyramid
Lake, Nevada, its occurrence at tiinberline on Shasta must be acci-
dental. (Identified by F. Y. Coville.)

Stachys ingrata Greene.

Abundant in moist soil at Wagon Camp, on the border between the
Canadian and Transition zones. (Identified by Professor Greene.)

Chamaesaracha nana Gray.

This solauaceous plant, which has la ge white flowers and looks like
a dwarf potato, is common in an old burn near Wagon Camp, where
the Canadian and Transition zones overlap. (Identified bj^ Miss East-
wood.)

Castilleja miniata Dougl. Scarlet Painted Cup.

Common and conspicuous. A large form, probably the type form —
since the type came from the Blue Mountains of Oregon — is common in
moist places in the lower part of the Canadian zone and upper part of
the Transition. It was flowering abundantly in the uppermost grove of
ponderosa pines at Wagon Cami) the latter half of July. A smaller
form, apparently unnamed, abounds in the heather beds of the Hud-
sonian zone near and a little below timberline, where it was flowering
profusely the first half of August.

The scarlet painted cup is one of the handsomest flowers of the moun-
tain. Where the plants are abundant on the dark heather the effect
of the bright green corolla tubes protruding in slender spindles from
the vivid scarlet mass of bracts and calyx is superb. Early in August
the calliope hummingbirds were constantly hovering over these flowers.
(Identified by ^I. L. Fernald.)
21753— No. 10 21

162 NORTH AMERICAN FAUNA. [no. 16.

Castilleja affinis Hook. & Arn.

A plaut collected in the Hudsouian zone near timberliue just north
of E-ed Butte by Vernon Bailey has been identified as this species by
M. L. Feruald. Since the type locality of affinis is the low coast strip
near San Francisco or Monterey, the typical form would hardly be
expected to occur in the high timberline region of Shasta.

Orthocarpus pilosus Watson.

Common in places near timberline, particularly near 'The [South]
Gate,' between the heads of Panther Creek and Squaw Creek. (Iden-
tified by M. L. Fern aid.)

Mimulus implexus Greene.

Abundant in wet places throughout the Hudsonian zone, reaching
its greatest perfection in the neighborhood of timberline. In the shal-
low rapids of some of the mountain rivulets it grows in such profusion
that its leaves form extensive mucilaginous jjatcheSj which completely
fill the beds of the streams. Its large and showy yellow flowers were
in blossom from the latter part of July until the end of September.
(Identified by Professor Greene.)

Mimulus moniliformis Greene.

Two forms of large yellow Mimulus grow in the marsh at Wagon
Camp, in the lower edge of the Canadian zone. (Identified by Professor
Greene.)

Mimulus tilingi (Regel.) Greene.

Common in the marsh at Wagon Camp. (Identified by Professor
Greene.)

Mimulus primuloides Benth.

Common in patches in marshy places and along streams in the Cana-
dian and Hudsonian zones, but much more common in the Canadian
than in the Hudsonian. It was abundant at Wagon Camp and also in
some of the heather meadows along Squaw Creek and other streams.
(Identified by Miss Eastwood and Professor Greene.)

Pentstemon menziesi Hook.

One of the most characteristic and widely distributed plants of the
higher slopes, where it is common among the bare rocks all the way
around the mountain. It is common in the Hudsouian zone in the
neighborhood of timberline, but much more abundant in the Alpine,
and was found by Vernon Bailey as high as 10,200 feet. This species
is easily distinguished from the other Alpine species of the region by
its smooth entire leaves and relatively small bluish or violet blue flow-
ers, which become pink or reddish with age. (Identified by Professor
Greene.) The plant is by no means typical, but seems to be interme-
diate between menziesi and davidsoni. It may be a distinct species.

OCT., 1899.] PLANTS. 1 63

Pentstemon newberryi Gray.

Coimnou in places in the Hudsoniau zone and possibly in the upper
part of tbe Canadian; inucli less generally distributed than P. menzlesi
and usually occurring- at lower altitudes. On tbe north side of Shastina
it was flowering, the latter part of July, at an elevation of 7,600 feet,
and at the same time on the cold east slope of Mud Creek Canyon as
low down as 5,600 feet. Its flowers are much larger than those of
P. menziesi and red instead of bluish; its leaves are longer, and are
serrate instead of entire.

Pentstemon glaber utahensis Watson.

Abundant just below tiraberline on the curious Pimis alhicanlis plain
between North (late and Bolam Creek, on the north side of Shasta,
where it was flowering plentifully July 24. It was not seen elsewhere
on the mountain. Its blue-purple flowers are even larger than those
of P. newberryi and are very showy and handsome. (Ideutitied by
Professor Greene.)

Pentstemon deustus (Dough). Yellow Pentstemon.

Common in the rough black lava beds in the Shasta hr forest north
of Cascade Gulch, on the west side of the mountain, at an altitude of
about 7,500 feet, but not seen elsewhere. This species is a dwarf bush
5 or 6 inches high, with yellow flowers and strongly serrate leaves.
(Identified by Professor Greene.)

Pentstemon gracilentus Gray.

Abundant in places in the Shasta fir forest, particularly between the
canyons of Mud and Ash creeks, in the Canadian zone. (Identified by
Miss Eastwood and Professor Greene.)

Pentstemon confertus Dougl. (Not typical.)

Common near Wagon Camp. (Identified by Miss Eastwood and
Professor Greene.)

Veronica cusicki Gray.

Common just below timberline, particularly along the little streams
where it grows in the grass and heather on the overhanging banks. It
was flowering abundantly from the time of our arrival, the middle of
July, until about the middle of August. (Identified by Miss Eastwood
and Professor Greene.)

Boschniakia strobilacea Gray.

Collected at Wagon Camp by Miss Wilkins. (Identified by Profes-
sor Greene.)

Lmnsea borealis Linn. Twin Bell-flower.

Fairly common along one of the streams half a mile north of Sisson
Tavern, but not noticed elsewhere. This form was described by Torrey
as variety longijlora.

164 NORTH AMERICAN FAINA. [no. 16.

Sambucus melanocarpa (rray. Blackberry Elder.

Common in moist parts of the Transition zone, particularly in the
canyons. (Identified by Profes.sor (Ireene.)

Symphoricarpos racemosus Mich. Snowberry.

Common about Sisson Tavern and in cool moist places along the
base of Mount Eddy.

Symphoricarpos pilosus Greene. Mountain Snowberry.

Abundant in the upper part of the Transition zone, particularly
at Wagon Camj) and along Squaw Creek. (Identified by Professor
Greene.)

