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HARVARD UNIVERSITY

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LIBRARY

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Museum of Comparative Zoology

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University of Kansas Publications
museum of natural history

VOLUME 11 . 1958-1960

EDITORS

E. Raymond Hall, Chairman
Henry S. Fitch
Robert W. Wilson

Museum of Natural History

UNIVERSITY OF KANSAS

LAWRENCE

1960

Museum of Natural History

university of kansas

lawrence

m. ^m. zooL

Llahiih^

NOV 18 1960

H.. JO
UNiyEHSiTY

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA. KANSAS

1 960

Contents of Volume 11

CONTENTS

1. Systematic status of the colubrid snake, Leptodeira discolor Giinther. By
William E. Duellman. Pp. 1-9, 4 figures in text. July 14, 1958.

2. Natural history of the six-lined racenmner ( Cnemidophorus sexlineatus ) .
By Henry S. Fitch. Pp. 11-62, 9 figures in text. September 19, 1958.

3. Home ranges, territories, and seasonal movements of vertebrates of the
Natural History Reservation. By Henry S. Fitch. Pp. 63-326, 6 plates, 24
figures in text. December 12, 1958.

4. A new^ snake of the genus Geophis from Chihuahua, Mexico. By John M.
Legler. Pp. 327-334, 2 figures in text. January 28, 1959.

5. A new tortoise, genus Gopherus, from north-central Mexico. By John M.
Legler. Pp. 335-343, plates 7 and 8, 1 figure in text. April 24, 1959.

6. Fishes of Chautauqua, Cowley and Ellc counties, Kansas. By Artie L.
Metcalf. Pp. 345-400, 2 plates, 2 figures in text. May 6, 1959.

7. Fishes of the Big Blue River Basin, Kansas. By W. L. Minckley. Pp. 401-
442, 2 plates, 4 figures in text. May 8, 1959.

8. Birds from Coahuila, Mexico. By Emil K. Urban. Pp. 443-516. August
1, 1959.

9. Description of a new softshell turtle from the southeastern United States.
By Robert G. Webb. Pp. 517-525, 2 plates, 1 figure in text. August 14,
1959.

10. Natural history of the ornate box tvulle, Terrapene omata omata Agassiz.

By John M. Legler. Pp. 527-669, 16 plates, 29 figures in text. March 7,

1960.
Index. Pp. 671-703.

'''^'^ r«-

University of Kansas PuBLicXTroNs^i:ZiL_J

Museum of Natural History

Volume 11, No. 1, pp. 1-9, 4 figs.
July 14, 1958

Systematic Status of the Colubrid Snake,
Leptodeira discolor Gunther

BY
WILLIAM E. DUELLMAN

University of Kansas
Lawrence

University of Kansas Publications, Museum of Natxjral History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 11, No. 1, pp 1-9, 4 figs.
Published July 14, 1958

University of Kansas
Lawrence, Kansas

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA. KANSAS

tsse

27-6708

I • iUi/S.

JAN -5 1959

i-i

Systematic Status of the Colubrid Snake, -^—
Leptodeira discolor Giinther

BY

WILLIAM E. DUELLMAN

At the time of completing my study of the genus Leptodeira
(1958) I had seen no specimens of Leptodeira discolor, a species
described by Giinther in 1860 and subsequently referred to the
genus Hypsiglena by Cope (1887), Boulenger (1894), and Moc-
quard (1908), and to the genus Pseudoleptodeira by Taylor (1938).
Giinther's description was based on two syntypes ( British Museum
of Natural History numbers 1946.1.23.67 and 68) collected in
Oaxaca, Mexico, by Auguste Salle. Information concerning the
scutellation and coloration of the syntypes was provided by J. C.
Battersby; in my revisionary study {op. cit.) this information was
included in a short discussion of the species, which was referred to
incerta sedis until specimens could be examined and the relation-
ships of the species determined.

Through the courtesy of John M. Legler of the Museum of Nat-
ural History, University of Kansas, I have been able to study a
specimen of Leptodeira discolor obtained six miles southeast of
Tamazulapam, Oaxaca, Mexico, by J. R. Alcorn on June 22, 1955.
Superficial examination of the external characters of this snake
shows a striking resemblance to Leptodeira. The specimen has a
vertical pupil, divided anal, 21 scale rows, and two apical pits. The
enlarged posterior maxillary teeth are without a trace of a groove.
Examination of the hemipenis revealed that the organ was bifur-
cate and had a forked sulcus; these penial characteristics are diag-
nostic of the subfamily Xenodontinae and not the subfamily Colu-
brinae that includes the genera Hypsiglena and Leptodeira.

Examination of all available xenodontine genera indicates that
this snake belongs to a heretofore unnamed genus. In recognition
of the mental torment customarily sufiFered by workers attempting
to ascertain the relationships of the many genera of colubrid snakes,
I propose the generic name

Tantalophis, new genus

Leptodeira (in part), Giinther, Proc. Zool. Soc. London, pp. 317-318, 1860;
Garman, Bull. Essex Inst., vol. 16, p. 23, January 9, 1884; Dunn, Proc. Nat.
Acad. Sci., vol. 22, pp. 697-698, December, 1936; Duellman, Bull. Amer. Mus.
Nat. Hist., vol. 114 (1), pp. 95-96, February 24, 1958.

Hypsiglena (in part). Cope, Bull. U. S. Nat. Mus., no. 32, p. 78, 1887;
Giinther, Biologia Centrali-Americana, Reptilia, pp. 137-138, pi, 49, fig. A,

(3)

4 University of Kansas Publs., Mus. Nat Hist.

October, 1894; Boulenger, Catalogue Snakes British Museum, vol. 2, p. 211,
September 23, 1894; Mocquard, in Dumeril and Bocourt, Mission Scientifique
Mexique I'Amerique Centrale, vol. 3, p. 871, 1908; Amaral, Mem. Inst. Butan-
tan, vol. 4, p. 183, May, 1930.

Pseudoleptodeira (in part) Taylor, Univ. Kansas Sci. Bull., vol. 25, no. 15,
p. 343, June 1, 1938.

Type Species. — Leptodeira discolor Giinther, Proc. Zool. Soc. London, pp.
317-318, 1860.

Diagnosis. — A xenodontine colubrid snake having a bifurcate hemipenis with
a forked sulcus spermaticus, many longitudinal folds on basal portion, and small
spines and calyces on distal part; 12 or 13 maxillary teeth followed by short
diastema and two somewhat enlarged maxillary teeth lacking grooves; small
parotid gland; normal colubrid skull; no hypapophyses on posterior vertebrae;
elliptical pupils; two apical pits; smooth scales; normal colubrid head shields;
divided anal; paired caudals.

The generic name comes from the Greek TavraXos, a mythological char-
acter symbohc of eternal tomient, and from the Greek 80ts for snake.

Tantalophis discolor (Giinther) New comb.

The synonymy for the species is indicated in the account of the genus. The
description below of the species is based on an adult male from 6 miles south-
east of Tamazulapam, Oaxaca, Mexico (University of Kansas Museum of Nat-
ural History No. 40143).

Scutellation. — Head shields normal; upper labials 7-7 (third and fourth
entering orbit); lower labials 9-9 (1-4 in contact with anterior chin-shield,
4 and 5 in contact with posterior chin-shield); preoculars 1-1 and not in con-
tact with frontal; postoculars 2-2; temporals 1-2-3, 1-2-3; nasals divided by a
distinct groove below nostril and faint groove above; portion of rostral visible
from above, one-third length of internasals; intemasals pentagonal and one-half
as long as prefrontals; parietal suture approximately as long as frontal; ventrals
178; anal divided; caudals 80. Scales in 21 rows at midbody and showing the
following reduction:

2-f 3 (130) 8 + 9 (162)

21 19 17 (178)

2-f 3 (130) 8-f 9 (152)

Coloration. — Dorsal ground-color hght brown and extending onto edges of
ventrals; transverse body blotches numbering 50, each 1/2 to 3 scales long and
separated by light interspaces IJs to 2 scales long; blotches brownish black and
extending onto second scale row; lateral intercalary spots forming dark smudges
on rows 1 and 2. Top of head black, Hecked with tan; nape cream, followed
by dark band six scales long; dark nape stripe from posterior edges of parietals
to first dark body band. Venter cream-tan; throat and labials cream; posterior
margins of all upper labials and of lower labials 1-3 black-edged (Figure 1).

Size and Proportions. — Head and body 312 mm. long; tail 118 mm., amount-
ing to 37.8 per cent of length of head and body.

Variation. — Data on the syntypes of Leptodeira discolor furnished by J. C.
Battersby give some indication of the variation in the species. Both are males,
and they have 175 and 180 ventrals, 88 and 89 caudals, 1 preocular, 2 post-
oculars, 1-2-3 temporals, 7 and 8 upper labials, 9 lower labials. They have

CoLUBRiD Snake, Leptodeiba discolor

Fig. 1. Dorsal and lateral views of the head of Tantalophis discolor (Giinther).

(KU No. 40143). X 7.

6

University of Kansas Publs., Mus. Nat Hist.

body lengths of 365 and 402 mm., total lengths of 509 and 555 mm., tail/body
ratios of 38.0 and 39.4. They have 51 and 54 dark blotches on the body.

Skull. — The skull is t>'pically colubrid and shows no modifications. The
quadrate has both a median and a lateral depression, forming a strong lateral
flange on the anterior edge; the columellar process is elliptical, and the supra-
columellar crest is robust. The posteroinferior vomerine process extends di-
rectly posteriorly and then angles sharply posterodorsally, enclosing an eUiptical
vomerine fenestra. The lateral processes of the premaxillary are shghtly
pointed; the median spine is relatively thin and high. The pterygoids bear
23 and 21 teeth that decrease in size posteriorly; the transpalatine articulating
process of the pterygoid is rounded, not robust; the lateral crest is high and
moderately robust; the depression in the ventral surface of the pterygoid is
moderate. There are 12 and 13 maxillary teeth that increase in size posteriorly;
these are followed by a short diastema and two larger, sohd teeth. The
prediastemal teeth are slightly curved and slender. The maxillary is laterally
compressed; the posterior knob is not robust; there is one foramen in the
lateral face of the bone (Figure 2). The 10 palatine teeth are almost uniform
in size; the dentary bears 19 teeth that decrease in size posteriorly.

Fig. 2. Lateral view of the left maxillary of Tantalophis discolor ( Giinther ) .

(KU No. 40143). X 17.

A thin and otherwise small parotid gland or "venom sac" exiends pos-
teriorly from beneath the eye to about the angle of the jaw; a minute duct
connects with the anteromedian siuiace and extends to the fleshy part of the
mouth at the posterior end of the maxillary ( Figure 3 ) .

Fig. 3. Lateral view of the head of Tan-
talophis discolor (Giinther), showing the
position and relative size of the parotid
gland. (KU No. 40143). X 3.

Hemipenis. — In situ the hemipenis extends to the posterior edge of the
thirteenth caudal. The unforked part of the organ is bedecked viath nimr»erous
heavy longitudinal folds alternating with thinner folds. The basal parts of
the two heads are covered with moderate sized spines, those closest to the base
and the sulcus being the smallest. The distal parts of the heads are covered

COLUBRID SnAXE, LePTODEIRA DISCOLOR 7

with calyces. The sulcus bifurcates on the unforked part of the organ at a
point about two-thirds of the distance from the base to the division of the
organ. The sulcus is a deep groove between heavy folds proximally and is a
shallower fiurow distally (Figure 4).

uJtD

Fig. 4. Hemipenis of Tantalophis dis-
color (Giinther). The organ was cut on
the ventral surface and opened. (KU
No. 40143). X 4.

Relationships. — Using Dunn's ( 1928 ) arrangement of the Ameri-
can colubrid snakes as a guide permits the taxonomist to group
Tantalophis with several genera, some of which occur in South
America and others in the West Indies. Although the significance
of such generic characters as scale pits and nature of the hemipenis
is not clear, these characters must, of necessity, be utiHzed in at-
tempting to ascertain the relationships of Tantalophis to other
colubrid snakes. Assuming that the primary divisions of the
American colubrids into subfamilies based on the nature of the
sulcus spermaticus and the presence or absence of hypapophyses on
the posterior vertebrae have some reality, Tantalophis must be
placed in the subfamily Xenodontinae comprising genera chiefly
South American in their distribution, but with several genera in
Middle America and a few in North America and the West Indies.

8 University of Kansas Publs., Mus. Nat Hist.

In order to limit the number of genera to be compared with Tantal-
ophis, only those xenodontines having apical pits and bifurcate
hemipenis are considered. These include Cyclagras, Drepanoides,
Hypsirhynchus, laltris, Leimadophis, Pseudahlabes, Siphlophis,
Tachymenis, Tomodon, and Trypanurgos. Aside from differences in
scutellation, Leimadophis, Siphlophis, and Trypanurgos have the
heads of the hemipenes terminating in a disc, and laltris has a pli-
cate hemipenis. Tomodon has basal spines on the hemipenis. The
hemipenes of the other genera have proximal folds, distal spines, and
distal calyces, not greatly unlike the condition found in Tantalophis.
Of these, Cyclagras, Hypsirhynchus, and Pseudahlabes have round
pupils and certain differences in scutellation. Drepanoides and
Tachymenis have elliptical pupils like those of Tantalophis, but
Tachymenis has only one apical pit, and Drepanoides has one apical
pit or none. In the above characters no especially close relation-
ship between Tantalophis and any one of these genera is apparent.
If the characteristics usually employed in distinguishing and re-
lating genera are ignored and other more subjective criteria are
used, the relationships of Tantalophis still remain obscure. Of the
xenodontine genera Tantalophis approaches Leimadophis in gen-
eral physiognomy; perhaps it represents an early divergent stock of
Leimadophis that has undergone radical changes in the hemipenis
and other characters. On the other hand, if the nature of the hemi-
penis is of no importance in defining supergeneric groups of colu-
brid snakes, Tantalophis may have its relationships with Leptodeira
and Hypsiglena. Although herpetologists have been working in-
tensively on American colubrids for many decades, the relation-
ships of the majority of the groups are not well understood. Until
the hemipenes and skulls of all of the forms have been studied and
compared, and the evolutionary significance has been determined
for the characters of the hemipenes, dentition, and apical pits, our
knowledge of the relationships of these snakes will be incomplete.

Remarks. — The individual on which this paper is based is the
only specimen of the species with definite locality data. It is from
a locality six miles southeast of Tamazulapam in northwestern
Oaxaca. This town lies at an elevation of about 6500 feet in the
upper reaches of the Balsas Basin, an arid interior valley that ex-
pands in its upper end to form a broad basin of rolling and dissected
terrain ranging from about 4000 to 6800 feet in elevation. The
countryside around Tamazulapam is arid and supports plants of the
genera Prosopis, Acacia, Ipomoea, and Cassia, and also columnar
cacti. Oaks and pines grow on the limestone hills rising above the

CoLUBRiD Snake, Leptodeira discolor 9

rolling valley. Tantalophis may be endemic to the Balsas Basin, as
are many other species of reptiles. However, if the snake has its
relatives to the south in lower Central America and South America,
such a distribution seems unlikely, even for an apparent relict.

Acknowledgments. — For permission to study and report on this
specimen I am indebted to Dr. E. Raymond Hall and Mr. John M.
Legler. I am grateful to Dr. Laurence C. Stuart for many helpful
suggestions and to Dr. Norman E. Hartweg for placing at my dis-
posal the facilities of the Museum of Zoology at the University of
Michigan.

LITERATURE CITED
Amaral, a. do

1930 Estudos sobre ophidios neotropicos XVIII — Lista remissiva dos
ophidios da regiao neotropica. Mem. Inst. Butantan, 4:129-275.

BOULENGER, G. A.

1894 Catalogue of the snakes in the British Museum (Natural History).

London, 2:xi -f 382, pis. 1-20, figs. 1-25.
Cope, E. D.

1887 Catalogue of the batrachians and reptiles of Central America and

Mexico. Bull. U. S. Nat. Mus., 32:1-98.

DUELLMAN, W. E.

1958 A monographic study of the eolubrid snake genus Leptodeira.

Bull. Amer. Mus. Nat. Hist., 114:1-152, pis. 1-31, figs. 1-25, maps

1-25.
DuMERiL, A. M., and Bocourt, F.

1870-1909 fitudes sin- les reptiles. Mission scientifique au Mexique et

dan TAmerique Centrale, Recherches zoologiques. Paris, Pt. 3,

l:xiv + 1012, pis. 1-77.
Dunn, E. R.

1928 A tentative key and arrangement of the American genera of Colu-

bridae. Bull. Antivenin Inst. Amer., 2 (l):18-24.
1936 Notes on North American Leptodeira. Proc. Nat. Acad. Sci., 22

(12): 689-698.
Carman, S.

1884 The North American reptiles and batrachians. Bull. Essex Inst.,

16:1-46, 3 figs.

GUNTHER, A. C. L. G.

1860 On new reptiles and fishes from Mexico. Proc. Zool. Soc. London,
pp. 316-319.

1885-1902 Biologia Centrah-Americana. Reptilia and Batrachia. Lon-
don, pp. XX + 1-326, pis. 1-76.
Taylor, E. H.

1938 Notes pn Mexican snakes of the genus Leptodeira, with a proposal
of a new snake genus, Pseudoleptadeira. Univ. Kansas Sci. Bull.,
25:315-355, pis. 30-34, figs. 1-7.

Transmitted March 11, 1958. Contribution No. 11 from the Department of
Biology, Wayne State University, Detroit 2, Michigan.

n

27-6708

' All^-

University of Kansas Publications
Museum of Natural History

Volume 11, No. 2, pp. 11-62, 9 figs, in text, 9 tables
September 19, 1958

Natural History
of the Six-lined Racerunner
(Cnemidophorus sexlineatus)

BY

HENRY S. FITCH

University of Kansas

Lav^rence

■1958

UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY

Institutional libraries interested in publications exchange may obtain this
series by addressing the Exchange Librarian, University of Kansas Library,
Lawrence, Kansas. Copies for individuals, persons working in a particular
field of study, may be ootained by addressing instead the Museum of Natural
History, University of Kansas, Lawrence, Kansas. There is no provision for
sale of this series by the University Library which meets institutional requests,
or by the Museum of Natural History which meets the requests of individuals.
However, when individuals request copies from the Museum, 25 cents should
be included, for each separate number that is 100 pages or more in length, for
the purpose of defraying the costs of wrapping and mailing.

* An asterisk designates those numbers of which the Museum's supply (not the Library's
supply) is exhausted. Numbers published to date, in this series, are as follows:

Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950.
*Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140
figures in text. April 9, 1948.
Vol. 3. *1. The avifaima of Micronesia, its origin, evolution, and distribution. By Rol-
lin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951.
*2. A quantitative study of the nocturnal migration of birds. By George H.
Lower>', Jr. Pp. 361-472, 47 figures in text. June 29, 1951.

3. Phvlogenv of the waxwings and allied birds. By M. Dale Arvey. Pp. 473-
530, 49 figures in text, 13 tables. October 10, 1951.

4. Birds from the state of Veracruz, Mexico. Bv George H. Lowery, Jr., and
Walter W. Dalquist. Pp. 531-649, 7 figures in text, 2 tables. October 10,
1951.

Index. Pp. 651-681.
*Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31
figures in text. December 27, 1951,
Vol. 5. 1. Preliminary survey of a Paleocene faunule from the Angels Peak area. New
Mexico. By Robert W. Wilson. Pp. 1-11, 1 figure in text. February 24,
1951.

2. Two new moles (Genus Scalopus) from Mexico and Texas. By RoUin H.
Baker. Pp. 17-24. February 28, 1951.

3. Two new pocket gophers from Wyoming and Colorado. By E. Raymond
Hall and H. Gordon Montague. Pp. 25-32. February 28, 1951.

4. Mammals obtained by Dr. Curt von Wedel from the barrier beach of
Tamaulipas, Mexico. By E. Raymond Hall. Pp. 33-47, 1 figure in text.
October 1, 1951.

5. Comments on the taxonomy and geographic distribution of some North
American rabbits. By E. Raymond Hall and Keith R. Kelson. Pp. 49-58.
October 1, 1951.

6. Two new subspecies of Thomomys bottae from New Mexico and Colorado.
By Keith R. Kelson. Pp. 59-71, 1 figure in text. October 1, 1951.

7. A new subspecies of Microtus montanns from Montana and comments on
Microtus crmicaudus Miller. By E. Raymond Hall and Keith R. Kelson. Pp.
73-79. October 1, 1951.

8. A new pocket gopher (Genus Thomomys) from eastern Colorado. By E.
Raymond Hail. Pp. 81-85. October 1, 1951.

9. Mammals taken along the Alaskan Highway. By Rollin H. Baker. Pp.
87-117, 1 figiu-e in text. November 28, 1951.

*10. A synopsis of the North American Lagomorpha. By E. RajTiiond Hall. Pp.
119-202, 68 figures in text. December 15, 1951.

11. A new pocket mouse (Genus Perognftthus) from Karwas. By E. Lendell
Cockrum. Pp. 203-206. December 15, 1951.

12. Mammals from Tamaulipas, Mexico. By Rollin H. Baken Pp. 207-218.
December 15, 1951.

13. A new pocket gopher (Genus Thomomys) from Wyoming and Colorado.
By E. Raymond Hall. Pp. 219-222. December 15. 1951.

14. A new name for the Mexican red bat. By E. Raymond Hall. Pp. 223-226.
Dfcomber 15, 1951.

15. Taxonomic notes on Mexican bats of the Genus Rhogeessa. By E. Ra>Tt»ond
Hall. Pp. 227-232. April 10, 1952,

16. CfimnicnI.s on the taxonomy and geographic distribution of some North Amer-
ican woodrats (Genus Neotoma)., By Keith R. Kelson. Pp. 233-242. April
10, 1952. ^ ,'

17. The subspecies of tho Mexican red-bellied squirrel, Scinrus aurcogaster. By
Keilb R. Kelson. Pp. 243-250, 1 figure in text. April 10, 1952.

18. Geographic range of P<Tomysctis melanophrvs, with description of new siib-
spocies. By Rollin H. Baker. Pp. 251-258, 1 figure in text. May 10, 1952.

(Continued on inside of bock CQver)

University of Kansas Publications
Museum of Natural History

Volume 11, No. 2, pp. 11-62, 9 figs, in text, 9 tables
September 19, 1958

Natural History
of the Six-lined Racerunner
(Cnemidophorus sexlineatus)

BY

HENRY S. FITCH

University of Kansas

Lawrence

1958

Untversity of Kansas Publications, Musexh^ of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 11, No. 2, pp. 11-62, 9 figs, in text, 9 tables
Published September 19, 1958

University of Kansas
Lawrence, Kansas

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA. KANSAS

I Bse

Natural History
of the Six-lined Racerunner
( Cnemidophorus sexlineatus )

BY
HENRY S, FITCH

Contents

PAGE

Introduction 14

Methods 15

Relationships 17

Habitat 18

Cycles of activity and temperature relationships 20

Daily and seasonal activity 20

Effect of temperature 21

Hibernation 24

Movements 26

Burrows 28

Food habits 31

Breeding 32

Growth 39

Proportions of the tail and its regeneration 43

Molt 46

Predation and parasitism 48

Escape reactions 48

Natural enemies 51

Parasites 53

Populations 54

Summary 56

Literature cited 59

(13)

14 UNivERsiTi' OF Kansas Publs., Mus. Nat. Hist.

Introduction

Opportunity to investigate the interrelationships of many com-
mon species of the same ecological community was aflPorded the
writer in 1948 with the creation of the University of Kansas Natural
History Reservation. On this area six miles north northeast of the
University of Kansas campus at Lawrence, the species of vertebrate
animals have been investigated in an order somewhat proportional
to their abundance and conspicuousness. Since the Reservation
has been protected as a natural area, ecological succession has pro-
ceeded; species of native plants and animals that were scarce or
restricted in distribution on the area have, in some instances,
greatly increased. Concurrently, certain other species that had
thrived under the habitat conditions created by cultivation or heavy
grazing have decreased or disappeared. Among the latter species
adversely affected by the successional changes, is the six-lined race-
runner.

Belonging to a genus the species of which are primarily inhab-
itants of deserts or other arid regions, this racerunner has in the
course of its evolution, penetrated far to the north and east of most
of its congeners, into a region which, under original conditions was
chiefly deciduous forest. It exists partly in disjunct populations,
selecting mainly habitats of xeroseral aspect such as sand dunes,
beaches, and the edges of cultivated fields. On the Reservation,
deeply incised gullies in fallow fields, heavily grazed pastures, and
exposed soil and rock of a limestone quarry created favorable hab-
itat, and racerunners were numerous in localized colonies. Under
protection, after 1948, herbaceous vegetation grew luxuriantly in
the former pastures, and progressively encroached onto the barren
areas of soil and rock on the eroded gully banks and at the quarry.
In the course of this change, the racerunner's habitat shrank and
its numbers were correspondingly reduced. Because of variation
in the annual weather cycle, there were occasional reversals of the
general trend; drought delayed the course of succession and per-
mitted the racerunner to regain some of its losses, in numbers and
in area occupied. However, in the summer of 1957, rainfall was
above average, and racerunners on the area were further reduced,
to the point where few were recorded, and no significant additions
were made to the data already accumulated. With the conviction
that the species is disappearing from the area, and that future field
work there can be more profitably devoted to animals of other
kinds, I have undertaken the preparation of this report, despite the

Six-lined Racerunner

15

fact that much still remains to be discovered concerning the ecology
of the six-lined racerunner.

My study is based upon the records of 230 individual racerunners
caught, or seen and recognized in the field, a total of 581 times. I
have collected and observed racerunners in many other places in
Kansas. Also I have examined a series of 226 racerunners (mostly
from Kansas ) in the University of Kansas Natural History Museum
for certain supplementary items of information.

Credit is due to my son, John H. Fitch, and my daughter, Alice
V. Fitch, for assistance with field work, particularly in the capture
of live racerunners, which required the combined efforts of two or
more persons for best results in maneuvering the lizards into traps.

Methods

In the course of routine field work, racerunners were observed from time
to time. Some were caught when they were discovered inactive beneath
rocks. Others were caught in traps that were set primarily for animals of
other kinds, pitfalls made of gallon cans sunk in the ground at strategic loca-
tions, and especially cylindrical wire funnel traps (Fitch, 1951). In 1952
and later years attention was concentrated on a colony living along an old
diversion ditch near the headquarters of the Reservation. Many funnel traps
with several screen drift fences were installed to intercept those racerunners
that used the bare soil of the ditch banks as travelways. The traps were
checked several times daily at the season when the lizards were most active,
and occasionally a lizard that was moving near a trap was chased toward and
into it.

Fig. 1. Diagram of "corral trap," used for capture of racerunners, viewed
from above and behind. The enclosure is open on top, except for a small piece
of wire mesh at each of the two ends to prevent the lizard from climbing out.

Later, portable "corral traps" were devised, each consisting of a piece of
hardware cloth wire of )* inch mesh, about one foot wide and six feet long,
set up on one edge and bent into a bilobed enclosure. The two lobes were
compressed to narrow, angular shape, and in the apex of the V between them.

16 University of Kansas Publs., Mus. Nat. Hist.

the ends of the strip almost met, leaving a narrow entrance (Fig. 1). This
device was used with the assistance of a second person; when a racerunner
was sighted, one person kept it under observation while anotlier set up the
trap, usually at a distance of 30 to 50 feet. The edge of the wire was worked
into the loose soil, so that the lizard could not easily escape by passing beneath,
and an eight-inch square of hardware cloth was set horizontally over the end
of each lobe to hinder the lizard from climbing or jumping out. A board
several feet long and six or eight inches wide was set up on edge, projecting
from each wing of the trap as a "drift fence." Both persons then maneuvered
to the side of the racerunner opposite the enclosure, and advanced cautiously,
directing the animal's movements with slender poles, and steering it toward
the enclosure without alarming it unduly by sudden movements. When the
racerunner had progressed to a point between the two converging boards, it
could usually be driven into the enclosure by a direct approach.

Even after the quarry's entry into the enclosure, a capture was by no means
assured. Within a few seconds the racerunner's attempts to escape would be-
come frantic. It could burrow under the wire with a few rapid strokes of its
forefeet, and could make an exit even more speedily by climbing or jumping
the barrier. Almost perfect timing was required to drive the lizard into one
comer and seize it as it climbed the wire. Many of those driven into the
enclosures escaped; nevertheless the method was rather effective, and served
in the capture of a large proportion of the racerunners that were recorded. In
collecting various species of desert lizards, I have used the same device with
notable success. Important advantages of the "corral trap" method were that
the individual racerunner's normal activities were only briefly interrupted and
it was not exposed to unfavorable weather, predators, or to lack of food. Sig-
nificant records of body temperature were obtained from the lizards driven into
corral traps while those caught in other traps usually had been confined so long
that their temperatures were altered beyond their control.

Smaller numbers of racerunners were caught by hand. Most of these were
hatchlings; they permitted closer approach than did the adults, and had less
endurance. After a hatchling had been chased several times in succession from
one shelter to another it seemed to tire and become less inclined to run but
was more inclined to hide. It might permit approach to within a few feet
while skulking behind a clump of grass. Sometimes I was able to seize such
a lizard, by waiting until it momentarily closed the eye that was directed
toward me. Chances of catching a racerunner in this way were best when the
air temperature was relatively low; the animal then was reluctant to run, and
was slower than usual in its movements. Several had much lower temperatures
than that which racerunners usually maintain.

In the field I usually carried a quick-reading Schulthcis thenuometer and at
each capture of a racenmner, in a corral trap or by hand, I recorded body
temperature by inserting the bulb into the cloaca for approximately 15 seconds,
meanwhile holding the lizard through several thicknesses of cloth, to prevent
conduction of heat from my hand.

Raccnmners were permanently marked for individual recognition by clip-
ping off two toes (always on different feet) near their bases and deriving a
formula from them. The clipped toes healed rapidly and seemed to incon-
venience the raccnmners little or not at all. Probably those individuals marked
were somewhat handicapped by loss of two of their ten toes, but speed and

Six-lined Racerunner 17

agility were not noticeably affected, and it seems that the mutilation involved
in marking was not often a decisive factor in the survival of the individual.
Racerunners were marked also with black ink in the early stages of the study,
and later with bright colored enamel paint. Loss of these colors served as
an indication that a lizard had molted; also the color served to render the
individual recognizable, as each had paint on a different part, such as the
left side of the base of the tail, the right thigh, or the nape. The colors were
conspicuous for a day or more but dulled rapidly as a result of adhering of
soil, and wearing off of the paint. After a week a mark was usually so obscure
that it could not be recognized with certainty unless the lizard was in hand.
The sight records obtained from such individuals served to supplement
much more numerous records based on actual captures, with regard to move-
ments and molt. Ordinarily each racerunner captured was transferred to a
small cloth bag and brought back to the laboratory, where it was measured,
weighed, marked with paint, and toe-clipped (if it happened to be an indi-
vidual not marked previously). Those caught at the diversion ditch near the
Reservation headquarters were ordinarily returned the same day but those from
more remote study areas were of necessity most often kept a day or two in
the laboratory. The racerunners caught in 1949 were mostly released without
being brought back to the laboratory. They were weighed with spring scales
accurate to within two grams, but only those weights obtained by means of
a laboratory balance accirrate to 0.1 gm. were actually used in preparing the
following account.

Relationships

The six-lined racerunner is a member of the large family Teiidae, which
includes about 37 genera and 200 species (Darlington, 1957:216). All the
genera occur in South America, and only two (Cnemidophorus and its near
relative Ameiva) range much farther north. The South American tegu is a
giant of the family (three feet long), and its genus, Tupinambis, is known
from the Oligocene and PHocene of Argentina. Cnemidophorus is the only teiid
genus occurring north of the Neotropical region, and is known from the Pliocene
of North America. In its modern distribution the genus centers in desert
regions of the Mexican Plateau and the southwestern United States. Of ap-
proximately 20 species in this genus, the six-lined racerunner is the most
northern in its over-all range although a widespread, desert living, western
species, C. tigris, attains a latitude in central Oregon approximately the same
as that reached by C. sexlineatus in Minnesota and Wisconsin. Of live natural
species-groups within the genus, recognized in the most recent revision (Burt,
1931), the sexlineatus group is one of the most advanced, characterized by
peculiarities in the arrangement of the cephalic shields, and by the stripes on
back and sides; it ranges throughout Guatemala, most of Mexico, and the
central and southern United States, except the far West. C. sacki, vdth many
subspecies, occupying most of Mexico and much of the southwestern United
States, is the central species of this group, and may most closely resemble the
ancestral type from which the others have been derived. C. sacki differs from
sexlineatus in larger size, more sluggish behavior, and spotted pattern, with
partial loss of stripes. C. sacki octolineatus of Arizona, New Mexico, and
adjacent regions shows no differences between the sexes in coloration or bodily

18 University of Kansas Publs., Mus. Nat. Hist.

proportions whereas in C. sacki gularis of Texas there is striking difference
between the sexes. The male of gularis has an orange throat and deep blue
on the belly in contrast to the cream color of the ventral surface of the female.
C s. gularis seems to be the nearest relative of C. sexlineatus and the ranges
of the two overlap in appro.ximately the eastern half of Te.xas and much of
Oklahoma.

Habitat

The habitat of the six-lined racerunner has been amply described by earlier
writers. Burt (1931:91-92) quoted the statements of nearly all those authors
whose publications preceded his own. Of eleven authors, quoted by Burt
who described the habitat in ten different states representative of the entire
range, eight mentioned sandy situations which seemed to be the chief feature
that habitats had in common, although the racerunner is not invariably con-
fined to places where there is sand. Three of the authors alluded to high or
hilly habitat, two described the habitat as dry, two mentioned plowed fields,
and two mentioned rocky hills. One statement contrasting with the general
trend of all the others was that of Strecker (1915:24) describing the species
as ". . . partial to wooded bottom lands" in the southwestern part of its
range near Waco, Texas, although he also described the species as common in
cutover pine woods (which presumably were sandy uplands) in the more
eastern part of the state.

The following briefly quoted statements, from more recent authors, describe
the habitat in 13 states, representative of the entire geographic range. Coastal
Virginia: ". . . along sandy roads and in borders of fields . . .", Rich-
mond and Coin (1938:308). Eastern North Carolina: ". . . beach drift
. . . on coastal banks . . . isolated sandy areas", Robertson and Tyson
(1950:136). South Carolina: ". . . open and grassy portions of the dry
pinewoods . . .", Malnate (1944:729). Florida: ". . . not entirely
restricted to xeric situations . . . plowed up in deep damp muck . . .
wet sand under logs on lake beaches", Carr ( 1940:74). Indiana: ". . . dry,
almost arid, habitat . . . sand dune country . . . crests of the Bald
Knobs . . .", Minton (1944:449). Mammoth Cave region of Kentucky:
". . paths and in old fields where there are clearings", Hibbard ( 1936:

280). Western Tennessee: ". . . in both uplands and river bottoms,
usually near forests or thickets . . . sandy and gravelly regions . . .",
Parker (1948:25). Mississippi: "dry, sandy or gravelly soil . . . com
and cotton fields . . ." Cook (1943:13). Southeastern Minnesota and
adjacent Wisconsin at the northernmost stations of occurrence: "dry hillsides
. . . cinder fills along railroad . . . steep gravel sides of a ravine
with red cedar and sandburs . . . sand beach of the river at the foot of
a steep sand and gravel bluff", Breckenridge (1943:95). Nebraska: ". . .
dry, sandy places . . .", Hudson (1942:40). Southeastern Missouri:
". . . dry, open woods and glades . . .", Boyer and Heinze ( 1934:192).
Southeastern Oklahoma: ". . . weeds along the roads, in open fields, along
the fence rows, and along the edges, and in open glades of the forest." Trow-
bridge (1937:294). Lake Texoma area in Oklahoma and northern Texas:
. well drained areas . . free of timber . . . especially sand

dunes", Bonn and McCarley (1953:469).