Campanula wilkinsiana Greene.' Shasta Bluebell.

This new bluebell, recently described by Professor Greene from speci-
mens collected by us on the upper part of Squaw Creek, is common in
the lower heather meadows, and less so in the marshy place at Wagon
Camp. It is singular that the only Campanula found on Shasta should
be new, the usual Sierra and Cascade species being absent. The pres-
ent si)ecies, which is named after Miss Lewanna Wilkins, averages 4 to
5 inches in height and has a rather small flower. Its zone position is
Hudsonian and Canadian.

Achillea lanulosa Nutt.

Common in the Transition zone at and below Wagon Camp. (Identi-
fied by O. L. Pollard.)

Achillea borealis Bong. Dwarf Alpine Yarrow.

Occurs in places on the stony pumice slopes above timberline. Be-
tween the heads of Mud and Squaw creeks Vernon Bailey found it at
an altitude of 9,000 feet. (Identified by C. L. Pollard.)

Anaphalis margaritacea (L.) B. & H.

Common in dry open jjlaces in the chaparral from Sisson to Wagon
Camp. (Identified by Professor Greene.)

Antennaria geyeri Gray. Geyer Everlasting.

A large specimen of this handsome everlasting was collected by Ver-
non Bailey in the Transition zone just below Wagoh Camj) September
25. The species was not noted elsewhere. (Identified by Miss East-
wood and Professor Greene.)

Antennaria media Greene [ = A. alpina Auct.]. Alpine Everlasting.

Common in open parts of the Hudsonian zone in the neighborhood
of timberline on most parts of Shasta and on Shastina. On ordinary
slopes it ranges from an altitude below 8,000 feet up to about 9,000.
(Identified by Professor Greene.)

' Pittonia, IV, pp. 38-39, March 17, 1899.

OCT., 1899.] PLANTS. 165

Arnica merriami Greene,'

Common in the heather meadows and along the edges of streams in
most parts of the Hadsoniau zone. This new species has just been
described by Professor Greene from specimens collected by us on the
moist banks of the small west arm of Upper Sijuaw Creek, under the
alpine hemlocks, where it is very common.

Arnica longifolia D. C. Eaton.

Common in moist places in the Canadian zone, chiefly on the banks
of streams. (Identified by Professor Greene.)

Arnica viscosa Gray.

This singular species, wliich differs strikingly in appearance and
habit from most members of its genus, grows in dense patches, usually
2 or 3 feet in diameter, among the bare lava rocks on steej) slopes near
timberline. On the south side of Shasta it was fairly common and
flowering abundantly in August. (Identified by Miss Eastwood.)

Artemisia tridentata Nutt. True Sagebrush.

Sagebrush is unknown in the region about Shasta except in Shasta
Valley, which it invades and overspreads from the Klamath country
on the north. It reaches the extreme south end of Shasta Valley and
penetrates a short distance into the bordering forest of ponderosa pines,
where it mixes with Kunzia tridentata. Pushing southeasterly over
Shasta Valley it reaches the gap at Sheep Rock. Its zone position is
Upper Sonoran and Transition.

Artemisia ludoviciana 2vutt.

A form, ot Artemisia Judoviciana is common in places just below Wagon
Camp, along the upper edge of the Transition zone. When the species
and subspecies now lumped under the above name are ])roperly defined,
several useful zone phints may be added to their appropriate lists.

Machaeranthera shastensis Gray. Shasta Alpine Aster.

[^= Aster sliastensis Auct.].

A dwarf alpine species abounding on the bare rocky x)umice slopes
in the neighborhood of timberline, and common also on the borders of
the heather beds in the glacial meadows. This is the type form. A
related but unnamed form, usually a foot and a half or more in height,
is common and widely distributed on the lower slopes, chiefly in the
upper part of the Transition and lower part of the Canadian zones.

Oreastrimi alpiginum (Gray) Greene.

Common in the neighborhood of timberline, particularly in moist
places. (Identified by Professor Greene.)

' Pittonia, IV, p. 3(5, :\rarch 17, 1899.

1G6 NORTH AMERICAN FAUNA. |N'Mf!.

Chsenactis nevadensis (Kellogg) Gray. Alpine Cbienactis.

This curiovis composite, with whitish tubular flowers, much divided
sticky leaves, and a smell like that of alcoholic s])ecimeus, occurs here
and there in the pumice sand and among the broken lava shale above
timberline. It was first seen in flower on the north side of Shastina
July 24 (alt. 9,000 feet), and early in August was flowering plentifully
on the south slope of Shasta above the head of Squaw Creek (alt.
9,300 feet). (Identified by Professor Greene.)

Chrysothamnus bloomeri (Gray) Greene.

One of the commonest and most widely distributed idants of the
higher slopes, where it occupies pumice soils from the bottom of the
Hudsonian zone up to several hundred feet above timberline. It was
rarely found above an altitude of 9,000 feet.

This i^lant is very long lived, and although the part above ground
is relatively small it springs from a large woody base resembling that of
the true sagebrush. The root is enormous. The main taproot (some-
times double) usually sinks so deeply into the soil that I was unable
to dig one up without breaking off the terminal point. In mature
plants the main root measures about 200 millimeters in circumference
where it enters the ground, and is 500 to 600 millimeters in length. It
gives off numerous small and slender rootlets some of which exceed
600 millimeters in length. The height of the plant above ground
rarely exceeds 150 millimeters.

Chrysothamnus bloomeri angustatus (Gray) Greene.

Common at Wagon Camp and in various parts of the Transition zone,
usually scattered through the manzauita chaparral. Vernon Bailey
collected it at Sheep Rock. (Identified by Professor Greene.)

Chrysothamnus occidentalis Greene. Rabbit Brush.

Fairly common in open spots on the lower slopes throughout the
Transition zone, where it is mixed with Kunzia tridentata, Arctostaphy-
Jos patula, and Ceanothus velutinus. This large white-stemmed spe-
cies occurs sparingly in dry soil at Sisson, and thence to Edgewood
and Shasta Valley, in the upper edge of the Upper Sonoran zone, where
it is common among the sagebrush. It was obtained at Sheep Rock
by Vernon Bailey. (Identified by Professor Greene.)

Chrysothamnus viscidiflorus Nutt.

Common in the upper part of the Upper Sonoran zone in Shasta
Valley. (Identified by Professor Greene.)

Erigeron armeriaefolium Turcz.