Burt (1931:92) described the habitat in Kansas as including rocky hillsides,

Six-lined Racerunner 19

fields cultivated for corn and wheat, upland meadows, sandy riverbanks, chalk
cliffs, railroad embankments, road beds, sand dunes, rock outcroppings, and
wooded hillsides. Burt indicated that in Kansas the racerunner occupies a
greater variety of habitats than does any other kind of lizard, and speculated
that a high moisture content of the soil surface was the chief limiting factor
as the species was rare in places where the soil was loamy. Twenty-five years
later, Smith (1956:206) made the following statement regarding the habitat
in Kansas: "The habitat is in relatively dry areas, where sandy or other loose
soil, short grass, and other kinds of low vegetation occur. The chief require-
ment seems to be a certain degree of dryness. The land may be flat or hilly,
and the soil coarse or fine." Near Fort Leavenworth, in the northeastern
comer of Kansas, Brumwell (1951:204) found the racerunner to be the most
abundant lizard, living in woodland, especially about rotted logs and rock
ledges.

My own observations indicate somewhat different habitat preferences in
different parts of the state. In the Chautauqua HUls area representing ap-
proximately the westernmost extent of the original deciduous forests, and
located about % of the distance east across the state from the western boundary,
and in the southern one-third, the species is especially abundant. Here the
rocky woodlands support mostly scrubby oaks, especially post oak and black-
jack oak, and underbrush is sparse. Farther east and north, in more mesic
types of woodlands, with more continuous leaf canopy, and more underbrush
shading the ground, the racerunner is confined to open situations. The race-
runner does not thrive in a typical prairie habitat, either of tall-grass type or
of short-grass. Where it occurs in prairies, it requires patches of open ground
such as are produced by excessive grazing or trampUng (especially about salt-
licks or watering places of livestock) or such as occur naturally, along eroded
gully banks and stream edges, or on slopes and hiUtops where soil is shallow
and rocky, and vegetation is sparse.

In northeastern Kansas I have found the racerunner to be most abundant
in sandy situations, and also fairly nmnerous about heavily grazed rocky pas-
tures. Often it is found in cultivated fields but usually near their edges. The
species is abundant in sandy lowland areas near the Kansas River. In the great
flood of July, 1951, when the river's floodplain was entirely inundated, the
population of racerunners was mostly destroyed, and they did not recover
their former numbers for several years, despite the favorable habitat conditions
created by extensive sandy deposits. In May, 1957, I found racerunners abun-
dant along the south bank of the Kansas River a few miles west of Lawrence,
in Douglas County. Here groves of young cottonwoods and thickets of willow
had sprung up since the flood, creating an abundance of shelter. About the
razed remains of an old building destroyed in the flood, the Hzards were espe-
cially numerous, and when disturbed, they sought shelter beneath the piled
debris of boards and shingles. The sand there was riddled with their burrows.
Often several racerunners were in sight at the same moment.

On the Reservation in 1948, racerunners were localized, and occupied only
a small part of the square mile. None was seen in most of the cultivated
fields. They were mostly in rocky and brushy parts of the heavily grazed
pastures, and occasionally appeared on roads and gully banks. Some were
present in a much eroded upland field, which had been fallow for several
years.

20 UMVERsrri' of Kansas Publs., Mus. Nat. Hist.

Cycles of Activity and Temperature Relationships
Daily and seasonal activity

Various observers have noted that raceninners are not uniformly
active throughout the day, but have periods of relatively concen-
trated activity in the open and spend a greater part of the time in
retirement. Taylor (1935:211) wrote that in Arkansas near Devall
Bluff, in July, racerunners emerged daily about 11 A. M. to begin
feeding, and disappeared into their burrows about 3 P. M, Oliver
(1955:133) UTOte that in Florida in April the racerunner becomes
active about 9:45 A. M. and remains more or less active until
about 4 P. M., but that late in summer when midday temperatures
are higher, it is abroad only in early morning and late afternoon.

Milstead (1957:111) studying four other species of Cnemido-
phorus (perplextis, sacki, tessellatiis, and tigris) in western Texas,
found that in summer the daily activity began between 7:00 and
9:00 A.M., continued until from 11:00 A.M. to 1:00 P.M., and
was resumed near 5:00 P. M. He concluded that soil temperature
was more critical than air temperature; the lizards were active at
soil temperatures of between 29° and 50° C.

By means of laboratory experimentation, Barden (1942) deter-
mined that under normal conditions of light and temperature, race-
runners attained a peak of activity between 10 and 11 A. M. When
kept in complete darkness (at 85°F) or in constant artificial light,
the lizards tended to retain the same cycle but this could be altered
by reversed illumination. Technique in this study involved sus-
pending the racerunners' cage from a sensitive brass spring, so that
movements of the cage were recorded on a kymograph.

In my own experience on the Reservation, the daily schedule of
racerunners is somewhat variable according to the individual, the
season, and especially the temperature. On some warm summer
days most of the population is active by 8 A. M., and by noon most
have returned to their burrows. In mid-afternoon, the hottest part
of the day, activity was usually at low ebb, and the few individuals
that remained active above ground kept to shaded spots and travelled
little. At temperatures higher than 90° F, when the sun was shining
brightly, the soil surface was probably too hot to permit the lizards
to forage comfortably; they were usually seen at somewhat lower
air temperatures. On cloudy days the racerunners did not emerge
at the usual time, but might remain in their burrows all day, or
might emerge, hours later than usual, whenever the sun was shin-
ing. For 25 individuals that I caught by hand, time of capture was
rather evenly distributed from 8 A. M. to 3 P. M.

Six-lined Racerunner

21

Opportunity and effort to obtain records o£ racerunners varied
considerably during the course of the study. Nevertheless, the
monthly totals in Table 1 show well defined trends of increasing
and decreasing activity in the course of the growing season. Only
a few records were obtained in April, but activity rapidly increased
in May to a peak in June, then abruptly fell off in July, August and
September, with only an occasional record in October (Fig. 2).

o
cc
o
o

UJ

o

01
Ul
CD

=5

250

,'. CAPTURES

; I
1 \
1 \

200

I <

/ \

150

/ ..\ INDIVIDUALS

100 /

~ /
/
/
/
/

50 /

\ \ -

\\
— 1 1 1 1 1-^

APR MAY JUN JUL AUG SEP OCT

Fig. 2. Seasonal incidence of rec-
ords for racerunners on the University
of Kansas Natural History Reservation.
Combined data for nine consecutive
years are represented. Activity begins
in late spring, reaches a peak in June,
and then declines rapidly.

Effect of Temperature
Bogert ( 1949 ) originally called attention to the fact that lizards,
in the course of tlieir normal activities usually maintain high and
fairly uniform body temperatures, the preferred level being char-
acteristic of the species, with differences between genera and fam-
ilies. Of various lizards studied by Bogert, the racerunner was
found to maintain temperatures higher than those of any other kind.
Bogert found that in Cnemidophorus sexlineatus of Florida, the
optimum body temperature and temperature range were both sim-
ilar to those of C. tigris Hving on the Arizona desert under much
different habitat conditions. Both species were found to have pre-
ferred body temperatures several degrees higher than the species

22

University of Kansas Publs., Mus. Nat. Hist.

of Sceloporus (S. woodi and S. magister, respectively) that shared
their habitats.

Of the one hundred body temperatures that I recorded ( Fig. 3 ) ,
three-fourths were in the five-degree range, 38° to 42°. Racerun-
ners will tolerate a range some 5" below the optimum before they
seek shelter, but will tolerate only about 2° above it. Milstead ( op.
ct^: 112) found that in four other species of Cnemidophorus in
Texas, body temperature in the normal range of activity varied
between 36° and 40°.

15 20 25 30 35 40 45
DEGREES CENTIGRADE

Fig. 3. Air temperatures at which
records of active racerunners were
obtained. The records are distrib-
uted over a 20-degree range, but are
concentrated between 33° and 35°C.

The lowest air temperature at which I found a racerunner in the
open was 20.4°; the lizard, chased into a trap, was found to have a
body temperature of 37.1° C. At the same time another found
already in a trap but basking in the sunshine, had a body tempera-
ture of 35.3°. Early in the afternoon of June 1, 1956, at an air
temperature of 21.5°, three racerunners, caught while they were
active, had body temperatures of 37.0°, 38.6°, and 39.6° C. On May
26, 1956, an adult female, which had just sought shelter under a
board after the sun had disappeared behind a cloud, was found to
have a temperature of 36.4° when the air was 29.0° C. On June 1,

Sec-lined Racehunner

23

1955, when the sky was overcast (air 25.0% soil surface 28.2° C) only
one raceninner could be found active along the diversion ditch
where many usually could be seen at times favorable for their ac-
tivity. The lone racerunner was noticeably slower than usual in its

30 35 40 45

DEGREES CENTIGRADE

Fig. 4. Body temperatures of ac-
tive racerunners; the range is narrow
and the preferred level is high, be-
tween 40° and 41°C.

movements because of low temperature; nevertheless it easily
avoided capture by hand. On June 5, 1954, a recently caught adult
male was weighed and measured in the laboratory at a body tem-
perature of 26.5°C. He was able to run and dodge with some agility
when released on a table top, but was so much slower than normally
active racerunners that he was easily recaptured. As I reached for
him, he turned and snapped at my fingers. On May 4, 1955, an
adult caught the day before and kept overnight in the laboratory
was found to have a body temperature of 19.5° when handled.
It was stiflF and slow, although it struggled vigorously. When
released on a table, it could not rise up on its legs to run, but pro-
gressed with a sprawling gait and was easily recaptured. On Sep-
tember 23, 1952, a juvenile was weighed and measured in the labora-

24 UNrv'ERSiTY OF Kansas Publs., Mus. Nat. Hist.

tory at a temperature of 14°. When released on a table, it tried to
run, dragging itself with stiff, heavy movements, and after it had
progressed a few inches and bumped into the wall, it subsided into
quiescence.

In 1955 racerunners became active in a warm period of late April
and early May. Nevertheless, in the second week of May, with the
return of somewhat lower temperatures, none was seen; even tem-
peratures in the low seventies (F) were too cold for the lizards to
emerge, despite bright sunshine.

Hibernation

Feeding and reproductive activity are concentrated in late spring
and early summer. After July, activity wanes and the adult race-
runners spend more time in their burrows and are seen with de-
creasing frequency; they are rarely seen in September even though
summer weather may still prevail. Whether such racerunners
emerge regularly each day, even for brief periods of activity, seems
doubtful. They probably burrow to depths where temperatures
are fairly stable, and far below those that they require for full
activity, then lapse into a semitorpid state, which passes into true
dormancy with the onset of winter. Because the racerunner's lower
limits of temperature for various activities such as locomotion,
feeding, and mating, are all relatively high, the species emerges
relatively late in spring. On the Reservation it was consistently
the last species of common reptile to appear each spring. Cook
(1943:13) wrote that in southern Mississippi racerunners seldom
emerge from hibernation before the first of April, whereas in the
northeastern part of the state they usually are not in evidence until
mid-April. In northern Florida the racerunner's hibernation begins
in October and ends in March or April (Oliver, 1955:118). Neill
(1948:109) stated that in Richmond County, Georgia, tlie racerun-
ner remains active late in the autumn, but is the last species of
reptile to emerge in summer. Neill described hibernation burrows,
which usually end beneath a large rock. The lizards found in these
hibernation dens were sometimes single and sometimes in twos
and threes. The hibernating racerunners were able only to writhe
and gape when uncovered. Carr (1940:74) discussing habits of
the species in Florida, wrote: "They hibernate under six or eight
inches of sandy soil; two specimens have been found in gopher
[turtle] holes." In Indiana, Minton (1944:449) found that race-
runners seldom emerge before mid-May and retire to hibernation

Six-lined Racerunner

25

in early September. Burt (1933:191) recorded finding three race-
runners ( two adults and one juvenile ) with a collared lizard ( Cro-
taphytus collaris), all hibernating together under a deeply em-
bedded large rock, in Cowley County, Kansas, on March 6, 1933.
These lizards were completely dormant and immobile.

No hibernating racerunners were found in my study, but in each
of several years specimens were found in spring under deeply em-
bedded rocks before any were found in the open. Such racerunners
were in distinct nest cavities and often were in contact with the
lower side of the rock, and were warm to the touch as a result of
absorbed heat. The cavities occupied had no evident tunnels lead-
ing either to the outside or to possible hibernation sites at deeper
levels, but it seemed most probable that the lizards had not yet
emerged, having merely worked their way up from a lower level,
compacting the soil as they progressed, until attaining contact with
the comfortable warmth of the rock's lower surface.

Over a seven-year period. May 1 was the average date for earliest
record of a racerunner in the open (Table 1), and the average

Table 1. — Year by Year Record of Events in the Annual Cycle of the
Six-lined Racerunner on the University of Kansas Natural History
Reservation.

Earliest records

First
recorded
hatchling

Last

recorded

adult

Last

Year

Under
shelter

In

open

recorded
hatchling

1949

August 13
September 3
September 13
August 13
August 9
August 4

September 19
September 1

1950
1951

April 14

April 26
May 2
April 29
May 8
May 6
April 21
April 21

October 10

1952

September 16
September 21
September 18

September 21

1 QF)^

May 3
April 24
April 18

1954
1955

October 3

1956

August 9

1957

May 10

deviation from this date was a little less than six days. For five dif-
ferent years of late season records for adults, September 15 was
the average date, with an average deviation of five days. At the
latitude of the Reservation the length of the growing season for the
racerunner is thus about 4/2 months, but for an individual it prob-
ably averages somewhat shorter. Approximately two-thirds of the
entire year is spent in hibernation.

26 University of Kansas Publs., Mus. Nat. Hist.

Mo\TMENTS

The only published statement knowTi to me concerning home
range in the six-lined racerunner is that of Oliver (1955:112), evi-
dently based on his original observations in Florida: ". . . does
not appear to maintain a definite territory, but wanders over a
fairly large home range of 200 to 300 feet in diameter, perhaps
more." In a much quoted statement concerning speed of the race-
runner, Hoyt (1941:180) related an instance of one that ran on the
road in front of a truck, gaining and then maintaining a speed of 18
miles per hour, for a period of a little more than a minute. It must
have travelled for approximately % mile in the same direction dur-
ing the observation, but there is no reason for believing that this
movement was entirely within the normal home range. The speed
attained by this individual is possibly the maximum actually re-
corded for any reptile, although a few desert inhabiting lizards
are generally supposed to be even swifter.

In the course of my study, 85 of the racerunners that were marked
were subsequently recorded at different locations, affording in-
formation on their movements; 50 were recaptured only in the
same season, 31 others after the lapse of one hibernation, four in
three different seasons, and one in four different seasons. In some
instances span of individual records was longer than these figures
indicate, since a marked racerunner might be missed for an entire
growing season, or even two, and then recaptured subsequently.

Because of the nature of the terrain on the Reservation, the race-
runners recorded by me usually did not move freely in all direc-
tions, but tended to have their movements channeled linearly along
rock outcrops or the barren banks of gullies or ditches. The shape
of the home range was hence long and narrow, and the size may
have been considerably different from that in locations where fa-
vorable habitat features were more generally distributed.

As a typical example of the movements recorded for the marked
racerunners, a male first recorded on the ditch banks on May 20,
1955, still well below average adult size, was recorded two more
times in the following month and on 23 additional occasions in late
May, June and early July of 1956. All the 26 records were along
the ditch; 16 including the first and last were at the same central
location. Frequency of capture at this particular spot perhaps
resulted not so much from the racerunner's preference as from the
fact that a screen drift fence extended diagonally across the ditch

Six-lined Racerunner 27

bottom, with funnel traps facing in either direction, which were
fairly eflFective in intercepting travel. With respect to this central
location, the other ten in the order they were recorded, were: 280
feet east, 30 feet west, 70 feet east, 305 feet east, 240 feet west, 40
feet east, 70 feet west, and 260 feet east. On several occasions
within a few minutes after being released from a trap, this race-
runner was seen in one of the more remote parts of the home range;
hence it was obvious that the entire area was covered frequently.
The ditch extended in both directions beyond the lizard's known
range, and this individual probably would have been either trapped
or observed at more remote locations if it had ranged consistently in
those directions. The 545 foot stretch of ditch along which it was
recorded probably was almost the full length of the home range,
and the width of the area with bare soil or sparse vegetation used
by the racerunners along this ditch averaged approximately 25
feet; hence the computed home range was 13,625 square feet, or .31
acre.

No other racerunners were caught so many times or at so many
different locations as this one. Only a few were recorded along
such a long segment of the ditch. Presumably in most instances
the data are inadequate to show the full extent of the home ranges
of the individuals involved. The racerunners that frequented the
ditch probably had home ranges that were limited to this area,
because nearby areas were, in general, not suitable habitat.

If recorded locations for an individual were distributed at random
along the length of the elongate home range, any two records would
be separated, on the average, by a distance equivalent to approxi-
mately 35 per cent of the area's full length. Even though indi-
vidual racerunners were able to cover their whole ranges within a
few minutes, and were not dependent on home bases, they tended
to concentrate their activities at certain places where conditions
were especially favorable for feeding, basking, or other essential
functions. Traps were not randomly distributed either, and a dis-
proportionately large number of captures were made at a few espe-
cially strategic locations. For 188 recorded movements of male
racerunners between successive captures (or sight records), dis-
tance averaged 125 feet. The comparable figure for 102 movements
of females was 171.3 feet. These figures exclude several excep-
tionally long movements between the hilltop quarry and the ditch
at the base of the hill, which seem to represent shifts from one home
range to another. Home range lengths along the ditch of 343 feet

28 University of Kansas Publs., Mus. Nat. Hist.

for males and 440 feet for females are indicated, with actual areas
of approximately .20 acre and .25 acre.

Home ranges of these sizes seem remarkably small for an animal
that moves so rapidly and frequently as the racerunner. Milstead
(1957:116-117) measured a home range of .53 acre for a marked
individual of the desert living Cnemidophorus tigris for which 17
recoveries were recorded. Excluding one outlying point of re-
covery, this lizard's recorded locations all fell within an area of .26
acre. Milstead concluded that the home range may be divided
into an area of greater activity and a contiguous area of lesser
activity. My records of the six-lined racerunner also show a defi-
nite tendency to cluster in one part of the home range for most
individuals.

In general, the conditions of my study were not well adapted for
showing the longer movements made by racerunners, including
exploration, possible migration, and shift from one home range to
another. However, in the drought years of 1953, 1954, and 1955,
when habitat conditions became more favorable for racerunners
than in the several preceding years, the lizards appeared abruptly
in places where they had not been seen before — in places in 1950
and 1951 where vegetation had been too dense for them. This
prompt exploitation of newly available habitat demonstrated the
potential vagility of the species. Six of the marked racerunners
(three of each sex) were recorded to make fairly long movements
that must have involved shifts in range; in every instance the shift
was between the quarry and the ditch. For males distances in feet
were: 700, 600 (and back again in the same season), and 1170. For
females distances in feet were: 750, 750 (and back again in the same
season), and 900.