Rather common near timberline. Specimens were collected above
Squaw Creek, near the head of Mud Creek Canyon, and on the north
slope of Shastina. (Identified by Professor Greene.)

OCT., 1899.] PLANTS. 1()7

Erigeron compositus trifidus Hook.

Common on tlie slopes of broken shale and pnmice above tinibeiiine,
where it often grows in mats of Silene suhsdorji close nnder the edges
of rocks. On the south side of Shasta it was not observed lower than
9,200 feet, but on the cold north slope of Shastina it was found as low
as 8,900 feet. Its yellow flowers are rather conspicuous, and were
noted from the latter part of July until late in August. (Identified by
Professor Greene.)

Erigeron inornatus Gray.

This tall and much-branched Erigeron was found in Mud Creek
Canyon and along Squaw Creek, a little below the fall, at an altitude of
6,800 feet. (Identified by Professor Greene.)

Eupatorium occidentale Hook.

Collected by Vernon Bailey and Miss Wilkins on Horse Camp Trail
on the boundary between the Transition and Canadian zones. (Identi-
fied by Professor Cireene.)

Heleniastrum rivulare Greene.

Common on damp ground at Wagon Camp, on the boundary between
the Canadian and Transition zones. (Identified by Miss Eastwood.)

Hulsea nana Gray.

A common and cliaracteristic i^lant of the barren alpine slopes above
timberllne, where its showy yellow heads are conspicuous among the
bare rocks. Its leaves are sticky, crinkled, and relatively smooth. It
was found on most of the high ridges above timberline all the way
around the mountain, and was obtained on the north side of Shastina
at an altitude of 9,000 feet. Its highest limit on the southwest side, as
observed by Vernon Bailey, is 11,300 feet. (Identified by Miss East-
wood and Professor Greene.)

Hulsea larseni Gray.

Common on the pumice slopes above timberline, growing with H.
nana, from which it may be easily distinguished by the leaves, which
are covered with a dense whitish pubes(ience. (Identified by Professor
Greene.)

Madia "bolanderi Gray.

Very common in the marsh at Wagon Camp, where it was in blossom
all summer. Common also along some of the streams in the upper
part of the Transition zone, where the low temperature resulting from
the water carries narrow tongues of Canadian zone species down to alti-
tudes below their normal limits. (Identified by Miss Eastwood.)

Senecio trigonophyllus Greene.

Common along Panther Creek in the Shasta fir belt. (Identified by
Miss Eastwood and Professor Greene.)

168 NORTH AMERICAN FAUNA. [no. 16.

Senecio canus Hook.

Common on the rock-strewn ]mmice slopes above timberline. (Identi-
tied by Miss Eastwood and Professor Greene.)

Solidago elongata ]!^utt.

Abundant in open grassy places at Wagon Camp, where it was in
flower in August and September. (Identified by Professor Greene.)

Agoseris monticola Greene.^ Alpine Dandelion.

Abundant in the neighborhood of timberline on the stony slopes and
basins, where its yellow dandelion-like flowers were conspicuous from
the end of July until the middle of September. On warm southerly
slopes it was observed as high as 9,750 feet. Its wavy leaves are some-
times entire, sometimes indented or cut. This species has just been
described by Professor Greene from specimens collected by us at tim-
berline on Shasta.

Crepis intermedia Gray.

Collected by Miss Wilkins in Mud Creek Canyon near the mouth of
Clear Creek, where its zone position is either Hudsonian or Canadian.
(Identified by F. V. Coville.)

Hieracium albiflorum Hook.

Common in places along the upper part of the Transition zone. Col-
lected at Wagon Camp, on Squaw Creek, and on a warm slope in Mud
Creek Canyon near the mouth of Clear Creek. (Identified by Miss
Eastwood and Professor Greene.) A dwarf alpine form, apparently not
yet named, occurs above timberline and is fairly common above the
head of Squaw Creek, growing with H. horridirm.

Hieracium cynoglossoides nudicaule Gray.

Very common in the Transition zone Just below Wagon Camp.
(Identified by Professor Greene.)

Hieracium horridum Fries.

Common in places on the higher slopes at and above timberline,
forming small and densely hairy tufts among the rocks. The plant,
particularly when young, is completely covered by a very dense growth
of stifl" silky white hairs which give it a woolly appearance.

Hieracium gracile Hook.

Common in the heather beds just below timberline, where its conspicu-
ous yellow flowers were in blossom the eat ly part of August. (Identified
by Miss Eastwood and Professor Greene.)

Hieracium greenei Gray.

Fairly common just below Wagon Camp in the Transition zone. The
type locality of this species is in the Scott Mountains a little west of
Shasta. (Identified by Professor Greene.)

1 Pittonia, IV, p. 37, March 17, 1899.

OCT., 1899.]

PLANTS.

169

Ptiloria lactucina (Gray) Greene. {=StepJianonieria lactucina Anct.)

This curious composite, with milky Juice and conspicuous pinkish
flowers, suddenly appeared on burns near Wagon Camp the latter part
of August. It was not observed elsewhere. (Identified by Miss East-
wood and Professor Greene.)

Fig. 46.— Monument on summit of Sbasta. (Photogiapbed by W. H. Osgood.)

21753— No. 1() 22

I N D K X .

[Names of new species in black-face type. For zone position of species, see alphabetical lists,

pages 54 — 79.]

Abies amabilis, 82.

concolor lowiana. 34, 37, 138.

lasiocarpa, 82.

niagniflca, 37, 81.

nobilis, 82.

shastensis, 36-38, 138.
Accipiter atricapillus striatuhis, 111.

cooperi, 111.

velox, 111.
Acer circinatum, 153.

glabruni, 153.

macrophyllum, 153.
Achillea borealis, 164.

lanulosa, 164.
AcGxiitnm columbianuni. 146.
Actitis niacularia, 110.
.Slgialitis vocifera, 110.
Aeronautes melanoleucus, 117.
Agelains phrenicens, 122.
Agoseris nionticola, 16, 168.
Alder, 141.

])aperleaf, 141.

white tree, 141.
Alectoria fremonti, 136.
Allium sp. ?, 139.
Allium validum, 139.
Aluus incana var. virescens, 141.

rhombifolia, 141.

sinuata, 141.

tenuifolia, 141.
Amelanchier alnifolia, 150.
Ammodramus sandwichensis alaudinus, 124.
Anaphalis margaritacea, 164.
Anas bcscbas, 109.
Anorthura hiemalis pacificus, 80, 131.
Antelope, prong-horn, 102.
Antennaria alpina, 164.

geyeri, 164.

media, 164.
Anthus pensilvanicus, 130.
Antilocapra americana, 102.
Aphelocoma californica, 119.
Aplodontia, 92-93.
Aplodontia major, 92-93.

major raiuieri, 74, 81.
Apocynum pumilum, 159.
Aqnila chryaaetoa, 112.
Aquilegia truncata, 146.
Arabia, flat-pod, 146.
Arabia platysperma, 146.