The slope between the quarry and the ditch is woodland with
brushy tangles. Little sunlight reaches the ground, and the habitat
is unfavorable for racerunners. However, on a few occasions the
lizards were seen on a trail that led from the ditch to the quarry.

Burrows

The six-lined racerunner is well equipped for digging, having
well developed and powerful forelegs with long, powerful claws,
and spends much time underground in its burrows, even at the
season of greatest activity. The burrows have several important
functions. They provide nocturnal shelters (as these lizards are
strictly diurnal and heliophilic), and are insulated retreats that are

Six-lined Racerunner 29

used at any time when the Hzard is inactive because of unfavor-
able weather conditions. For example, the burrow's shelter is
sought at times when air and soil temperatures are so high that
the racerunner cannot readily hold down its body temperature to
the preferred level of around 40°C, since temperatures only slightly
higher would be lethal. Also, in times of emergency when the lizard
is hard-pressed by a pursuer despite its fleet and elusive running,
a burrow constitutes the final refuge.

Where burrows of other small animals, such as mice, voles, or
moles are abundant, the racerunner utihzes them and needs to do
less digging than in places where such ready-made shelters are
lacking. Wliere there are flat rocks, the racerunner digs beneath
them and at the end of a short tunnel it excavates a disk-shaped
cavity, just large enough to squeeze into when the long tail is
curled up. Where rocks and burrows of mammals are absent, the
racerunner does much digging. Burt (1928:43) stated that the
burrows are used repeatedly. Boyer and Heinze (1934:192) de-
scribed the burrows as forming a network of runways in dry open
woods in southeastern Missouri in places where racerunners were
abundant. Blanchard (1922:6) describing the habits of the race-
runner in sandy areas of western Tennessee, stated: "The burrow
is short and has two openings, and when the lizard is inside, one of
these openings is partially filled with sand thrown out from within."
Cook (1943:13) stated that in Mississippi the racerunner makes
free use of mole runs in old abandoned fields. Edgren (1955:141)
described the short, shallow tubular burrows that are used in the
sand dune region of northern Illinois to escape unfavorably high
surface temperatures. Taylor (1935) described the burrows of
racerunners where the species was abundant in a cotton field near
Devall Bluff, Arkansas; from two to four tunnels would meet in a
somewhat enlarged cavity usually eight to ten cm. below the sur-
face (maximum depth 14 cm.). Pearson and Nelson (1952:188)
examined 40 burrows of the oldfield mouse (Peromyscus polionottis)
in Marion County, Florida, and found that at least one-third had
recent tracks of racerunners leading into the entrances. The lizards
were found to have excavated lateral tunnels of their own, which
led nearly to the surface and thus could serve as emergency exits.

At the old diversion ditch where my own observations were most
concentrated the racerunners while foraging kept mostly to the
nearly barren soil of the ditch bottom and banks, but when alarmed,
they would retreat to nearby sheltering vegetation. Burrows and

30 Universit\' of Kansas Publs., Mus. Nat. Hist.

surface runways of the prairie vole (Microtus ochrogaster) were
numerous in the grass beyond the bank, and racerunners that were
chased often took refuge in them. Also, they often escaped into
burrows that were in the more barren areas. Some of these were
old burrows of indeterminate origin, but most were probably made
originally by the mole (Scalopus aquaticus). The mole does not
leave open surface entrances; its tunnels become accessible when
the layer of soil roofing them collapses, or when the lizards dig into
them. In its foraging the mole excavates and soon deserts exten-
sive tunnel systems, which then become available to the racerunner
and many other small animals. Being a voracious predator, the
mole would almost certainly kill and eat any racerunners it en-
countered in passing through its runways or digging new tunnels.
Other small digging insectivores (Blarina brevicauda), mice (Per-
omyscus and Reithrodontomys) and even voles (Microtus) might
sometimes prey on racerunners found in this manner at times when
they were too sluggish to escape quickly.

Besides the old burrows of moles and other small mammals, there
were many burrows on the ditch bank that were obviously exca-
vated by the racerunners themselves. These burrows were of diam-
eters only shghtly greater than the lizards, and were concentrated
in bare soil in what seemed to be the hottest and driest situations
that could be found. They were on the sunny north bank of the
ditch, and none was ever noted on the south bank, which was
shaded throughout most of the day. Often burrows were noted to
be plugged from within with finely pulverized soil. At times when
I attempted to dig out such burrows, the lizard inside usually burst
through the surface crust a few inches away and escaped. Ordi-
narily most of the burrows were not plugged and those dug out
usually were unoccupied. Probably a racerunner frequently digs
new burrows, and its attachment to any one burrow probably is
temporary.

A hole is defended by the occupant against trespassers. On Sep-
tember 9, 1953, in a brief cloudy period, I saw a juvenile enter a
burrow beneath a flat rock, and almost instantly it darted out again
chased by another of about the same size. Within a few seconds
the pursuer returned and re-entered its burrow.

In the summer of 1957 several racerunners were kept in a circular
enclosure ten feet in diameter, with an 18-inch wall of sheet metal,
buried in the ground for several inches. The lizards excavated
many burrows in a sand pile at the center, and a few among rocks
near the wall. Each individual seemed to have its own burrow.

Six-lined Racerunner 31

Food Habits

Burt (1928:58-61) summarized early published literature (Hol-
brook, 1882; Cope, 1866; Hartman, 1906; Ruthven, 1907; Ditmars,
1915; Taylor, 1916) concerning the racenmner's food, along with
his own observations on captives offered a variety of foods, and his
analysis of contents of 15 stomachs of specimens from Kansas. Dit-
mar's statement that "Adults are not adverse to feasting on eggs of
small birds that nest on the ground" was rightly questioned by
Burt, as the smallest eggs of ground nesting birds are much larger
than items normally eaten by the racerunner. Other early authors
referred to the racerunner's insectivorous habits, and its swiftness
in securing such agile prey as flies, tiger beetles, cockroaches,
spiders, and grasshoppers. Burt listed many types of insects ob-
tained with a sweep net and offered to captive racerunners, that
were eaten with relish. Insects that were rejected included lady-
bird beetles, large butterflies, and ground beetles (unless their
elytra were removed). The 15 stomachs that Burt examined con-
tained: grasshoppers and crickets {Melanoplus, Nomotettix, grylHd,
tettigoniid) 41.5%; spiders (salticids, lycosids, etc.) 24.5%; lepidop-
terans (9 adult moths, 2 pupae and 12 caterpillars) 19.9%; snails
{Polygyra, pupillids) 7.0%; beetles (Sitonia hispidulus, Agonoderis
obliqulus, Arma, Baris) 2.5%; ants and wasps (Ponera, Cremato-
gaster, Formica, Chalcis, Sigalphtis) 1.9%; flies (tachinids) .5%.

Boyer and Heinze (1934:192) mentioned a racerunner that dis-
gorged a cutworm, and Breckenridge (1943:96) mentioned one
that contained a medium-sized grasshopper. Force examined
stomachs of 29 specimens from Okmulgee County, Oklahoma.
Eleven had no food. In the remaining 18, food was found in the
following percentages: locustids 52, arachnids 15, myrmicids and
formicids 16, elaterids 3.5, unidentified larvae 9, mayfly and mos-
quito larvae each 1. Hudson (1942:40) recorded that five from
Nebraska had eaten spiders, grasshoppers, beetles and their larvae,
and bugs.

Racerunners that were foraging moved about jerkily, probing
with the long snout beneath objects and into holes and crevices,
as if guided partly by olfactory sense. Stebbins (1948:199) in his
study of nasal structure in lizards, examined specimens of another
Cnemidophorus species ("tesselatus" = tigris) , and finding the
nasal chamber completely lined with olfactory epithelium, con-
cluded that, on a structural basis, members of this genus would
appear to have a keen sense of smell. That these Hzards depend

32 University of Kansas Publs., Mus. Nat. Hist.

more on scent and less on sight than do some others in finding their
prey is evidenced by the taking of such immobile prey as pupae.
Foraging racerunners dig frequently with the forefeet, and some of
the prey at least is found beneath the surface. As compared with
some others, these lizards are dainty feeders, which spend a long
time killing, and worrying their prey, and sometimes reducing it
to small morsels, instead of gulping down relatively large objects.
On May 20, 1955, an adult male that had been under observation
about ten minutes as he foraged along a ditch bank, was seen to
find a 1/2-inch larva of a sphingid moth. Immediately he attacked
it, biting it, shaking it, and tossing it about. However, the victim
was too bulky for this lizard to subdue or swallow. After struggling
with it for approximately two minutes, he left it, still alive though
perhaps seriously injured.

Breeding

Sexual behavior in various kinds of lizards kept under laboratory
conditions, was studied by Noble and Bradley ( 1931 ) . The species
observed included Cnemidophorus sexUneatus, and another teiid,
Ameiva chrysolacma. Both were similar in their courtship and
mating, illustrating the authors' findings that each family of lizards
has a characteristic behavioral pattern that varies somewhat in dif-
ferent genera and species, but to a lesser degree than families differ
from each other.

The report by Noble and Bradley is an important contribution
and has done much to promote a better understanding of the be-
havior of lizards. Nevertheless, it is my impression that their
results were influenced to a large degree by the conditions of cap-
tivity, such as crowding and relatively low temperature, and that
these results have been interpreted too literally as applying under
natural conditions.

Noble and Bradley stated that the male racerunner commences
sexual activity by rubbing his pelvic region on the ground in lateral
movements, and he may continue this rubbing as he runs in a cir-
cuitous course the shape of a figure eight. Thus stimulated he
may pursue any other racerunner that comes within sight, and
overtaking it he may poke or nip its neck with his pointed snout.
The male has well-developed femoral pores, and having attained a
dorsal position straddling the mate, he employs leg rubbing as
further stimulation, also continuing the pelvic rubbing on the fe-
male's back. He thrusts his tail beneath that of the female to attain

Six-lined Racerunner 33

cloacal contact, and as copulation begins, he shifts to a flank grip,
with his body arched in a lateral flexure over the posterior part of
the female's back. A few seconds after coitus has begun, the male
begins forward thrusts of the pelvis, about two per second. In
the instances observed by Noble and Bradley, copulation in Cnemi-
dophorus lasted from five to fifteen minutes, considerably longer
than in Ameiva. In 15 instances of attempted matings, eight of
the courted lizards were males and seven were females. Therefore
Noble and Bradley concluded that racerunners do not recognize
sex in other individuals, except by behavioral responses. Fighting
also was observed, and in this phase of behavior the racerunner
differed from Ameiva, in which the males do not fight and the sexes
are similarly colored. An aggressive male of Cnemidophorus would
rise high on his legs, thus showing off the bright colors of his sides
and would rush at others to bite them with a more vigorous and
direct approach than that used in courting, and without the prelim-
inary pelvic rubbing and neck poking.

The laboratory temperatures in the study by Noble and Bradley
averaged 82.2°F, which is now known to be below the normal ac-
tivity range of the six-lined racerunner. Significantly, the authors
noted that courtship and fighting occurred most often just after
treatment with artificial ultra-violet illumination, when presumably,
the lizards had temporarily raised their body temperatures. From
the text and photographs it is my impression that most of the lizards
in the crowded cages were lying in a semi-torpor, too sluggish
either to escape or to make the normal responses ordinarily evoked
by the approach of another individual. Under these conditions, the
aggressor lacking the usual cues, might easily mistake the sex of the
individual approached. Also, in lizards as a group, a more powerful
male may express his dominance by homosexual mating with a
weaker individual, who is unable to escape or defend himself effec-
tively. In the cages, containing many individuals of each sex, ol-
factory cues must have been much confused, and like other reptiles,
the racerunner probably does not readily shift its attention from one
type of response to another. The long time required for comple-
tion of copulation probably was a result of the unfavorably low
temperature, as field observations have not borne out the laboratory
findings in this regard.

In adult six-lined racerunners the striking differences in coloration
and proportions probably serve as the basis for social releasers,
eliciting courtship, fighting, and other types of behavior. The fe-

34 University of Kansas Publs., Mus. Nat. Hist.

male retains the juvenal pattern little altered except that the bright
blue of the hatchhng's tail fades to brown, and the vivid longitu-
dinal hght stripes of the hatchling become dull, less sharply set oflF
from the ground color. In the adult male however, the stripes fade
more completely and those on the sides may almost disappear;
meanwhile the dorsum and sides become increasingly sufiFused with
greenish coloration, shading into blue-green on the flanks. The
male's head is broader and more robust than that of the female,
and even when the female's body is not distended with eggs, it is
of more plump appearance than that of the male.

The conspicuous sexual dichromatism of the six-lined racerunner
is not shared by most other members of the genus and family, nor is
it associated with elaborate behavioral displays such as are common
in lizards of some other families. Sexual relations seem to be pro-
miscuous with a minimum of courtship behavior. Association be-
tween members of a pair was of brief duration in the instances that
I observed.

Copulation was observed on several occasions. On May 30,
1953, at 8:40 A. M. at the west end of the quarry, my attention was
attracted by rustling in dry leaves, to two racerunners struggling
on the ground. The male had grasped the female near the base of
her tail on one side, but the two were facing in opposite directions.
The female was thrashing and twisting in vigorous attempts to
escape. After about 20 seconds the male suddenly shifted to a
dorsal neck grip. For the next three minutes he rode the female,
as she carried him about in short jerks alternating with brief pauses,
moving in circles a few inches in diameter. From time to time the
male's hindquarters made waggling movements, and several times
he shifted his grip slightly. This phase was terminated when the
male suddenly shifted his grip about two inches posteriorly, to the
female's flank, and arched his body about her hindquarters, com-
mencing copulation, which lasted about 1% minutes. Afterward,
the male released the female, and as she darted away, he followed
her for a few inches, with his tail arched and his vent pressed against
the ground.

On June 7, 1956, a pair was found copulating on bare soil of the
dry ditch bottom. At the disturbance of my approach, they broke
apart, and the female ran, with the male in pursuit. The female hid
in concealing vegetation of the ditch bank. A half-grown juvenile,
basking nearby, was startled by the sudden movements and darted
aside; instantly the male's attention was transferred to it, and he

Six-lined Racerunner

35

chased it for 50 feet along the bank. On June 24, 1956, an adult
male was seen chasing a much smaller, but perhaps sexually mature,
female along the ditch bank. Several times the larger hzard caught
the smaller by the side, and turned it on its side or back. The
smaller lizard was quiescent most of the time that it was held down,
but when the male attempted to shift his grip, it broke loose and
darted away.

On July 1, 1956, an adult male and female were placed in a cage
with a small female (63 mm. snout-vent length) which had laid
eggs the day before. The eggs had been removed from the cage.
The young female had been kept there for several days, and had
become familiar with her surroundings. Almost immediately she
darted at the larger female and bit her. In the next few minutes
she repeated the attack many times, approaching the larger lizard
with menacing demeanor — standing high off the ground, wdth snout
depressed and body tilted toward her adversary. The larger fe-
male, being in strange surroundings, merely attempted to escape
without retahating. Toward the adult male the young female's
attitude was strikingly different; she avoided his proximity, but with
no display of hostility.

MAY

jure JULY AUG.

Fig. 5. Records of adult female
racerunners grouped in five-day inter-
vals; those known to be gravid are
shown in lined columns, and those not
gravid are shown in solid black.

Breeding begins a few weeks aftetr emergence from hibernation,
and extends through much of the grovnng season. For a week or
more before laying, the enlarging ovarian eggs, and those in the
oviducts after ovulation has occurred, so noticeably distend the
body of the female that her gravid condition is obvious. Over an
eight-year period such gravid females were recorded in May on
the 27th, 28th, and 30th; in June on the 8th (2), 10th, 11th (2), 13th,
16th, 20th (2), 21st (2), 22nd (1), 23rd (3), 24th, 25th (4) and

36

Univ'ersity of Kansas Publs., Mus. Nat. Hist.

28th (2); in July on the 1st (3), 6th, 9th (2), 11th (4) and 13th (2);
and in August on the 12th (2). The concentration of records near
the end of the third week in June and again in the second week of
July suggest that two main broods are represented. Probably the
same females in some instances, contribute to both, with an inter-
vening interval of two to three weeks betvveen successive clutches.
In 1955 and especially in 1949 the trend of records was somewhat
advanced as compared with other years. Individuals were not
recaptured with sufficient regularity to permit tracing the history
of their breeding, in most instances. One large adult female re-
corded as gravid on July 1, 3, 5, 7, and 9, was again recorded as
gravid with what was probably a second clutch, on July 19. Another
large female was recorded as gravid on June 23 and 25, 1954, and
again on July 6. In view of the racerunner's short growing season
in the latitude of northern Kansas, it is remarkable that females
mature sexually and produce eggs while still in their first year. Rec-
ords of three such individuals, gravid when less than eleven months
old, are recorded in Table 2.

Table 2. — Lengths and Weights of First-year Females Nearly Ready to
Lay Their First Clutches of Eggs, in Late June.

Date

June 28, 1954
June 23, 1955
June 21, 1956

Snout-vent
length
in mm.

Weight

in
grams

Still another female, 63 mm. in snout-vent length when captured
in the last week of June, 1956, laid two eggs while in confinement
on June 30. These two eggs were of normal size and constituted
the entire clutch. As in other reptiles, size of clutch tends to be
proportional to size of the female. The measurements of the 24
museum specimens found to be gravid are not entirely comparable,
because of shrinkage of several per cent after preservation. Of
these 24 gravid females, half were considered to be fully adult
(lengths 85, 85, 76, 71, 70, 70, 69, 69, 68, 68, 67, 67) and the re-
maining half were considered to be first-year individuals (lengths,
in mm. 65, 64, 64, 63, 62, 61, 61, 61, 60, 60, 58, 54). For the group
of larger individuals, judged to be fully adult, clutches averaged
4.4 eggs, while for the smaller lizards judged to be in their first
year, clutches averaged only 2.0 eggs (1 to 3, see Fig. 6).