Arctomys caligatus. 69.

flaviventer. 79, 80, 89.
Arctostaphyloa neyadensis, 157.

patula, 30, 157-158.

viscida, 53.
Ardea herodias, 109.
Arenaria, 80.
Arenaria compacta, 81.

jjongesta, 81.
Arnica chamissonia, 80.

longit'olia, 165.

merriami, 16, 165.

vi.scosa, 165.
Artemisia Indoviciana, 165.

rothrocki, 81.

trideutata, 165.
Asarum hartwegi, 143.
Aah. mountain, 150.

Oregon, 159.
Aspleniura tilixtVemina. 136.
Aster, Shasta alpine, 165.
Aster shastensis, 165.
Astragalinus psaltria. 124.

tristis salicamans, 124.
Aulaconininm audrogynum, 136.
Avalanches, 26-27.

Badger, 105.

Basin alopea, 52.

Basaariscus astutus rajitor, 106-107.

Bat, big-eared, 88.

California, 89.

large brown, 89.
Bear, black, 107.

grizzly, 107.
Beaver, 92.

Belltiower, twin, 163.
Betula occidentaiis, 141.
Bighorn, 103.
Bikukulla unirtora 146.
Birch, 141.
Blackberry, 150.
Blackbird, Brewer, 122.

red-winged, 122.

yellow-headed, 121.
Bluebell, Shasta, 164.
Blueberry, 159.

dwarf alpine, 158-159.
Bluebird, arctic, 133-134.

mountain, 133-134.

171

172

INDEX.

Bluebird, western, 133.

Boreal fauna of Shasta and Sierra-Cascades,

sources of, 85-86.
Boreal species :

common to Shasta and Sierra-Cascades, 73-74.
conmion to Shasta and Sierra, not known from

Cascades, 74-75.
common to Shasta and Cascades, not known

Irom Sierra, 75.
of Shasta, not known from Sierra or Cas-
cades, 75.
exclusively, of Shasta, 76-77.
Hudsonian-Alpine, of Shasta, 77-78.
Hudsonian-Alpiue common to Shasta and

Sierra-Cascades, 78.
Hudsoniau- Alpine common to Shasta and

Sierra, not known from Cascades, 78-79.
Hudsonian-Ali)ine common to Shasta and

Cascades, not known from Sierra, 79.
common to Sierra and Cascades, not known

from Shasta, 79-82.
of Sierra, not known from Shasta or Cascades,

80-81.
of Cascades, not known from Shasta or Sierra,
81-82.
Boschniakia strobilacea, 163.
Boxwood, Oregon, 1.52.
Branta canadensis, 109.
Brodiasa, 140.

Bryanthu.T empetriformis, 158.
Bubo Virginian us, 114.
Buck-brush, 153-154.
Bunting, lazuli, 127.
Burns, efi'ects of, 47.
Bush-tit, California, 132.
Buteo borealis calurus, 1 12.
swainsoni, 112.

Cacomistle, 106-107.
Callospermophilus chrysodeirus, 90.

saturatus, 69.
Calochortus maweanus, 139.

nudus, 139.
Calypte anna, 117.
Campanula scouleri, 80.

wilkinsiana, 16, 164.
Canis lestes, 103.

ochropufe, 103.
Canyons, 21-23.

Ash creek, 22, 23.

Bolam, 22.

Brewer, 22.

Diller, 22.

Mud creek, 22, 23.

Whitney, 22.

Cardamine bellidifolia pachyphylla, 146.
Carex, alpine, i:>9.
Care.x; breweri, 139.
Carpodacus cassini, 123.

mexicanus frontalis, 123.

mexieanus obscurus, 123.
Carum gairdneri, 155.
Cassiope, 80.
Castanopsis chrysophylla, 142.

chrysophylla var. minor, 142.

sempervirens, 142-143.

Castilleja atfinis, 162.

miuiata, 161.
Castor canadensis, 92.
Cat, California wild, 104.

ring-tail, 106-107.

wild, 104.
Catchfly, Alpine, 146.

Hudsonian, 145.
Cathartes aura. 111.
Catherpes mexicanus punctnlatus, 130.
Ceaiiothus, wedge-leaf, 153.
Ceanothiis eordulatus, 153.

cuneatus, 153.

integerrimus, 154.

(Cerastes) prostratus, 1.54.

velutinus, 30, 153-154.
Cedar, incense, 35, 138.
Ceoi)hloBus pileatus abieticola, 116.
Cerasus demissa, 150.

emarginata, 151.

glandulosa, 151.
Cercis occidentalis, 151.
Cercocarpus ledifolius, 148.

parvifolius, 53.
Certhia familiaris occidentalis, 131.
Cervus occidentalis, 102.
Ceryle alcyor, 114.
Chfenactis, alpine, 166.
Chainactis nevadensis, 166.
Chwtura vauxi, 117.
Chamsecyparis lawsoniana, 138.
Chama'saracha nana, 161.
Chat, long-tailed, 130.
Cheilanthes gracillima, 136.
Cheiranthus perennis, 146.
Cherry, bitter red, 151.

bush, 151.
Chickadee, chestnut-backed, 132.

mountain, 132.
Chimaphila menziesi, 156.

umbellata, 156.
Chinquapin, Sierra, 142-143.
Chipmunk, Allen, 90-91.

Klamath, 90.
Chokecherry, western, 150.
Chondestes grammacus strigatus, 125.
Chordeiles virginianus, 116.
Chrysamphora californica, 147.
Chrysopsis breweri, 81.
Chrysothamnus bloomeri, 166.

bloomeri angustatus, 166.

occidentalis, 53, 166.

visciditiorus, 53, 166.
Cinclus mexicanus, 130.
Circus hudsonius. 111.
Cistothorus palustris paludicola, 131.
Claytouia, 80.
Clintouia, 80.

Coccothraustes vespertinus montanus, 122-123.
Coccyzus americanus occidentalis, 114.
Colaptes cafer, 116.
Collomia granditiora, 160.
Columba fasciata, 111.
Columbia Gap, 69.
Columbine, red, 146.
Contopus bore tlis, 118.
richardsoni, 118.

JNUEX.

173

Cony, <.in-100.

Codt, 109.

Corallorhiza bigelovi, 141.

Comua nattalli, 156.

pubescens, 1.56.
Corviis aiuericanus, 119.
Corylus rostrata californica, 141.
Cow parsnip, 156.
Coyote, mountain, 103.

valley, 103.
Crataegus rivularis, 150.
Creeper, western brown, 131.
Crepis intermedia, 168.

nana, 80.
Crossbill, Sierra red, 12;S-124.
Crow, 119.

Clark, 110-121.
Cryptanthe geminata, 160.
Cryjitogramme acroaticboides, 136.
Cuckoo, California, 114.
Cupressus niacnabiana, 138.
Cyanocepbalus cyanocepbalus, 121.
Cyanocitta stelieri, 118.

stelleri frontalis, 74.
Cyano.spiza amcena, 127.
Cycladenia bumilis, 159.
Cymopteris terebintbinus, 156.
Cypress, Lawson, 138.
Cystopteris fragilis, 136.

Dafila acuta, 109.

Dandelion, alpine, 168.

Darlingtonia, 147.

Datura meteloides, 53.

Deer, Columbia black-tail, 101-102.

mule, 102.
Delpbinium sonnei, 146.
Dendragapus obscurus fuliginosus, 1 10-111.
Dendroica a'stiva, 128.

auduboni, 128-129.

occidentalis, 129.

townsendi, 80.
Diller Cauyon, 12.
Dipodomys californicus, 97.
Dipper, 130.
Dogwood, Oregon, 156.
Dove, mourning, 111.
Draba, Sierra alpine, 147.
Draba breweri, 147.

lemmoni, 81.
Drcsera rotundifolia, 147.
Dryobates pubescens gairdneri, 114.

villosus byloscopus, 114.
Dryoi)teris aculeata scopuliua, 136.
Duck, cinnamon teal, 109.

mallard, 109.

pintail, 109.

sboveller, 109.
Eagle, bald, 112-113.

golden, 112.
Elder, blackberry, 164.
Elk, 102.
Empidonax difficilis, 118.

bammondi, 118.

wrigbti, 118.
Epilobium, rose, 155.
Epilobium brevLstylum, 155.

Epilobium clavatum, 1

glaberrimum, 155.

bornemanni, 155.

obcordatum, 155.

oregonense, 155.

pringleanum, 155.

spicatum, 155.
Eretbizon epixantbus, 98.
Ereunetes occidentalis. 110.
Erigeron armeri.e folium, 166.

compositus trifidus, 167.

inornatus, 167.

salsuginosus, 80.
Eriodictyon glutinosuni, 53.
Eriogonum large-leaf alpiiie, 144

naked stem, 144.

small-leaf alpine, 143-144.

yellow, 143.
Eriogonum niarifolium, 143.

nudum, 144.

polypodum, 143-141.

pyrobvfolium, 144.
Erysimum asperum ]>erenne, 146.
Erythronium, 80.
Eulopbus pari.sbi, 81.
Eupatorium occidentale, 167.
Eutamias alpinus, 81.

amcenus, 90.

senex, 90-91 .

speciosus, 81.

speciosus callipeplus, 81

speciosus frater, 81.

townsendi, 81.
Everlasting, alpine, 164.

Geyer, 164.
Evernia vulpina, 130.
Evotomys, mountain, 95.
Evotomys gapperi saturatus. 69.

mazama, 95.

Falco columbarius -suckleyi, 113.

mexicanus, 113.

sparveriu.s, 113-114.
Falcon, prairie, 113.
Eeatber River or Quincy (iap, 70
Felis oregoneusis, 104.
Fincb, Cassin purple, 123.

bouse, 123.
Fir, Douglas, 34-35, 137.

Shasta, 36-39, 138.

white, 34, 37, 138.
Fire-weed, 155.
Fisher, 106.

Flicker, red-sbafted, 116.
Flycatcher, Hamnioml, 118.

olive-sided, 118.

western yellow-bellied, 118.

Wright, 118.
Forest fires, 46-47.
Forests, 30-40.

Shasta fir belt, 36-39.

white-bark pine belt, 39-46.

yellow-pine belt, 30-35.
Fox, mountain red, 103.

Townseud gray, 103-104.
Fragaria bracteata, 148.

chiloensis, 148.

174

INDEX.

Fraxiiiiis oregana, 159.
Fritillaria atropnri)Ui ea, i;!9.
Fulica araericana, 109.

Gallinago delicata, 109.
Garrya fremonti, 53.
Gaultheria myrsinites, 82.
Gayophytiim raniosissimuiu, 155.
Gentian, blue, 159.
Gentiana newberryi, 80.

simplex, 159.
Geothlypis tolmiei, 129.

triclias occidentalia, 129.
Geum, 80.

Gilia aggregata, 159-ltiu.
Ginger, wild, 143.
Glacial basins, 20-21.
Glaucidium gnoiiia caliloruicuiu, 114.
Goldfincb, Arkansas, 124.

willow, 124.
Goose, Canada, 109.
Goplier, pine- woods, 97.

Sierra pocket, 95-97.
Gosbawk, western, 111.
Grass, alpine, 139.

blue-eyed, 141.
Grass-ol-Parnassus, 147.
Grebe, pied-billed, 109.
Grosbeak, black beaded, 126.

western evening, 122-123.
Grouse, sooty, 110-111.
Gulo luscus, 79, 80, 105.

Habenaria leucostacbys, 141.

unalascbensis, 141.
Halia'etu.s leucocepbalus, 112-113.
Harpidium exannulatum, 130.
Hastingsia alba, 139.
Haw, black, 150,
Hawk, Cooper, 111.

uiarsb, 111.

sbarp-sliiuned. 111.

sparrow, 113-114.

Swainson, 112.

western I'ed-tail, 112.
Heather, red, 158.
Heleniaatriini rivulai'e, 107.
Heliotropium currassavicum, 53.
Hell-diver, 109.
Hellebore, white, 140.
Helininthopbila celata lutescens. 128.

rubricapilla gutturalis, 128.
Helodromas solitarius <',innainoiiieiis, 110.
Hemlock, black alpine, 42-46, 137.
Hemp, wild, 152.
Heracleum lanatum, 156.
Heron, great blue, 109.
Heucbera, 80.
Hieraeium alhitlorum, 168.

cynoglossoides nudicaule, 168.

gracile, 168.

greenei, 168.

horridum, 168.
Hirundo erytbrogastra, 127.
Holodiscus discolor, 148.
Horkelia pseudocapitata, 149.
Hosackia americana, 151.