Six-lined Racerunner

37

The eggs and nesting habits of the six-lined racerunner are well
known, especially through the careful studies of Brown (1956) in
Piedmont North Carolina. By digging in old sawdust piles left from
transitory sawmill operations, Brown was able to find nests in quan-
tity. The sawdust piles were preferred over more natural nesting
sites in sandy soil, and had unusual concentrations of nests. For
55 such nests found in sawdust, mean depth was 5.6 inches, varying
from three to eleven inches and usually the nest was situated in a

T — r

to
o
^.

u.
o

a.

Mi
00

J.

J.

_L

J L

55 60 65 70 75 80 85
SNOUT-VENT LENGTH IN mm.

Fig. 6. Number of eggs per clutch
(determined from gravid museum
specimens) correlated with sizes of
females. First-year females (usually
less than 65 mm. in snout-vent length)
produce smaller clutches than do fully
mature females.

moist zone. A typical nest cavity was closed ofiF, with a low domed
ceiling, and had a volume of twice or even three times that of the
enclosed eggs. Daytime temperatures at the depths where the eggs
were situated were always near 90 °F, while surface temperatures
were much higher, and deep within the piles temperatures ap-
proached or exceeded 100 °F. Number of eggs within a clutch
was most typically three ( 29 instances ) , but might be two ( 18 in-
stances), four (15 instances), one (3 instances), or five (twice).
By gradual absorption of moisture an average egg enlarged from

38 University of Kansas Publs., Mus. Nat. Hist.

16.2 .X 9.3 mm. and .78 grams at laying to 18.4 .\ 13.0 mm. and 1.76
grams at hatching. The average hatchhng had a snout-vent length
of 32 mm., tail 61 mm., and weight .84 gram, with no significant
difiFerences in size or proportions between males and females.

Hatching dates in the clutches studied by Brown ranged from
late June to early September, witli concentrations near mid-July
and in early August. Post-laying incubation of approximately 60
days was indicated. Although Brown assumed that an individual
female produces only one clutch per season, the distribution of
hatching dates seems to suggest that two or more clutches are
produced. It is evident that the breeding season begins at least a
month earlier in the region of Brown's study than it does in north-
eastern Kansas. Also, it is of longer duration and less concentrated.

Milstead (1957:120) studying four other species of Cnemido-
phorus (sacki, perplexus, tessellatus, and tigris) in western Texas,
reached a rather surprising conclusion regarding the production of
eggs : "The size of mature eggs in the oviducts implies that it would
be a physical impossibility for more than two eggs to mature simul-
taneously." He found one to four eggs in most of the gravid fe-
males examined, but was of the opinion that where the larger
number was present they represented more than one clutch.

Of five clutches that I recorded on the Reservation, one had six
eggs, one had four, two had three, and one had two, an average of
3.6, contrasting with the average of only 2.9 found by Brown in
a much larger series of clutches, but in the 24 gravid museum spe-
cimens that I examined, mostly from Kansas and Arkansas, the
average clutch was only 2.6. Five was the maximum number found
in a clutch by Brown, who cited records from the literature ( Boyer
and Heinze, 1934; Cook, 1945; Minton, 1944 ) of six eggs per clutch,
and mentioned the possibility of larger average clutch size in other
parts of the range. It seems that one of the two records by Cook
of six eggs in a nest pertained to a composite clutch, since three of
the eggs were on the point of hatching while the other three had
small embryos. For 13 clutches reported in the literature, from
Indiana (Minton, 1944), Kansas (Marr, 1944), Mississippi (Cook,
1940), Missouri (Boyer and Heinze, 1934), Nebraska (Hudson,
1942) and Oklahoma (Force, 1930) the average was 4.5, even
higher than in my own sample from the Reservation. It seems
that size of clutch is subject to geographic variation, but the chief
variable may be composition, by age, of the population of females
contributing to any sample of clutches. Age composition of course
varies according to the time and place. For the total of 112 clutches

Six-lined Racerunner

39

recorded in my study or reported in the literature, there were 339
eggs, an average of 3.03 per clutch.

Growth

Earliest appearances of hatchling racerunners were in the first
week of August, but in most years they did not appear until the
second week of August, Each year within a few days after the
first appearance, young all about the same size, had become abun-
dant. By September, young of noticeably diflFerent sizes were in
evidence; some near the minimum size were from late broods, and
others from early broods were partly grown. The smallest young
were 32 to 35 mm. in snout-vent length and weighed approximately
one gram. Most rapid growth occurs in the interval between hatch-
ing and the time of retirement into hibernation. From the size of
the largest hatchlings caught in September, it is evident that the
most successful young average an increment of approximately half
a millimeter per day onto the snout-vent length. By the time of
hibernation, after approximately six weeks of growth, they are
about midway in snout-vent length between newly emerged hatch-
lings and average-sized adults. These half -grown young weigh a
little less than half an much as small adults. Sizes of selected indi-
viduals of these half-grown young are shown in Table 3.

Table 3. — Sizes of Most Advanced Hatchlings in Early Autumn of Three
Different Years, Illustrating That, in Some, Nearly Half the
Growth From Hatching (Snout- vent Length 32 to 35 mm.) to Ma-
turity ( 72 mm. ) Is Made Within a Few^ Weeks in Late Summer Before
the First Hibernation.

Year

Snout-vent
length
in mm.

Date

of
record

Most

probable date

of hatching

1952

53
52
53

September 16
September 9
September 18

August 13
August 9
August 4

1953

1954

These three young had each increased approximately 20 mm. in
snout-vent length, assuming they were of average hatching size at
the start, and that their emergence corresponded approximately
Mdth my earliest records of hatchlings in the same years. Some
young from broods hatched late in the season make relatively little
growth before they are forced to hibernate.

Emerging, usually in May, after their first hibernation, the young
resume their rapid growth, but the rate gradually decreases as they

40

University of Kansas Publs., Mus. Nat. Hist.

approach minimum adult size. By the end of the summer all are
adults. The five individuals whose histories are set forth in Table
4, show the variation in size and the trends in growth in these first
year young after emergence from hibernation.

Table 4. — Gro\vth in Young Racerunners After Hibernation.

Individual

AND

Sex

Dates

of
capture

Snout-vent
length
in mm.

Tail

length
in mm.

Weight

in
grams

No. 1, female. . .

May 9, 1956
May 21, 1956
June 26, 1956
August 9, 1956

391^
44
55
68

65

74

101

130

1.3

1.8
3.1
7.0

No. 2, female.. .

May 21, 1956
July 3, 1956
August 14, 1956

42
55
69

81^
113
138

1.6
3.4
8.1

No. 3, male. . . .

May 31, 1956
June 20, 1956
August 2, 1956

48

50^

70

80

91

133

2.3
2.9

No. 4, female . . .

June 1, 1956
July 9, 1956

46
61

85
113

2.2
4.3

No. 5, male. . . .

June 29, 1956
August 12, 1956

62
71

46
(recently broken)

57
(+60 regenerated)

3.9

8.8

Table 5. — Growth in Individual Racerunners Approaching Adolescent
Size. In This Group Average Gain Per Day Was Almost % mm. For
Intervals That Averaged Almost a Month.

Increase

Increase

m

m

snout-vent

Growth

snout-vent

Growth

Sex

length from

interval

Sex

length from

interval

beginning

in days

beginning

in days

to end of

to end of

interval'

interval*

cf

52 to 58

23

9

49 to 61

31

d"

53 to 58

17

cf

49 to 59

20

&

53 to 59

35

9

46 to 61

38

cf

56 to 58

29

9

50 to 56

13

9

50 to 56

44

1. Total gain 77% mm.
0.31 mm.

Total elasped time 250 days. Average increment per day

Six-lined Racerunner

41

Slower growth in larger young is illustrated by a group of 26 that
averaged 55.2 mm, in snout-vent length at the beginning and 68.1
mm. at the end of growth intervals that averaged 34 days, as shown
in Table 6.

Table 6. — Growth in 26 Racerunners of Adolescent Size. On The
Average, These Grew Less Than /« mm. per Day.

Increase

Increase

in

in

snout-vent

Growth

snout-vent

Growth

Sex

length from

interval

Sex

length from

interval

beginning

in days

beginning

in days

to end of

to end of

interval*

interval *

cf

51 to 70

64

&

57 to 61

14

(f

58 to 64

38

d^

583^ to 70

49

9

61 to 61H

22

7

62 to 68

20

d"

59 to 63

29

&

65 to 69

12

cf

60 to 65

20

cf

62 to 66

14

&

56 to 68

69

&

64 to 68

20

cf

54 to 67

68

cf

65 to 70

21

9

55 to 68

44

cf

64 to 67

27

cf

58 to 63

23

&

58 to 71

48

9

55 to 69

42

9

63 to 71

41

9

56 to 61

41

cf

62 to 64

19

cf

50^ to 70

53

cf

64 to 70

23

9

59 to 62

11

cf

62 to 71

44

1. Total gain 199% mm. Total elapsed time 876 days. Average increment per day
0.226 mm.

In these large young, approaching adult size, the growth rate
had slowed to less than half the daily length increment of hatch-
lings. Eight even larger young increased in snout-vent length from
66.6 to 73.1 mm. in intervals that averaged 35 days, thus having an
average increment of .18 mm. per day as small adults.

Twenty-seven young adults recorded in May and June, all known
to be second year individuals because they had been caught and
marked in early stages of growth, averaged 72.8 ± .51 mm. in snout-
vent length, and ranged from 77 mm. to 68 mm. Eleven marked
adults that were recaptured when they were known to be late in
their third years (having hibernated in three winters) averaged
75.6 dz .78 mm., indicating a growth rate of 2.8 mm. beyond the
average size of second year individuals. Five fourth-year adults
averaged 78.9 mm. (77)2 to 81); four fifth-year adults averaged 81.5
mm. (79 to 84), and the only known sixth-year individual had a
snout-vent length of 83 mm. The largest racerunner recorded had
a snout-vent length of 84 mm., and it was one of those known to

42 Unrtersity of Kansas Publs., Mus. Nat. Hist.

be in its fifth year. In each series males and females averaged ap-
proximately the same size, and the small differences were not sta-
tistically significant.

As growth proceeds, there are changes in proportion and these
are somewhat different in the sexes. Relative size of the head as a
ratio of total length, and the shape of the head are among the more
important changes, and there are smaller changes in the propor-
tions of torso and limbs. No attempt was made to analyze these
changes, but length of the tail was one of the items regularly re-
corded in each lizard handled, and, in those racerunners having
intact tails, the relative length of tail varied according to age
and sex, being longer in adults than in young, and longer in mature
males than in mature females.

<
a:

1

1 1 1

_80

WEIGHT...-—-'- _,.-.--

/'.'-•'""length 8

70

/V

60

//

t
#

'7 6

}

50

0

4

*

t

0

*

3.

/

,./

/ ,.'

1 ,'

2

1 .'

1

•?

■

30 1 ,

L.

-J lJ 1 1 1 U 1 i 1 lJ 1 1 L-

6 / ^ .. «>

Ui

growing seasons

Fig. 7 Correlation of average length and average weight with age in six-
hned racerunners through four growing seasons. Each growing season is di-
vided into 4Js months. Growth is most rapid during the first year, but con-
tinues, at a decreasing rate, for several years, perhaps throughout the indi-
vidual's lifetime.

Six-lined Racerunner 43

Proportions of the Tail and It's Regeneration

The racerunner's tail is flagelliform, and in adults is approxi-
mately twice the combined length of the head and body. In closely
related species, Cnemidophoriis per plexus and C. inornatus, the tail
is relatively even longer, and it may be nearly three times the snout-
vent length. In the racerunner, as in the majority of other Hzards,
the tail is fragile, breaking easily if it is struck or pinched. If
grasped by the tail, the racerunner promptly snaps off the appen-
dage in order to escape. The detached tail performs jerky squirm-
ing movements for several minutes, a circumstance obviously ad-
vantageous to the lizard in allowing it to escape while the atten-
tion of a predator is distracted. However, in the racerunner the
detached tail is less lively and conspicuous than in other common
lizards including skinks, anguids, and iguanids. Also, the tail is
less fragile tlian in some of these kinds.

Brown (1956:38) found that in hatchling racerunners in North
Carolina, snout-vent length averaged 32 mm., and tail length aver-
aged 61 mm., or 190 per cent of the former measurement. In all
the hatchlings examined in the course of my field study, relative
length of tail was considerably less. To test the possibility that this
character is subject to geographic variation, I measured series of
museum specimens, including five hatchlings from Louisiana, Ar-
kansas and Oklahoma, in which tails averaged 199 per cent of
the snout-vent length, and 13 hatchlings from Kansas the tails of
which averaged 166 per cent of the snout-vent length. Comparable
differences may exist in adults, although the material available to
me is not well adapted to demonstrate them. In 12 adults (more
than 61 mm. ) from Arkansas, tails averaged 211.5 per cent of snout-
vent lengths whereas in 18 adults from Kansas tails averaged 197.6
per cent of the snout-vent length.

Relative length of tail increases as the racerunner grows. For 12
small young 30 to 40 mm. in snout-vent length, the tail averaged
167 per cent of the snout-vent length. This increased to an average
of 199 per cent in males of more than 70 mm. in snout-vent length
( Fig. 8 ) . In males that are partly grown and those that are adult,
tails consistently average somewhat longer than in females of com-
parable size, but there is overlapping. These figures are based
on live racerunners measured in the course of my field study; the
figures in the preceding paragraph apply to preserved museum

2—3118

44

Untv'ersity of Kansas Publs., Mus. Nat. Hist.

specimens that cannot be measured so accurately, as tliey can-
not be relaxed, and are subject to some shrinkage and distortion.

X

200

42

^ 38

.--0

LLl

1 . '

_1
1-

Z
UJ

>

190

2
20 ,.-'

40

\-

180

-'$

3

~*

/

O

^

cn

/

0^

17

0/

]/l2

_l

.<r

H-

160 .

1

,

40 50 60 70
SNOUT- VENT LENGTH, mm.

Fig. 8. Relative tail-length in live
racerunners of different sizes. Tails
are shortest in hatchlings, and increase
in relative length until in large adult
males they are nearly twice the com-
bined length of head and body. Rec-
tangles indicate standard errors of
the means; figures indicate size of
each sample.

These factors may reduce by several per cent the snout-vent meas-
urement, thus resulting in a high figure for relative tail length.

Many adult racerunners retain the tail intact, whereas others
have had their tails broken or regenerated one or more times.

Table 7. — Percentages of Racerunners Retaining Intact Tails in Various
Size Groups, From Hatchlings to Large Adults.

Size group, snout-vent length in mm.

30

to
39

40
to
49

50

to
59

60
to
69

70

to
79

80
to

84

Number in sample ....

Percentage retaining
tails intact

14
100.0

20
95.0

39
92.2

55

87.1

84
77.3

8
50.0

Six-lined Racerunner

45

Probably the incidence of broken tails varies according to the
time and place, under the influence of such factors as numbers and
kinds of natural enemies present, and the population density of
the racerunners themselves, influencing to some degree the amount
of intraspecific competition and actual fighting. Table 7 shows the
decreasing percentage retaining intact tails as greater size and age
is attained, but even in the largest and oldest group, half retain
their original tails. Differences between the sexes are not evident
in this connection.

In individuals having regenerated tails, length of the regenerated
part varied greatly in proportion to the original part and the body
length (Fig. 9). Records of marked individuals have shown that
most rapid growth of the regenerating tail takes place within a
few weeks after a break. Presumably, therefore, the majority of

a

o

UJ

o

< z

cc ili

z (X

UJ UJ
O Q.

q: ^

-100

X

-80

o

z

LU

_l

-60

1-

z

UJ

>

-40

1-

Z)

o

z
in

-20

J.

• •• •

• ••

•••.

/ . ••.•;•• •

V

I

J l!

X

J.

• •

±

20 40 60 80 100 120 140 180
TAIL -STUB AS PERCENTAGE OF SNOUT- VENT LENGTH

Fig. 9. Relative lengths of original and regrown parts of the tail in individual
racerunners that have lost and regenerated varying lengths of their tails.

regenerated tails recorded were not still in process of active growth.
Size and age of the lizard at the time a break occurs, and the abun-
dance of food in the early weeks of regeneration are among the fac-
tors which might have important effects on the amount of regen-
eration.

Individual histories of several of the racerunners that yielded
most significant records concerning regeneration of their tails are
set forth in Table 8.

46

University of Kansas Publs., Mus. Nat. Hist.

Table 8. — Rate and Amount of Regeneration in Racerunners Having

Broken Tails.

Individual

Dates

of
capture

Snout-
vent
length

Tail length

Xo. 1, young male

May 24
June 19
June 28
July 25
July 31

54
60
60
67
67

35 (+ 68 broken off)

40 (+ 2 regenerated)

41 (4- 8 regenerated)
50 (+ 48 regenerated)
50 (-1- 49 regenerated)

No. 2, voung male

June 29
August 12

62
71

46 (broken stub)

57 (+ 60 regenerated)

No. 3, small adult male ....

June 15
July 1
Julv 28
Api-il 26

71H
73
74
74

87 (broken in capture)
87 (+ 11 regenerated)
91 (+ 13 regenerated)
91 (+25 regenerated)

No. 4, adult male

June 17
July 15
July 28
August 14

76
76

77
77

92 (broken in capture)

95 (+ 63^ regenerated)

96 (+ 22 regenerated)
96 (+ 28 regenerated)

No. 5, large adult male

June 28

July 7
May 22

80
80
80

96 (+ 4 regenerated)
96 (+ 23 regenerated)
96 (+35 regenerated)

No. 6, small adult female . . .

June 23
May 20
July 11
June 5

63
74
74
74

76 (+ 19 regenerated)

77 (+45 regenerated)
77 (+45 regenerated)
77 (+ 51 regenerated)

No. 7, small adult female . . .

June 2
June 24
August 8

71
71

77

68 (recently broken stub)
70 (+ 16 regenerated)
75 ( + 39 regenerated)

Molt
The racerunner, like other hzards, undergoes periodic molt in
which the outer layer of epithelium loosens and breaks off in small
translucent flakes. The molt begins on the head and progresses
posteriorly. Several days sometimes elapse from the beginning to
the completion of the process. The distal part of the tail usually
retains its layer of old skin after shedding has been completed on
the head, body, and limbs.