Hnlsea algida, 81.

larseni, 167.

nana, 167.
Hummingbird, Anna, 117.

calliope, 117.

rufous, 117.
Hylocichla aonalaschkie auduboui, 133.

ustulata, 81.
Hypericum, dwarf, 154.
Hypericum anagalloides, 154.

Icteria virens longicauda, 130.
Icterus bullocki, 122.
Iris, 80.

Itinerary, 10-14.
Ivesia, 80.

Jay, California, 119.

Oregon, 119.

piuon, 121.

Steller, 118.
Jerboa, Sierra, 98.

valley, 99.
Junco, Sierra, 125.
Junco hyemalis connectens, 74.

hyemalis thurberi 125.
Juncus, Parry, 139.
Juncus orthophyllus, 80.

parryi, 139.
Juniper, dwarf alpine, 138.

western, .53, 138-139.
Juniperus nana, 138.

occidentalia, 138-139.

Ealmia glauca micro])hylla, 1.58.

Killdeer, 110.

Kingbird, 117.

Kingfisher, 114.

Kinglet, ruby-crowned, 132.

western golden-crowned, 132.
Klamath Gap, 69.

efficiency of, as barrier to boreal 8pecie.s, 83-84.
Kunzia tridentata, 149.

Lauius ludovicianus excubitorides, 128.

Lappula nervosa, 161.

Lark, dusky horned, 118.

Larkspur, 146.

La.sionycteris noctivagan.s, 79, 80.

Laurel, alpine, 158.

Ledum glandulosum, 81.

Lepus californicus, 101.

klaiiiatlieiisis, 16, 100-101.

nuttalli, 100.
Leucosticte, gray-crowned, 124.
Leucosticte tephrocotis, 124.

tephrocotis littoralis. 74, 81.
Leucotho'J davisia>, 81.
Libocedrus decurrens, 35, 138.
Life zones, 52-68.

Alpine, 67-68.

Canadian, 61-64.

Hudson ian, 64-67.

Transition, 54-61.

Upper Sonoran, 53-64.
Ligusticum grayi, 156.
Lilac, California, 154.

INDEX.

175

Liliuin i)arvnTn, 139-140.

waslnngtoLiianimi, 140.
Lily, tiger, 130-140.

Washington, 140.
Linnaea boreali.s. 16:!.

borealis var. longirtora, 60, 163.
Linum lewisi, 152.
Lion, mountain, 104.
Listera, 80.
Lonicera conjugali.s, 80.

involucrata, 80.
Lopliortjx californicus vallicola, 110.
Lotus aniericanus, 151.
Loxia ciirvirostra bendirei, 123-124.

ciirviro.stra minor, 74.
Lupine, dwarf alpine, 152.

dwarf lowland, 152.

silvery, 151-152.
Lupinus albicaulis sylvestris, 151.

albifrons, 152.

argenteus decumbena, 152.

elnieri, 151.

lyalli, 152.

minimus, 152.

ornatus, 151-152.
Lutkea pectinata, 149.
Lutra hudsonica, 106.
Lutreolavisou cnergumenos, 105.
Lychnis californica, 81.
Lynx californicus, 104.

canadensis, 81.

fasciatus pallescens, 10, 104.

Machaeranthera shastensis, 165.

Madia bolanderi, 167.

Magpie, 118.

Mahogany, mountain, 148.

Mallard, 109.

Manzanita, dwarf mountain, 157.

green, 157-158.
Maple, big-leaf, 153.

bush, 153.

Oregon, 153.

vine, 153.
Marmot, mountain, 89.
Marten, Pacific, 106.
Martin, western, 127.
Meadowlark. western, 122.
Megascops asio bendirei, 114.
Melanerpes formicivorus V)airdi, 116.

torquatus, 116.
Melospiza lincolni, 126.

melodia montana, 125.
Mentzelia Isevicaulis, 53, 155.
Menziesia ferruginea, 82.
Mephitis occidentalis, 105.
Merlin, black, 113.
Mertensia, 80.

Merula migratoria propinqua, 133.
Microtus (Arvicola) arvicoloides, 81.

(Chilotus) bairdi, 81.

californicus, 94.

dutcheri,81.

montanus, 95.

mordax,95
Mimulus implexus, 162.

moniliformis, 162.

Mimulus priniuloides, 162.

tiling!, 162.
Mink, Pacific, 105.
Mitella pentaudra, 147.
Moisture, ett'ects of scanty, 19-20.
Mole, California, 88.
Monardella odoratissima, 161.
Monkshood, 146.
Montia fontana, 81.
Mouse, big-eared, 94.

common white-footed, 94.

house, 93.

lemming, 95.

mountain pocket, 98.
Mud hen, 109.
Mus musculus, 93.
Mustela cauriua, 106.

pennanti, 106.
Myadestes townsendi, 132-133.
Myotis californicus, 89.

evotis, 88.

lucifugus longicrus, 89.

yumaneusis saturatus, 89.

Neotoma cinerea, 94.

fitscipes, 94.
Neurotridius gibbsi major, 16,88
New species, 16.
Nicotiana sp. ?, 53.
Kighthawk, 116.
Nucifraga columbiana, 119-12!.
Nutcracker 119-121.
Nuthatch, pigmy, 131.

red-breasted, 131.

slender-billed, 131.

Oak, black, 143.

Ochotona schisticeps, 99-100

Odocoileus columbianus, 101-102.

hemionus, 102.
Onion, large wild, 139.
Oreamnos montanus, 69.
Oreastrum alpiginum, 16.').
Oreobroma nevaden.sis, 81.

pygmaea, 81.

triphylla, 145.
Oreortyx pictus plumiferus, 110.
Oreospiza chlorura, 126.
Oriole, Bullock, 122.
Orthocarpus pilosiis, 102.
Osmorrhiza nuda, 156.
Otocoris alpestris nierrilli, 118.
Otter, 106.
Ouzel, water, 130.
Ovis canadensis, 103.
Owl, burrowing, 114.

California pigmy, 114.

California screech, 114.

great horned, 114.
Oxyria digyna, 144.

Pachystima myrsinites, 152.
PiEonia browni, 146.
Pivony, wild, 146.
Painted cup, scarlet, 161.
Parnassia californica. 147.
Parus gambeli, 132.

176

INDEX.

Paiu.s lufe.seens, 132.

Passcrella iliaca megarliynclia, 126

iliaca uuala.seliensis, 126.
Pedicularis. 80.
Peltipliylliim peltatuni, 148.
Pentstemou, yellow, 16.3.

conlertus, 1G3.

(laviilsoni, 162.

deu.stus, 163.

glaber utahensis, 163

gracilentus, 163.

inenziesi, 162.

newberryi, 163.
Perisoreiis obsciirus, 119.
Pcrogiiatlms moUipilosiis, 98.

parvus, 98.
Peroiiiy.scu8 boyli, 94.

oieas, 69.

gambeli, 94.

truei, 94.
Personnel, 15.

Petrocbelidon lunitVons, 127.
Pewee, western wood, 118.
Phacelia, dwarf alpine, 160.
Pliacelia frigida, 10, 160.

magellanica, 160.
Pbahenoptilus nuttalli. 116.
Phegopteris alpestn's, 136.
PLenacomjs oiophiliis, 95.
Philonotis, 136.
Phlox, alpine, 160.
Phlox douglasi diffu.sa. 160.
Phcebe, Say, 117.
Phyllodoce breweri, 81.

etnpetriformis, 158
Pica pica hndsonica, 118.
Picoides arcticus, 115.
Pika, 99-100.
Pigeon, band-tail, 111.
Pine, Jeffrey, 31-32.

knobeone, 33-34, 137.

lodge-pole, 38-39, 137.

nioiintain, white, 38, 136.

ponderosa, 30-32, 137.

silver, 38, 136.

sugar, 32-33, 137.

white-bark, 39-42, 137.

yellow, 30-32, 137.
Pinodrops, 157.

Pinicola enucleator calif'ornica, 81.
Pintail, 109.
Finns albicaulis, 39-42, 137.

attenuata, 33-34, 137.

balfouriana, 81.

flexilis, 81.

jeffreyi, 31-32.

lambertiaua, 32-33, 137.

monticola, 38, 136.

murrayana, 38-39,137.

ponderosa, 30-32, 137.
Pipilo fuscuscrissaiis, 126.

maculatus niegalonyx, 126.
Pipit, 130.

Piranga ludoviciana, 127.
Pitcher plant, California, 147.
Pitt River Gap, 69.
Pleurieospora fimbriolata, 157.

Plum, white, 151.
Podilymbus podicei)s, 109.
Polemouium pulcLelluni, 1(10.
Polygala cornuta. 1.'2
Polygonum, broad leaf 145.

Shasta, 144-145.
Polygonum bistortoides, 80.

newberrji, 145.

paronychia, 145

shastense, 143, 144-14.">.
Poor will. 116.

Poplar, western balsam. 141.
Populus treniuloides, 80.

trichocarpa, 141.
Porcu])ine, 98.
Porzana Carolina, 109.
Potentilla, dwarf alpine, 149.

large yellow, 150.
Potentilla flabellifoliii, 149.

glandulosa, 149, 150.

pseudorupestria, 149.
Primula, 80.

Primula suttrutescens, 81.
Procyoii psora piioificn, 16, 107.
Progne subis hesperia, 127.
Prongliorn antelope, 102.
Prunus subcordata, 151.
Psaltriijarus minimus californicua, 132.
Pseudotsuga mucronata, 34-35, 137.
Pteris a(iuiliua lanuginosa, 136.
Pterospora andromedae, 157.
Ptilorialactucina, 169.
Pulsatilla occidenfalis, 146.
Puma, Oregon, 104.
Pussj-paws, 145.
Putorius arizonensis, 106.

cicognani streatori, 81.

Masliingtoni, 69.
Pyrola pallida, 16, 156.

jiicta, 156.

secunda, 1.56.

Quail, California valley, 110.

plumed mountain, 110.
Quercus californica, 143.

chrysolepis, 142.

garryana, 143.

kelloggi, 143.

vaccinii'olia, 81, 142.

wislizeni, 143.
Querquedula cyanojitei'a, 109.
Quincy Gap, 70.

Rabbit, California jack, 101.

Klamath, 100-lUl.

sagebrush cottontail, 100.
Rabbit brush, 53, 166.
Raccoon, Pacific, 107.
Rail, sora, 109.
Raillardella, 80,
Raillardella scaposa, 81.
Ranunculus, 80.

oxynotus, 81.
Rat, bushy-tail wood, 94.

kangaroo, 97.

round-tail wood, 94.

INDEX.

177

Red-bud, 151.
Regulus calendula, 132.

satrapa olivaceus, 132.
Ueitlirodoiitonijs klainathensis, 16,93
Rharamis californica, 154.

rubra, 1.54.
Rbododendron albitiorum, 82.
Rhus trilobata, 152.
Ribes amictum, 148.

cereiim, 148.

erytbrocarinim, 82.

klamathense, 148.

lacustre, 82.

viscosis-slnnim, 148.
Robin, western, 133.
Rock slopes, 25-26.
Rose, California, 150.

wild, 150.
Rosa californica, 150

gymnocarpa, 150.
Rubus lasiococcus, 82.

nutkanus, 150.

parviflorus, 150.

vitifolius, 150.

Sagebrush, true, 53, 165.
Sagina saginoides, 145.
Salix barclayi, 80.

lasiandra, 141.

nuttalli, 141.

sitchensis, 141.
Salpinctes obsoletus, 130.
Sambucus nielanocarpa, 164.
Sandpiper, spotted, IK).

western, 110.

western solitary, 110.
Sapsucker, redbreasted, 115.

Williamson, 116.
Sarcodes sau;:uiue.i, 157.
Saxifrage, alpine rock, 147.

giant water, 148.
Saxifraga bryophora, 147.

nivalis, 80.

l)unctata, 80.

tolmiei, 147.
Sayornis saya, 117.
Scapanus alpinus, 81.

californicus, 88.
Sciuropterus alpinus klamatliensis, 92
Sciurus alboliuibatus, 91-92.

californicus, 91.

douglasi cascadenais, 81 .

fossor, 92.
Scolecophagus cyanocephalus, 122.
Scutellaria nana, 161.
Selasphorus rufus, 117.
Senecio canus, 168.

trigonophyllus, 167.
Serviceberry, 150.
Sewellel, 92-93.
Shasta, avalanches, 26, 27.

basin slopes, 52.

birds, 1U9-134.

canyons, 21-22.

eflects of burns, 47.

eflects of scanty moisture, 19-20.

forest tires, 46-47.