Six-lined Racerunner 47

To determine how often shedding occurred, I marked the skins
of most of the racerunners that were examined. In 1949 and 1950
indehble black ink was used, but it was unsatisfactory, becoming
obscured within a few days. In subsequent years I applied enamel
paint of a bright color ( usually red or yellow ) contrasting with the
color of the skin, to an area perhaps half an inch square, on one
leg, one side of the tail base, the flank, nape, or some other con-
spicuous location to serve the dual purpose of individual recogni-
tion in the field, and of indicating when the racerunner shed its
skin. This paint was not wholly satisfactory either, as the color
was soon dulled by bleaching and by adhering soil, and the paint
tended to peel or flake off.

Occasionally lizards were in process of shedding at the time of
capture or at least retained old skin on the distal part af the tail
indicating recent shedding. Others, from their glossy "new" ap-
pearance, were known to have just completed shedding. However,
for most, the time of shedding was unknown. No individual was
recorded twice in consecutive sheddings. The paint merely served
to show whether the lizard had, or had not molted since the pre-
ceding capture when paint was applied. Lizards marked with
paint were recaptured while still retaining the color after the fol-
lowing intervals of days: 28, 26, 22, 18, 17, 15, 14, 14, 13, 13 and 11,
in addition to many shorter intervals of less than a week. Others
that were recaptured after the following intervals of days had
elapsed had lost their paint: 62, 62, 50, 45, 42, 40, 36, 35, 27, 20,
20, 17, 16, 13, and 13. No racerunner recaptured after a hiberna-
tion period retained paint from the previous season. Some of those
marked with paint probably were almost ready to shed, and hence
lost the color within a few days after their release, whereas others
perhaps had shed a short time before they were painted. The fact
that none was recaptured retaining paint after more than 28 days
suggests that this is near the maximum interval elapsing between
molts in the season when the lizards are active. Some may molt
at considerably shorter intervals. As in other kinds of lizards, grow-
ing young probably shed their skins more frequently than do adults.
Some of the racerunners found to have lost paint between captures
may have shed twice in the intervening time.

One of the best records was that of a female recorded seven times
in the summer of 1956, from May 9, soon after emergence from hi-
bernation, when she was only a little larger than the size at hatch-

48 University of Kansas Publs., Mus. Nat. Hist.

ing, to August 13 when she was of small adult size. Her records

follow.

May 9 Marked with red paint.

May 21 In process of completing molt; paint renewed.

June 1 Paint retained.

June 26 Paint gone; renewed.

August 8 Paint gone; renewed.

August 9 Paint gone; renewed.

August 13 Paint retained.

For this rapidly growing young racerunner, four molts are indi-
cated in the three-month period covered by the records, and a
fifth molt might reasonably be expected before retirement into hi-
bernation.

A young male more than half grown when first captured on May
1, 1956, was recorded seven times within an interval of a little less
than two months, as listed below.

May 12 Marked with red paint.

May 28 Paint gone; renewed.

May 30 Paint retained.

June 10 Paint retained.

June 25 Paint retained.

June 26 Paint gone; renewed.

July 3 Paint retained.

This racerunner is known to have shed twice within the first
half of its season of activity — a period of a little less than two
months. The two molts are known to have been separated by an
interval of at least 29 days, and probably somewhat more. That
no additional molts occurred within this period is amply demon-
strated by the captures on May 30, June 10, and June 25, when this
lizard retained its paint.

Predation and Parasitism

Escape reactions
The racerunner depends primarily on speed to escape from its
enemies. No actual observations on the pursuit and capture of
one by a predator are known to me. The escape behavior evoked
by the presence of persons may differ from the responses to some
natural enemies. Racerunners found in the field nearly always dem-
onstrated alertness and timidity. Ordinarily they detected a human
intruder before he was aware of the lizard's presence; the lizard
might be noted merely as a flash of motion as it darted away to
shelter. Only on rare occasions, when a racerunner was preoccu-
pied with catching prey, fighting, or mating activities was it ob-

Six-lined Racerunner 49

served unaware at close range. Racerunners seemed able to detect
the approach of a person from a distance of at least 50 feet, or per-
haps considerably farther. At such mild alarm, the lizard might
cease its previous activity, sometimes crouching against the ground
on the spot where it happened to be and sometimes darting to
nearby shelter such as a clump of grass. The racerunner might
permit approach to within a few yards. Lying flattened against
the ground, it was inconspicuous. I have never seen a racerunner
in the open until it moved. The crouching racerunner might wait
until a person had approached within a few yards before darting
away. A racerunner approached by a person often seemed confi-
dent of its ability to escape, and was uneasy rather than alarmed.
It might resume or continue its foraging, while obviously attentive
to the intruder, holding its head cocked toward him, and moving
furtively, skulking through screening vegetation, or dashing across
small bare areas between such coverts. Where the racerunner's
habitat was in a narrow band, along a road, or bank of a ditch or
gully, one might run ahead of a person for as much as 100 yards,
maintaining a minimum distance of several yards with occasional
spurts alternating with the usual jerky, half-running and half-
walking gait. Behavior varied somewhat according to individual
temperament, age, sex, and season, the temperature, and the actions
of the person. If sudden movements were avoided by the observer,
the racerunner seemed to become accustomed to his presence, and
might even approach to within three or four feet of a person stand-
ing still. In such instances, however, it was obvious that the lizard
was still alert and suspicious.

A sudden attempt to overtake or seize a nearby racerunner was
rarely successful, and only resulted in the almost instantaneous dis-
appearance of the lizard. Running full speed it would move so
rapidly that it became a blur and characteristically it darted in an
erratic course from one clump of vegetation to another, with fre-
quent changes in direction. It might enter a burrow, or might stop
beneath screening vegetation, but in either case it could not be
readily relocated. If, upon resumption of activity, it was again
approached, it was much more wary than before, maintaining a
greater distance between itself and the person following, and pan-
icking more easily.

On May 15, 1952, one seen on the road near the Reservation head-
quarters ran along the wheel track for about 30 feet, then turned
oflF abruptly and ran several yards through brome grass more than
a foot high. The grass was not thick, and stalking the hzard, I was

50 University of Kansas Publs., Mus. Nat. Hist.

able to observe its tactics. It would crouch motionless until I came
within perhaps five feet, then would spring up and dash away, spin-
ning around and changing direction after a few feet or sometimes
only a few inches. After several such short, erratic dashes, it would
stop abruptly and then was difficult to see. Soon it eluded me.

Racerunners that were chased into burrows usually emerged
again promptly. They seemed to have no central home base upon
which they depended for shelter, but each directed its movements
with respect to a succession of escape coverts as it foraged. Usually
a racerunner is able to attain shelter in a dash of only two or three
seconds, whenever the need arises. In crossing open areas where
there is no cover, the racerunner usually travels in long, rapid dashes.

Edgren (1955:141) described how, in sandy areas in Illinois, race-
runners that had burrowed in the sand would "explode" tlirough the
surface, at any disturbance, and would dash away to sheltering
vegetation. The burrows in which the lizards rested were shallow,
nearly straight, parallel to the surface, and about VA times the length
of the lizard which rested inside. In western Tennessee, Blanchard
(1922:6) recorded these racerunners as using short burrows dug
in the sand. When a racerunner was inside, it partially filled one
of the burrow openings, with sand thrown out from within. The
lizards spent the night in such burrows, and could be easily caught
in early morning before they had become active. To escape they
would rush into grass or brush.

Burt and Hoyle (1934:202) recorded that a six-lined racerunner
chased into a stream in Beckham County, Oklahoma, escaped by
swimming 12 feet to the opposite bank. Dillon and Baldauf (1945:
174) recorded one which after it was alarmed, scurried through
dry leaves to the edge of a stream, where it jumped two feet into
the water, and took shelter beneath a submerged rock near the
farther bank. Stille (1947:143) in catching racerunners in the sand
dunes of northern Indiana, found that they would often escape by
running into the water of small ponds and marshes and hiding on
the bottom, especially when they were chased downhill toward the
water.

Burt (1933:192) described the defensive behavior of racerunners
that were found in shallow pits beneath flat rocks along prairie
ledges in Kansas, on cool days in spring. The lizards, not fully
active, and unable to escape by running, would back into crevices,
facing outward, with jaws gaping threateningly.

Six-lined Racerunner 51

Natural enemies

In confinement, various predators have been recorded to eat
racerunners, but the significance of such records is doubtful. Burt
(1933:191) wrote that a captive box turtle {Terrapene ornata) ate
one that was placed in the turtle's cage. It is well known that these
turtles avidly accept animal food including small vertebrates, and
in many areas both the turtles and racerunners are abundant. A
turtle would be unable to catch a racerunner that was fully active,
but the box turtle can carry on normal activities at temperatures
many degrees lower than those required by the racerunner (Fitch,
1956:439). Therefore, the turtle might be able to dig out and
devour racerunners that had taken shelter beneath flat rocks or in
shallow burrows in loose sand. Hurter (1911:130) recorded that
a captive collared lizard (Crotaphtjtus collaris) ate a racerunner,
and Pritchett (1903:284) noted another instance. The collared
lizard is notorious for eating small lizards, and may be an important
predator on racerunners in certain areas where the ranges and
habitats of the two overlap. Klauber (1956:610) quoted Loewen
(1947:53) concerning a captive massasauga {Sistmrus catenatus)
that ate two racerunners, as part of a varied diet that included
several kinds of small birds and mammals and even pieces of meat.

Wright and Funkhouser (1915:130) recorded racerunners from
the stomachs of two racers (Coluber constrictor) from the Okefino-
kee Swamp in Georgia. Marr (1944:484) recorded a racer (C.
flagellum) from Trego County, Kansas, that had remains of at
least two racerunners in its stomach. I found a racerunner in the
stomach of a half grown yellow-bellied racer (Coluber constrictor)
caught in an upland field area of the Reservation on June 15, 1955,
and a juvenal yellow-bellied racer was found to have eaten a hatch-
ling racerunner on September 13, 1955. The racers C. constrictor
and C. flagellum both prefer habitats similar to that of the race-
runner, and are swift-moving diurnal predators that are notorious
lizard eaters. Burt (1935:331) collected a half grown prairie king
snake (Lampropeltis calligaster) near Lamont, Kansas, which dis-
gorged an adult racerunner. In a study of the food habits of snakes
at Fort Benning, Georgia, Hamilton and Pollack (1955 and 1956)
found the racerunner to be a common prey, eaten by no less than
nine species. Of those snakes that contained food, the following
numbers had eaten racerunners: Coachwhip (Masticophis flagel-
lum ) 23 ( of 45 ) ; black snake ( Coluber constrictor ) , 5 ( of 57 ) ; corn

52 Untv'ersity of Kansas Publs., Mus. Nat. Hist.

snake (EJaphe guttata), 5 (of 32); timber rattlesnake {Crotahis
horridus), 3 (of 26); pine snake (Pittiopliis melanoleucus) , 1 (of
5); grey rat snake (Elaphe obsolefa), 1 (of 13); king snake (Lam-
propcltis getulus), 1 (of 13); hog-nosed snake {Heterodon platy-
rhinos), 1 (of 33); pigmy rattlesnake {Sistrurits miharius), 1 (of
12).

In the contents of 104 armadillo stomachs collected in west-cen-
tral Louisiana in 1947 and 1948 (Fitch, 1949) I found one race-
runner, and several clutches of lizard eggs, some of which may have
been those of racerunners. On June 24, 1950, at a hilltop rock out-
crop (Rat Ledge) on the Reservation, remains of a marked adult
female were found where she had been killed and partly eaten by
a predator. The head and tail and one hind leg had been eaten.
The racerunner had been dug out from beneath a rock, and there
was much digging about other rocks in the vicinity. The predator
in this instance was probably a spotted skunk (Spilogale putorius),
judging from the sign, found frequently in this vicinity. Striped
skunks (Mephitis mephitis) and raccoons (Procyon lotor) also are
probable predators on the racerunner, and on the Reservation both
species have from time to time molested reptiles and invertebrates
caught in funnel traps. On May 27, 1955, a dead juvenal racerun-
ner partly eaten by an unknown mammalian predator, was found
on an exposed gully bank. On several occasions harvest mice
(Reithrodontomys megalotis) caught in funnel traps with race-
runners, killed and partly devoured them. In these instances the
harvest mice were impelled to attack by close confinement, and by
hunger in the absence of other food, but free-living individuals of
these and other mice may occasionally prey upon racerunners that
are inactive in their burrows because of low temperatures.

In an exhaustive study of the food habits of the coyote in Ne-
braska, Fichter, Schildman and Sather (1955:23) found that rep-
tiles made up a negligible percentage of the diet over the state as
a whole, but that in the vicinity of the Nebraska National Forest
(in a sandhill area a little north and west from the center of the
state), lizards were eaten frequently. Analysis of 1740 scats re-
vealed lizard remains in the following percentage frequencies ac-
cording to season: winter 4.3; spring 9.3; summer 15.9; fall 2.5.
Although the lizards were not identified as to kind, some probably
were racerunners. Other lizards known to occur in the same area
include only the earless lizard (Holbrookia maculata), the scaly
lizard (Sceloporus undtdatits) , and the skink (Etimeces intdtivir ga-
ins). It is significant that in this sample, lizards were well repre-

Six-lined Racerunner 53

sented not only in spring and summer but in fall and winter when
the population is dormant. This fact suggests that the coyote ob-
tains them by digging them out of the sand, and they could be
effectively hunted by this method on cool cloudy days, or at night,
in the summer. In the contents of 1190 coyote stomachs from Kan-
sas, Gier (1957:12) found lizard remains (Sceloporus) only twice.
In 118 scats of coyotes collected on the Reservation over a period
of years, there were no lizard remains, but most of these scats rep-
resented the colder half of the year ( Fitch and Packard, 1955 ) .

On June 13, 1957, the entire tail of an adult female racerunner
was found in a broad-winged hawk's nest, beside the young hawks.
The remainder of the lizard probably had been eaten by the hawks.
There seemed to be no chance that the lizard had merely shed its
tail and escaped, because the tail included the root, proximally be-
yond the point where fracture can occur. In 1954 more than 100
prey items were recorded in another broad-winged hawk's nest, and
they included no racerunners, although other lizards (Eumeces
fasciatus, E. ohsoletus, Ophisaurus attenuatus) were well repre-
sented. The sparrow hawk (Falco sparverius). Cooper's hawk
(Accipiter cooperi) and marsh hawk {Circus cyaneus) prey ex-
tensively on lizards of kinds that inhabit open situations, and might
be important as enemies of the racerunner where they share its
habitat, but no specific instances of such predation are known to me.

Parasites
Few racerunners were dissected in the course of my study, and
no internal parasites were recorded although there are, probably,
many kinds that infest this lizard. Harwood (1932:65) found the
rectums of two of four racerunners from Huntsville, Texas, to be
heavily parasitized with the oxyurid nematode, Pharyngodon war-
neri. Of ectoparasites, chiggers are by far the most conspicuous.
Almost all racerunners are infested with chiggers throughout the
greater part of the growing season. In early summer most infesta-
tions are extremely heavy. Chiggers attach in tight clusters, and
may occupy almost all areas of exposed skin between the scales.
On the ventral surface the scales are relatively large and the spaces
between them constitute especially favorable sites for attachments,
as do the axilla, the groin, anal region, ear opening, and eyelid. The
masses of chiggers attached to these areas of exposed skin contrast
with the greenish color of the scales giving the lizard the appear-
ance of having reddish markings. Many hundreds, or even thou-
sands, may be attached to the same racerunner simultaneously.

54 University of Kansas Publs., Mus. Nat. Hist.

Loomis (1956: 1386) examined 96 racerunners from eastern Kan-
sas (including many of the same individuals on which my own field
study is based) for chiggers. All were infested, with the excep-
tions of 13 of the 14 collected in May, and one each in June, July,
August and October. In the course of the season, average number
of chiggers per host on the infested racerunners changed as follows:
May, 10; June, 653; July, 288; August, 115; September, 14; October,
1. The trend in these numbers closely paralleled the numbers of
unattached larval chiggers, sampled by placing polished black
plastic squares on the ground for one minute and then counting the
chiggers running over them. These larvae are hatched in great
numbers with the advent of warm, humid weather in late May,
and their numbers continue to increase with the increments of
successive broods in June when weather conditions are most nearly
ideal. With increasingly dry weather and lower humidity charac-
teristic of late summer, chiggers wane in abundance.

Although some 46 species of chiggers occur in Kansas, all those
found on racerunners were the common pest species, Tromhiciila
alfreddtig,esi, which is the kind that commonly infests man. Dozens
of species of vertebrates including most of the mammals, birds and
reptiles that are common in eastern Kansas, are known hosts. Thus
this parasite is unspecific and not dependent on any one kind of host.
Nevertheless certain host species, including the cottontail {Stjlvi-
lagus floridanus), the box turtle (Terrapene ornafa), the black rat
snake (Elaphe obsoleta) and the racerunner, are especially favored,
and carry disproportionately large infestations. Although it is rela-
tively small, the racerunner perhaps carries more chiggers per square
inch of body surface than does any other kind of host. Therefore,
in localities where it is abundant, the racerunner may be an import-
ant factor in maintaining high populations of chiggers.

Populations

Most complete data are available for the small colony of race-
runners that inhabited the diversion ditch near the Reservation
headquarters. This ditch extended for approximately 1000 feet,
from the outlet of a x:)ond to a gully. At most times, in all seasons,
the ditch was dry or had only a slight trickle. Unusually heavy
summer rains occasionally caused the pond to overflow, and for a
period of hours the ditch might carry a large volume of water, even
a three or four foot depth with the result that many of the race-
runners were drowned or washed away. In the summer of 1951,
unusually heavy precipitation flooded the ditch repeatedly, and

Six-lined Racerunner 55

few racerunners survived. In 1952 and 1953, drought conditions
prevailed, with the result that racerunners thrived along the ditch,
and increased in numbers there by both reproduction and immigra-
tion. In 1954 there was heavier rainfall but it was not suflficiently
concentrated at any one time to flood the ditch. The summer of
1955 was, on the whole, dry, but on July 5 a rain of 3.15 inches
flooded the ditch; thereafter for the remainder of the summer, few
racerunners were in evidence. In 1956 they regained their numbers,
but in 1957 torrential flooding in June brought the colony to its
lowest ebb since 1951.