21753— No. ir> 23

Shasta, forests, 110-46.

general features, 17-18.

glacial basins, 20-21.

life zones, 52-68.

mammals, 87-108.

plants, 135-169.

rock slopes, 25-26.

streams, 23-24.
Sheep, mountain, 103.
Shoveller, 109.
Shrew, Monterey, 88.

Shasta. 87.

Sierra, 87.

white-bellied water-, 88.
Shrewmole, large, 88.
Shrike, white-rumped, 128.
Sialia arctica, 133-134.

mexicana occidentalis, 133.
Sibbaldia, alpine, 150.
Sibbaldia procumbens, 150.
Silene acaulis, 82.

bernardina, 81.

californica, 81

grayi, 145.

suksdorti, 140.
Siskin, pine, 124.
Sisyrinchium belluni, 141.
Sitanion cinereum, 139.
Sitta canadensis, 131.

caroliuensis aculeata, 131.

pygmaea, 131.
Skullcap, dwarf, 161.
Skunk, large, 105.

little spotted, 105.
Slope exposure, 47-51.
Slopes, basin, 52.

efl'ectof steej), 51.
Smelowskia, 80.
Smilax, California, 140.
Smilax californica, 140.
Snipe, AVilson, 109.
Snowberry, 164.

mountain, 10 ;.
Snow bush, 153.
Snow plant, 157.
Solidago elongata, 168.
Solitaire, Townsend, 132-133.
Sorbus occidentalis, 82.

sambucifolia, 150.
Sorex (Atophyrax) bendirei,8l.

moutereyensis, 88.

(Xeosorex) navigator, 88.

obscurus, 81.

shastensis, 16,87.

vagrans amcenua, 87.
Sorrel, alpine, 144.
Sparrow, Lincoln, 126.

mountain song, 125.

thick-billed, 126.

Townsend, 126.

western chipping, 125.

western lark, 125.

western savanna, 124.

white-crowned, 125.
Spatula cljpeata, 109.
Speotyto cunicularia hypogiea, 114.
Spermophilus beldingi, 81.

178

INDEX.

Speiuiophilus douglasi, 89-90.
Sphyrapicus ruber, 115.

tbjToideus, 116.
Spilogale lat ifrons, 105.
Spinus piuiis, 124.
Spirii'ii, alpine, 148.

red, 149.
Spira-a arbuscula, 82.

douglasi, 149.
Spizella socialis arizon.T?, 125.
Spraguea umbellata, 145.
Spring beauty, dwarf alpine, 145.
Spruce, Douglas, 137.
Squaw carpet, 154.
Squirrel, golden-mantled ground, 90.
Klaniatb flying, 92.
large tree, 92.
Oregon gray, 92.
Oregon ground, 89-90.
Sierra pine, 91-92.
Stachys ingrata, 161.
Stelgidopteryx serripennis, 127.
Stellaria crispa, 145.
Stellula calliope, 117.
Stephanomeria lactucina, 169.
Strawberry, large, 148.

small, 148.
Streams, 23-24.

Streptanthus orbiculatus, 147.
Streptopus, 80.

Sturnella magna neglecta, 122.
Sundew, 147.
Swallow, barn, 127.
oliti; 127.

rougb-winged, 127.
tree, 127.
violet-green, 127.
Wliite-bellied, 127.
Swift, Vans, 117.

wbite-tbroated, 117.
Symphoricarpos pilosus, 164.
racemosus, 164.

Tacbyciueta bicolor, 127.

tbalassina, 127.
Tanager, mountain, 127.
Taxideataxus, 105.
Teal, cinnamon, 109.
Thalictrum, 80.
Tbimbleberry, western, 150.
Thomomya alpiuus, 81.

mazama, 74, 81.

monticola, 95-97.

iiionticola pinetoriim, 16, 97.
Thrusb, dwarf hermit, 133.
Thryomanes bewicki spilunis, 131.
Timberline, 27-30.
Titlark, 130.

Tofieldia occidentalis, 140.
Tetanus flavipes, 110.
Towbee, California, 126.

green-tailed, 126.

spurred, 126.
Trocbilus alexandri, 117.
Troglodytes aedon parkmaui, 131.
Tritelia ixioides, 140.
Tsuga merteusiana, 42-46, 137.

Tsuga i)attoni, 42.
pattoniaua, 42.
Tyrannus verticalis, 117.

Urocyou callforiiicus toiviiseiidi, 16, 103-104.
Ursus americauus, 107.
horribilis, 107.

Vaccinium arbuscula, 159.

Cfespitosum, 158-159.

microphyllum, 82.

occidentale, 159.
Vagnera stellata, 140.
Valeriana sitcbensis, 82.
Veratruni californicum, 140.
Verbena bracteosa, 53.
Veronica cusicki, 163.
Vespertilio fuscus, 89.
Vicia americana, 152.

Viola blanda, 154 .

purpurea, 154.
Violet, alpine yellow, 154.

white, 154.
Vireo, Cassin, 128.

western warbling, 128.
Vireo gilvus swainsoni. 128.

solitarius cassini, 128.
Vole, California, 94.

mountain, 95.
Vulpcs macrourus, 103.
Vulture, turkey. 111.

"Warbler, Audubon, 128-129.

Calaveras, 128.

liermit, 129.

lutescent, 128.

Macgillivray, 129.

pileolated, 130.

western yellow-throat, 129.

yellow, 128.
Weasel, mountain, 106.

Willow, black, 141.

Nuttall, 141.

Sitkii, 141.
Willow herb, 155.
Wilsoni.i pusilla pileolata, 130.
Wolverine, 105.
Woodpecker, arctic three-toed, 115.

Cabanis hairy, 114.

California, 116.

Gairdner, 114.

Lewis, 116.

pileated, 116.

white-headed, 114-115.
Wren, canyon, 130.

Parkman, 131.

rock, 130.

tiile, 131.

Vigors, 131.

Western winter, 131.

Xanthium strumarium, 53.
Xantbocepbalus xanthocepbalns, 121.
Xeuopicus albolarvatus, 114-115.
Xerophyllum, 80.

INDEX

179

Yarrow, dwarf alpine, 164.
Tellowlegs, 110.

Zamelodia nielanocephala, 126.
Zapus niontanus, 74. 81 .

paciflcus, 99.

trinotatus, 69.

trinotatus alleni, 98.

Zeuaidura macroiira. 111.
Zones. Life, 52-68.

Alpine, 67-68.

Canadian. 61-64.

Hud.sonian, 64- 67.

Tran.sition. .54-61.

Upper Senoran, .53—54.
Zonofricliia leucopliry.s, 125.

c

B.P.L

NOV. lb ifcjyo

ii