For several years this largest remaining colony on the Reserva-
tion has thus existed on a somewhat precarious basis, subject to
drastic reduction by floods and yet largely dependent on the serai
habitat created by the scouring of the ditch bottom and banks in
such floods. The effective width of the barren or partly barren ditch
bottom and banks combined was estim.ated to average approxi-
mately 25 feet; hence, for the 1000-foot stretch of ditch the total
area was approximately 25,000 square feet, or .57 acre. For the
years 1953 to 1956 numbers of racerunners ( exclusive of hatchlings )
recorded were as shown in Table 9. The known population den-
sity represented by this colony in the four different years were:
65 per acre in 1953, 40 per acre in 1954, 49 per acre in 1955, and
72 per acre in 1956.

The relative numbers of the different age groups fluctuated
rather widely from year to year, as might be expected from the vi-
cissitudes to which this population was exposed, but with sharp
reduction from year to year, usually amounting to somewhere near
half in each age class, as shown by the following percentages for
the four annual samples combined.

57.0 per cent were first-year individuals

25.4 per cent were second-year individuals

10.0 per cent were third-year individuals

5.4 per cent were fourth-year individuals

1.5 per cent were fifth-year individuals
.7 per cent were sixth-year individuals

It should be made clear that the percentages listed above apply
essentially to a population of sexually mature racerunners. By "first-
year individuals" is meant those hatched in the preceding August
or September, and recorded any time, from May to October, in the
entire growing season following their first hibernation. A few
caught in May and early June were not much larger than newly
hatched young, but most were recorded later in the season and were

56

University of Kansas Publs., Mus. Nat. Hist.

more than half grown; many had already attained adult size or
were approaching it. Obviously a sample obtained in spring soon
after emergence from hibernation would have different composition
with a much higher percentage of first year individuals, still small,
after only a few weeks of active life.

InsuflBcient data are available to compute accurately the produc-
tivity of the population. Most females if not all, participate in the
annual breeding season; the yearlings making up fully half the
total may each produce one clutch of one or two eggs, while the
older females, perhaps comprising somewhat less than half, prob-
ably produce two clutches, usually of three or more eggs apiece. A
population of 100 racerunners might be expected to produce per-
haps 200 eggs per season. Even if the eggs are subject to heavy
losses during their incubation, the population might be doubled by
the annual hatch of young.

Table 9. — Composition, by Age Groups, of a Small Population of Race-
runners In Several Different Years.

Year

Age Group

1953

1954

1955

1956

Combined
sample

First year

20
8
4
3
2
1

10
8
4
1

22
5
1

22

12

4

3

74

Second year

Third year

Fourth year

Fifth vpar

33
13

7

2

Sixth vear. . . .

1

Total

38

23

28

41

130

Summary

The six-hned racerunner, a small teiid lizard, is the northernmost
representative of its genus and family, and occurs throughout much
of southeastern North America, but usually in a xeric type of habi-
tat that most closely resembles the desert regions inhabited by
other species of the genus Cnemidophorus. Over a nine year
period at the University of Kansas Natural History Reservation
230 racerunners were recorded a total of 581 times. Racerunners
became scarcer on this area after it was no longer grazed or cul-
tivated; herbaceous vegetation closed in and reduced areas of
bare soil and rock.

Six-lined Racerunner 57

Belonging to a group that is largely tropical, the racerunner
prefers a body temperature higher than that of most North Amer-
ican reptiles ( near 40° ) , and spends a relatively large part of the
annual cycle (half to more than two-thirds) in hibernation. Ac-
tivity is greatest in early summer. The racerunner is strictly
diurnal. It retreats into a burrow for the night, and also resorts
to the burrow by day at times when conditions above ground are
not favorable. In hot weather this lizard is most active in the
morning, and spends the hotter part of the day in its burrow. In
late afternoon the racerunner may again emerge from its burrow
for a second period of activity. The racerunner digs well and
makes its own burrows, but uses also those of various other small
vertebrates. The burrows are utilized to escape enemies, and to
avoid high or low surface temperatures. Air temperature of ap-
proximately 34° C seems to be preferred with a range of at least
19° to 39°, but soil surface temperatures are probably even more
critical. Activity is almost entirely limited to times when the sun
is shining, permitting maintenance of a high and fairly constant
body temperature by insolation.

Although racerunners cover ground rapidly in the course of
normal activities, each individual tends to keep to a small area.
Home ranges of .20 of an acre for males and .25 of an acre for
females are indicated, but occasionally the lizard may explore well
beyond the limits of its normal range, and may even shift to
a new location hundreds of yards away.

Racerunners shed their skins at intervals of about four weeks or
a little less. For young in their first growing season, four or five
molts probably are most typical. Adults shed somewhat less fre-
quently.

Racerunners do not maintain territories, and many of both sexes
share the same area; home ranges overlap broadly. Males fight
and have bright colors that are prominently displayed before such
encounters. In the breeding season promiscuity prevails. Egg-
laying takes place in June, July and early August, thus extending
over a large part of the racerunner's season of activity. Some indi-
viduals, at least, produce two (or perhaps even more) clutches of
eggs in the course of the summer, and young produced one sum-
mer breed the next summer when less than a year old, at which
age they have attained less than half the weight of fully matured
adults. Numbers of eggs regularly vary from one to six, depending
in part on the size of the female. First year females usually have

5S University of Kansas Publs., Mus. Nat, Hist.

only one or two eggs, occasionally three. For 97 clutches reported
in the literature or recorded in my study, the average was approxi-
mately 3.1 eggs per clutch. In the southern part of the range the
breeding season is longer, and there is some indication that clutches
average slightly smaller.

Typically the eggs are buried in sandy soil, at a depth of only a
few inches. Incubation time in the neighborhood of two months
is indicated. Hatchlings first appear in early August, but because
of the extended breeding season, some are near half-grown, that
is to say intermediate in length between hatching size and adult
size, by the time of retirement into hibernation. Others then are
still near the minimum size. Hatchlings resemble adults in minia-
ture, but the light stripes of hatchlings are more vividly outlined
against the dark ground color which lacks the greenish suffusion
of the adult and the tail of the hatchling is bright blue. Also body
proportions are somewhat different; the head of a hatchling is
relatively large with blunt muzzle, and the tail is relatively short.
By the time a racerunner has emerged from its second hibernation,
late in its second year of life, it is of typical adult size (average
72.8 mm. snout-vent length in May and June). For successive
years of age (and with successively smaller samples) growth to
75.6 zt .78 mm., 78.9 mm., 81.5 mm., and 83 mm. was recorded.

To escape enemies racenmners depend primarily on speed, and
upon seeking shelter in dense vegetation or in their burrows. The
most important natural enemies are snakes, notably the racers,
Coluber constrictor and C. fiagellum. Hawks (only Buteo platyp-
terus definitely recorded), skunks, coyotes, armadillos, and even
box turtles are known to eat six-lined racerunners.

Little is known of the diseases or internal parasites that affect
the racerunner, although nematode worms {Pharyngodon warneri)
have been found in the gut. By far the most important external
parasite is the common pest chigger, Eutrombictila alfreddugesi.
Especially in early summer, a single racerunner may carry hun-
dreds of these parasitic mites, and the racerunner seems to be one
of the favorite hosts, although this particular chigger attaches to
a great many kinds of vertebrates. Locally, where it is abundant,
the racerimner may be important in maintaining the numbers of
chiggers.

The racerunner is active and persistent in seeking out its prey,
which consists mostly of small insects, but also includes spiders
and snails. Most food items are relatively small in proportion to
the size of the racerunner. Sense of smell aids the racerunner in

Six-lined Racerunner 59

obtaining food; prey is often found beneath the soil surface. Prey
does not have to be moving to attract the attention of the racerun-
ner as is the case with many other kinds of Hzards.

In a partly isolated colony occupying a .57 acre area (a 1000-
foot long dry ditch), population density (exclusive of hatchhngs)
varied between 40 per acre and 72 per acre. Sex ratio was ap-
proximately 1 : 1, and the population comprised seven successive
annual age groups (including hatchlings). Loss, through natural
mortality factors, of roughly half the individuals in each age group
in the course of a year is indicated.

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D

27-3118

' (Continued from inside of front cover)

19. A new chipmunk (Genus Eutamias) from the Black Hills. By John A.
White. Pp. 259-262. April 10, 1952.

20. A new piiion mouse (Peromyscus truei) from Durango, Mexico. By Robert
B. Finley, Jr. Pp. 263-267. May 23, 1952.

21. An annotated checklist of Nebraskan bats. By Olin L. Webb and J. Knox
Jones, Jr. Pp. 269-279. May 31, 1952.

22. Geographic variation in red-backed mice (Genus Clethrionomys) of the south-
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Pp. 281-292, 1 figure in text. November 15, 1952.

23. Comments on tlie taxonomy and geographic distribution of North American
microtines. By E. Raymond Hall and E. Lendell Cockrum. Pp. 293-312.
November 17, 1952.

24. The subspecific status of two Central American sloths. By E. Raymond Hall
and Keith R. Kelson. Pp. 313-337. November 21, 1952.

25. Comments on the taxonomy and geograpl;iic distribution of some North Amer-
ican marsupials, insectivores, and carnivores. By E. Raymond Hall and Keith
R. Kelson. Pp. 319-341. December 5, 1952.

26. Comments on the taxonomy and geographic distribution of some North Amer-
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27. A synopsis of the North American microtine rodents. By E. Raymond Hall
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28. The pocket gophers (Genus Thomomys) of Coahuila, Mexico. By RoUin H.
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29. Geographic distribution of the pocket mouse. Perognathus fasciatus. By
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30. A new subspecies of wood rat (Neotoma mexicana) from Colorado. By
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31. Four new pocket gophers of the genus Cratogeomys from Jalisco, Mexico.
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32. Genera and subgenera of chipmunks. By John A. White. Pp. 543-561. 12
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33. Taxonomy of the chipmunks, Eutamias quadrivittatus and Eutamias xan-
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34. Geographic distribution and taxonoiny of the chipmiinks of Wyoming. By
John A. White. Pp. 584-610, 3 figures in text. December 1, 1953.

35. The baculum of the chipmunks of western North America. By John A.
White. Pp. 611-631, 19 figures in text. December 1, 1953.

36. Pleistocene Soricidae from San Josecito Cave, Nuevo Leon, Mexico. By
James S. Findley. Pp. 633-639. December 1, 1953.

37. Seventeen species of bats recorded from Barro Colorado Island, Panama Canal
Zone. By E. Raymond Hall and William B. Jackson. Pp. 641-646. De-
cember 1, 1953.

Index. Pp. 647-676.
*Vol. 6. (Complete) Mammals of Utah, taxonomy and distributidn. By Stephen D.
Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952.
Vol. 7. *1. Mammals of Kansas. By. E. Lendell Cockrum. Pp. 1-303, 73 figures in
text, 37 tables. August 25, 1952.

2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry
S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 fi^es in text. August
24, 1953. '

3. The silkv pocket mice (Perognathus flavus) of Mexico. By RoUin H. Baker.
Pp. 339-347, 1 figure in text. February 15, 1954.

4. North American jumping mice (Genus Zapus). By Philip H. Krutzsch. Pp.
349-472, 47 figures in text, 4 tables. April 21, 1954.

5. Mammals from Southeastern Alaska. By Rollin H. Baker and James S.
Findley. ' Pp. 473-477. April 21, 1954.

6. Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479-
487. April 21, 1954. '

7. Subspeciation in the montane meadow mouse. . Microtus montanus, in Wyo-
ming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text.
July 23, 1954.

8. A new subspecies of bat (Myotis velifer) from southeastern California and
Arizona. By Terry A. Vaughn. Pp. 507-512. July 23, 1954.

9. Mammals of the San Gabriel mountains of California. By Terry A. Vaughn.
Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954.

10. A new bat (Genus Pipisbellus) from northeastern Mexico. By Rollin H.
Baker. Pp. 583-586. November 15, 1954. m

11. A new subspecies of pocket mouse from Kansas, i *ilBy E. Raymond Hall. Pp.
587-590. November 15, 1954. , , ,

12. Geographic variation in the pocket gopher, Cratogepmy? castanopsi in Coa-
huila, Mexico. By Robert J. Russell and Btollin JH'.- Baker. Pp. 591-608.
March 15, 1955. . . .,V V '' ■^. '; V'^/- i

u .■,;■. 'V '•

(Contiriued on outside of back cover)

(Continued from inside of back cover)

13. A new cottontail (Sylvilagus floridanus) from northeastern Mexico. By RoUin
H. Baker. Pp. 609-612. April 8, 1955.

14. Taxonomy and distribution of some American shrews. By James S. Findley.
Pp. 613-618. June 10, 1955.

15. The pifcmv woodrat, Neotoma goldmani, its distribution and systematic posi-
tion. By Dennis G. Rainey and RoUin H. Baker. Pp. 619-624, 2 figs, in
text. June 10, 1955.

Index. Pp. 625-651.
VoL 8. 1. Life history and ecology of the five-lined skink, Eumeces fasciatus. By Henry
S. Fitch. Pp. 1-156, 26 figs, in text. September 1, 1954.

2. Myology and serology of the Avian Family Fringillidae, a taxonomic study.
By William B. Stallcup. Pp. 157-211, 23 figiues in text, 4 tables. November
15, 1954.

3. An ecological studv of the collared lizard (Crotaphytus collaris). By Henry
S. Fitch. Pp. 213-274, 10 figures in text. February 10, 1956.

4. A field study of the Kansas ant-eating frog, Gastrophr>'ne olivacea. By Henry
S. Fitch. Pp. 275-306, 9 figures in text. February 10, 1956.

5. Check-list of the birds of Kansas. By Harrison B. Tordoff. Pp. 307-359, 1
figure in text. March 10, 1956.

6. A population study of the prairie vole (Microtus ochrogaster) in northeastern
Kansas. By Edwin P. Martin. Pp. 361-416, 19 figures in text. April 2,
1956.

7. Temperature responses in free-living amphibians and reptiles of northeastern
Kansas. By Henry S. Fitch. Pp. 417-476, 10 figures in text, 6 tables. June
1, 1956.

^ 8. Food of the crow, Corvus brachyrh\Tichos Brehm, in south-central Kansas. By

Dwight Piatt. Pp. 477-498, 4 tables. June 8, 1956.
9. Ecological observations on the woodrat, Neotoma floridana. By Henry S.
Fitch and Dennis G. Rainey. Pp. 499-533, 3 figures io text. June 12, 1956.
10. Eastern woodrat, Neotoma floridana: Life history and ecology. By Dennis
G. Rainey. Pp. 535-646, 12 plates, 13 figures in text. August 15, 1956.
Index. Pp. 647-675.

Vol. 9. 1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 fig-
ures in text. December 10, 1955.

2. Additional records and extensions of ranges of mammals from Utah. By
Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80.
December 10, 1955.

3. A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rol-
lin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.

4. Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming.
By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956.

5. The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6
figures in text. May 19, 1956.

6. Additional remains of the multituberculate genus Eucosmodon. By Robert
W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.

7. Mammals of Coahuila, Mexico. By RolUn H. Baker. Pp. 125-335, 75 figures
in text. June 15, 1956.

8. Comments on the taxonomic status of Apodemus peninsulae, with description
of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 337-346,
1 figure in text, 1 table. August 15, 1956.

9. Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp.
347-351. August 15, 1956.

10. A new bat (Genus Leptonycteris) from Coahuila. By Howard J, Stains.
Pp. 353-356. January 21, 1957.

11. A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico.
By Robert J. Russell. Pp. 357-361. January 21, 1957.

12. Geographic variation in the pocket gopher, Thomomys bottae, in Colorado. ^
By Phillip M. Youngman. Pp. 363-387, 7 figures in text. February 21,
1958.

13. New hog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones,
Jr. Pp. 385-388. May 12, 1958.

More numbers will appear in volume 9.

Vol. 10. 1. Studies of birds killed in nocturnal migration. By Harrison B. Tordoff and

Robert M. Mfngel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 1956.

2. Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.

3. The forest habitat of the University of Kansas Natural History Reser\'ation.
By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures
in text, 4 tables. BTecembir 31, 1956.

4. Aspects of reproduction and development in the prairie vole (Microtxis ochro-
gaster). By Henry S. Fitch. Pp. 129-161, 8 figures in text, 4 tables. Decem-
bef 19, 1957.

5. Birds found on the Arctic slope of northern Alaska. By James W. Bee.
Pp. 163-211, pis. 9-10, 1 figure in text, March 12, 1958.

Vol. 11. 1. The systematic status of the colubrid snake, Lcptiodeira discolor Giinther.
By William E. Duellman. Pp. 1-9, 4 figs. July 14. 1958.
2. Natural history of the six-lined racerunner, Cnemidophorus sexlineatus. By
Henry S. Fitch. Pp. 11-62, 9 figs., 9 tables. September 19, 1958.

Mos. mm. 2801

:TBg^;i959

University of Kansas Publications , ;;

j [^' . rf

Museum of Natural History

Volume 11, No. 3, pp. 63-326, 6 pk, 24 figs., 3 tables
December 12, 1958

Home Ranges, Territories,

and Seasonal Movements of Vertebrates

of the Natural History Reservation

BY
HENRY S. FITCH

University of Kansas

Lawrence

1958

UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY

Institutional libraries interested in publications exchange may obtain this
series by addressing the Exchange Librarian, University of Kansas Library,
Lawrence, Kansas. Copies for individuals, persons working in a particular
field of study, may be obtained by addressing instead the Museum of Natural
History, University of Kansas, Lawrence, Kansas. There is no provision for
sale of this series by the University Library which meets institutional requests,
or by the Museum of Natural History which meets tlie requests of individuals.
However when individuals request copies from the Museum, 25 cents should
be included, for each separate number that is 100 pages or more in length, for
the purpose of defraying the costs of wrapping and mailing.

* An asterisk designates those numbers of which the Museum's supply (not the Li-
brary's supply) is exhausted. Numbers published to date, in this series, are as follows:

Vol. 1, Nos. 1-26 and index. Pp. 1-638, 1946-1950.

•Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444,
140 Egurcs in text. April 9, 1948.
Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, and distribution. By Rol'
lin H. Baker. Pp. 1-359, 16 fig\ures in text. June 12, 1951.

*2. A quantitative study of the nocturnal migration of birds. By George H.
Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951.

3. Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473-
530, 49 figures in text, 13 tables. October 10, 1951.

4. Birds from the state of Veracruz, Mexico. By George H. Lower>', Jr. and
Walter W. Dalquest. Pp. 531-649, 7 figiures in text, 2 tables. October
10, 1951.

Index. Pp. 651-681.
•Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates,
31 figures in text. December 27, 1951.

Vol. 5, Nos. 1-37 and index. Pp. 1-676, 1951-1953.

•Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. Bv Stephen D.
Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952.

Vol. 7. *1. Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, 73 figures in
text, 37 tables. August 25, 1952.

2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry
S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 figures in text. Augujrt
24, 1953.

3. The silkv pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker.
Pp. 339-347, 1 figiue in text. February 15, 1954.

4. Nortli American jumping mice (Genus Zapus). By Philip H. Krutzsch. Pp.
349-472, 47 figures in text, 4 tables. April 21, 1954.

5. Mammals from Southeastern Alaska. By Rollin H. Baker and James S.
Findley. Pp. 473-477. April 21, 1954.

6. Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479-
487. April 21, 1954.

7. Subspeciation in the montane meadow mouse. Microtus montanus, in Wyo-
ming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text.
July 23, 1954.

8. A new subspecies of bat (Myotis velifcr) from southeastern California and
Arizona. By Terry A. Vaughn. Pp. 507-512. July 23, 1954.

9. Mammals of the San Gabriel moimtains of California. Bv Terry A. Vaughn.
Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954.

10. A new bat (Genus Pipistrellus) from northeastern Mexico. By Rollin H.
Baker. Pp. 583-586. November 15, 1954.

11. A new subspecies of pocket mouse from Kansas. By E. Raymond Hall. Pp.
587-590. November 15, 1954.

12. Geographic variation in the pocket gopher, Cratogeoniys castanops, in Coahuile,
Mexico. By Robert J. Russell and Rollin H. Baker. Pp. 591-608. March
15, 1955.

13. A new cottontail (Svlvilagus floridnnus) from northeastern Mexico. By Rollin
H. B.iker. Pp. 609-612. April 8, 1955.

14. Taxonomy and distribution of some American shrews. By James S. Findley.
Pp. 613-618. June 10, 1955.

(Continued on inside of back cover)

University of Kansas Publications
Museum of Natural History

Volume 11, No. 3, pp. 63-326, 6 pis., 24 figs., 3 tables
December 12, 1958

Home Ranges, Territories,

and Seasonal Movements of Vertebrates

of the Natural History Reservation

BY

HENRY S. FITCH

University of Kansas
Law^rence

1958

Untvehsity of Kansas Publications, Musexjm of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 11, No. 3, pp. 63-326, 6 pis., 24 figs, in text, 3 tables
Published December 12, 1958

University of Kansas
Lawrence, Kansas

PRINTED IN

THE STATE PRINTINS PLANT

TOPEKA, KANSAS

I9E8

fm. GOMF. ZOOL

LfiRARV

FEB 2 1959

Home Ranges, Territories,

and Seasonal Movements of Vertebrates

of the Natural History Reservation

BY

HENRY S. FITCH

CONTENTS

PAGE

Introduction 65

Methods of study 68

Population densities 75

Habitats 82

Accounts of species 86

fishes 86

amphibians 88

reptiles 98

mammals 129

birds 170

Summary and conclusions 317

Literature cited 322

INTRODUCTION

In July, 1948, ecological investigations were initiated on the newly
created University of Kansas Natural History Reservation, a 590-
acre division of the original "Robinson Farm." Although all phases
of ecology have been given some attention, greatest eflFort has been
devoted to autecological studies of the more abundant kinds of
vertebrates.

Invariably in such studies a major amount of time and effort is ex-
pended in an investigation of spatial relationships. In order to
understand the ecology of a species it is necessary to recognize in-
dividuals and to trace individual histories. It is necessary to ascer-
tain how the typical individual animal is Hmited in time and space in
order to understand the population dynamics of the species. As
stated by Dice (1952: 231) most individual animals do not wander
about at random, but the great majority spend their lives in closely
limited areas, each confining his movements to a small part of the

(65)

66 UNi\'ERSiTi' OF Kansas Publs., Mus. Nat. Hist.

habitats that are fully suited to his existence, and over which he is
potentially free to roam.

From a practical standpoint it is important to understand the
daily and seasonal movements of an individual, and the spatial
requirements of a breeding population for those species that have
major economic bearing. Such information serves as a basis for
control measures for tliose kinds that are harmful, and as a basis for
encouragement through positive management for those kinds that
are beneficial.

In the course of routine field work, information concerning move-
ments has been obtained for most of the species of vertebrates re-
corded from the Reservation. However, the quantity and quality
of the information obtained varies greatly from species to species.
Ideally the information obtained for each species would have
answered the following questions: What is the size of a typical
home range and how much does it vary in size and shape? Is the
home range permanent; if not, how much and how often does the
animal alter it? How is the home range afiEected by weather, food
supply, and population density? What are the essential habitat
features that a home range must include? How do home ranges
differ between male and female, between adult and young, or be-
tween any other special groups within the species? Is the home
range in whole or in part defended as a territory? If so, what classes
of individuals (according to size, age, or sex) are active in such de-
fense, and what classes of individuals ( or what other species of ani-
mals) are objects of territorial hostility? Does the male share his
territory with a mate or with a family group? Is the territory a
phenomenon of the breeding season, or is it maintained throughout
the year?

For none of the species studied was the information obtained
sufiBciently extensive to answer all these questions completely. For
the rarer and more elusive species, the data obtained are scanty.
Nevertheless, for many of the more abundant species, the essential
items of information have been obtained.

Birds as a group have been studied in the past much more exten-
sively than the other groups of vertebrates, with regard to spatial
relationships. In general birds are more easily studied, because
they can be more readily observed. On the Reservation, observing
birds was unusually difiBcult because of the dense sheltering vege-
tation, and the shyness of most species. Attempts to trace move-
ments of individuals were often unsatisfactory. Because of the

Home Ranges of Vertebrates 67

large number of species present, the time available to devote to any
one species was limited. The mammals, and especially the reptiles
and amphibians, have been studied less, heretofore, and I made
concentrated efiForts to secure information concerning those kinds
that were abundant and available.

In the course of my field study, spanning more than nine years,
emphasis has shifted from time to time. Short lines of live-traps for
small mammals were first operated in 1948, and these operations
were much expanded in the spring of 1950. Live-trapping of rep-
tiles and amphibians was begun, on a small scale, in the summer
of 1949, and was gradually expanded. Observations on birds were
most concentrated in the years 1952 through 1955. Many of my
data concerning spatial relations of the narrow-mouthed toad, five-
lined skink. Great Plains skink, collared lizard, opossum, coyote,
prairie vole, eastern woodrat, and summer tanager have already
been published. The manuscript resulting from the present study
was completed in essentially its present form in 1956, but significant
items of data obtained since then have been incorporated. The spe-
cies accounts of amphibians and reptiles were written first, on the
basis of data on hand on July 1, 1955. Although more data were
collected subsequently for most species, the new figures were in-
corporated into the account only in those instances where they
seemed to alter substantially the trends previously revealed.

A faculty grant for the years 1956-1957, and 1957-1958 has aided
substantially in the completion of this project. Many persons have
contributed to the work in various ways. Richard E. Freiburg
(1951), Edwin P. Martin (1956), Robert L. Packard (1956) and
Dennis G. Rainey ( 1956 ) have published separate reports concern-
ing several species of vertebrates studied by them on the Reserva-
tion, and I have drawn freely upon their findings. Other persons who
have contributed data or have assisted me with field work include:
Maurice F. Baker, WilHam N. Berg, Alice V. Fitch, John H. Fitch,
Richard W. Fredrickson, Richard D. Harder, the late Robert M.
Hedrick, Donald W. Janes, Richard B. Loomis, John Poole, Louis
M. Sandidge, and my wife, Virginia R. Fitch who also typed much
of the manuscript. Mr. Robert M. Hedrick, Mrs. Connie Wynkoop,
and Mrs. Lorna Cordonnier completed the illustrations. Dr. E.
Raymond Hall, Director of the University of Kansas Museum of
Natural History, has provided valuable counsel and suggestions at
various stages of the work, and has made available the services
of the Museum staff artists.

68 University of Kansas Publs., Mus. Nat. Hist.

To supplement my original findings a large amount of published
literature has been examined, and the findings of other authors are
often cited in the species accounts for purposes of contrast or com-
parison. However, because of the large number of species involved,
a thorough survey of the literature for all was not feasible. For the
birds, especially, many pertinent published papers probably have
been overlooked.

METHODS OF STUDY

To facilitate ecological studies the University of Kansas Natural History
Reservation has been subdivided into 80 separate areas, having as their
boundaries, creeks, ravines, wire fences, rock walls, roads, trails, and, in some
instances, imaginary lines. Each subdivision has within it numerous specific
landmarks. Most of these are trees, but boulders, fence posts, logs and
other objects, even metal stakes especially placed for the purpose, have also
been utilized. Each record of an individual animal was located with reference
to some such named landmark. Distances sometimes were actually measured
with a steel tape, but nonnally they were estimated to the nearest ten-foot inter-
val. In studies of small mammals, traps were usually laid out in grids ( 50-foot,
30-foot and 20-foot intervals were used) with each position measured from a
definite landmark. Many of the landmarks were discernible on an aerial photo-
graph made on the scale of one inch to a hundred feet. A map on the same
scale, with two-foot contour intervals, was also available.

Various methods were used, depending on the species, in determining the
area covered by an individual animal. For a small minority of kinds it was
possible to keep an individual under continuous observation as it moved about,
carrying on its normal activities. The male dickcissel is perhaps the best
example. As he moves about his territory, with frequent shifts, usually perch-
ing on tlie topmost twigs of the largest shrubs or saplings, he soon reveals the
approximate extent of his territory. An individual may be recognized on
successive occasions, even though he is not distinctively marked, by his habit
of returning regularly to the same favorite perches.

The same method also applied well to the field sparrow, but was more difficult
because this species more often stayed in concealing cover, because the sexes
could not be readily recognized, and because the species was so abundant that
confusion might often arise concerning the identity of an individual that was
momentarily out of sight, or that encountered another.

Most other kinds of birds were less readily kept under observation, because
they were more wary or more secretive in their behavior. For many kinds,
including the wood thrush, the Kentucky warbler, and the several species of
vireos, the song of the male provided the chief basis for tracing movements
since the birds themselves were glimpsed only occasionally. However, under
favorable conditions, the frequent and regular bursts of song by the bird as
it moved about slowly in its foraging, served to keep the observer informed
as to its course. Little was learned concerning the movements of the females
in these particular species, but in each instance it is known that the members
of a pair are the joint occupants of a territory.

Home Ranges of Vertebrates 69

In many of the other species of birds that were studied, a combination of
sight and sound was used in keeping the individuals under observation and
tracing their movements. In some instances distinctive artificial marking of
individuals was an important aid. Besides the regular metal bands, provided
by the U. S. Fish and Wildhfe Service, colored celluloid bands, or dyed
feathers crimped onto the tail, or colored nylon ribbons, were used for sight
identification. In practice positive sight identification of individuals was made
only rarely, even with the aid of these various devices. The nylon ribbons
were most eflFective. Each was looped around the tarsometatarsus and sewn
snugly in place with a free end of approximately half an inch projecting out
behind, like the jess on a falcon. This colored ribbon, presenting several
times as much surface as an ordinary leg band, could be distinguished at a
greater distance. Most sight records from colored ribbons were obtained
for red-bellied woodpeckers, downy woodpeckers, and field sparrows.

The same types of ribbons, attached to the ears of cottontails (Janes, MS),
often served for sight records of these animals. Colored collars and ear disks
on gray squirrels and fox squirrels served the same purpose (Packard, 1956: 9).
For the six-lined racerunner, paints of various colors, carefully applied on dif-
ferent parts of the body surface, served effectively to render individuals dis-
tinguishable in the field for short periods. Through bleaching and wear these
colors rapidly dulled and after a few days they were no longer distinguishable.
The addition of bright colors, unnatural to the species involved in the attach-
ment of ribbons, bands, feathers, or paint, may of course, have rendered the
animals more susceptible to certain predators, and also may have affected
their social status adversely, reducing their chances of survival. Actually, no
evidence of such adverse effects was noticed, and it seems safe to assume that
the over-all ecology, and especially the movements, were not altered to any im-
portant degree, in most species at least.

For most kinds of mammals, reptiles, and amphibians, and some kinds of
birds, individual identification could be made with certainty only by recaptur-
ing and handling the animal. Many kinds are too wild, secretive, or nocturnal
to observe in the field. For birds, trapping was carried on principally in
winter. Procedure was not uniform, because of variation in the number of
traps and time available. Ordinarily 30 or more traps were set at fairly imi-
form intervals in a hue extending from the Reservation headquarters for a
quarter mile or more in one direction, then looping back to the starting point.
From time to time the trap hne was altered by moving one leg of it beyond
the other, and in the course of an entire winter the whole trap line was rotated
through 360 degrees about the central point of the headquarters. In the course
of a season, therefore, traps were well distributed over an area as large as 125
acres. For individuals ranging entirely within the area encompassed by the
trap lines the pattern of records obtained might show the extent of home
range (or territory) fairly well.

Most of the mammals recorded were caught in live-traps set in a grid pat-
tern or in parallel lines; the interval between traps was varied from time to
time and place to place, from ten to 120 feet. Mammals were marked by a
combination of ear-clipping and toe-clipping, or by toe-cHpping alone (Fitch,
1952: 32-33), in those species having small and inconspicuous ears. In the

70 Unimersity of Kansas Publs., Mus. Nat. Hist.

latter (Microtus ochrogaster and M. pinetorum) one toe was clipped on each
of four feet (excluding the minute "thumbs") and a total of 400 combinations
were possible without duphcation. In others only two toes were clipped in
combination with a distinctive ear mark. Because of the short life span of
small mammals in general, the animal marked with any one formula had died
long before there was need to use the same combination again. For some of
the most abundant species tlie same series were used repeatedly.

Toe-clipping was used also in the marking of frogs, toads and lizards. These
animals were secured by a variety of techniques. Many were caught by hand.
Others were taken in strategically placed funnel traps or pitfalls. The captures
in funnel traps and pitfalls were less randomly distributed than the records ob-
tained otherwise, as tliey were mainly along rock ledges or other natural ob-
stacles that might afiFect the movements of the animals. For the glass Uzard
the marking technique consisted of removing three scales from the edge of the
lateral fold, from which a formula was derived. The nick from the most an-
terior excision served as a base mark from which the positions of the other two
nicks could be recorded according to the number of scales between.

Snakes were marked by excising subcaudals in different combinations, count-
ing backward from the base of the tail. Only the proximal portion of the tail
was used, up to subcaudal number 20. When all possible combinations were
exhausted, they were used again in combination with ventral scutes cHpped
on either right or left sides. Snakes were obtained by various methods. Some
were found by chance in the course of routine field work, some were discovered
by raising sheltering objects such as rocks and boards beneath which they had
taken cover, and some were located by attention to the calls of birds, mammals
or frogs caught or pursued by the snakes or by the calls of blue jays and cer-
tain other birds that harrassed the snakes. Even more snakes were caught in
funnel traps. Success with these traps was greatest in September, October,
and sometimes early November, along the hilltop rock outcrops where many
snakes congregated to hibernate. So far as known, there were no major "dens"
on the area. A snake having gained a rock outcrop might travel along it
a longer or shorter distance, and might retreat into any deep crevice favorable
for hibernation, perhaps solitarily or perhaps in company with several others of
of its own species and other species. Attempts to trap snakes in these same
situations in spring, after their emergence from hibernation, were notably im-
successful. At this time of year there was rapid dispersal into other habitats.
In summer, funnel traps were set along specially constructed screen drift fences,
in grassland, thickets, and at woodland edge.

From this account it is evident that in the course of the study significant
movements of animals were actually observed only on rare occasions. For the
most part movements had to be reconstructed from the fragments of informa-
tion obtained by occasional captvires, in live-traps or otherwise. Obviosuly, in
most instances the animal did not move directly from one recorded point to
another, but usually covered a much greater distance, coming to the second
point by a roundabout route. When two or more such locations were recorded
for an individual within a short time, the assumption seemed justified that they
were probably within a home range (or territory) and partly defined its extent.
However, when much time had elapsed between two consecutive records, there
was no assurance that the animal had not shifted more or less permanently. Ex-

Home Ranges of Vertebrates

71

tensive comparison of long-term and short-term records was necessary to estab-
lish the probabihty of shifts in range.

In general, any point where an animal is recorded within its home range or
territory is located at random with reference to any other point of record, but
there are many exceptions. If records are based on Hve-trapping, trap habit
may become an important factor altering the behavior of the individual, and
may cause it to return regularly to one or a few traps while neglecting to visit
other parts of its home range. An animal that has a regular home base rehed
upon for shelter and escape, such as the home burrow of a vole, the stick house
of a woodrat, or the home blackberry patch of a cottontail, naturally tends to
stay within easy reach of this shelter, and in outlying portions of its range, ac-
tivities become increasingly diffuse. For a parent bird, the nest or yoimg simi-
larly provide a focal point of attraction, and activity tends to be much more
concentrated near the nest than it is in remote parts of the territory.

Other animals actually tend to move about more or less at random over their
territories or home ranges. Tracks of a foraging opossum, for instance, show
that the animal wanders circuitously in what seems to be an aimless course.
Opossums hve-trapped show little or no tendency to return to any one trap
regularly. A male wood thrush, singing as he forages, seems to cover his terri-
tory rather uniformly. In many other kinds of birds, the male uses a series of
territorial perches for singing, and these are usually well distributed over the
area to which his activities are confined.

The term "minimum home range" has been widely used in studies of small
mammals, in recognition of the fact that the area disclosed by investigation
nearly always is somewhat smaller than the total area utiUzed, because data

Fig. 1. Hypothetical home ranges of the same size, showing patterns of
records for: (A) an animal that nas a central home base and uses the vari-
ous parts of its range in inverse proportion to their distance from the home
base; and ( B ) an animal that wanders at random over its area.

72 UNivERSiTi' OF Kansas Publs., Mus. Nat. Hist.

are incomplete in varying degrees. Thus a minimum home range could be
plotted on tlie basis of only three records, providing that all three were at
separate locations, and that these locations did not fall in a straight line. How-
ever the triangular area encompassed by the three locations would almost cer-
tainly amount to only a small part of the total range. A fourth record of the
same individual might fall within the original triangle, or it might fall outside,
increasing the minimum home range and altering it to quadrangular form. Ob-
viously many more records would be needed before tlie minimum home range
disclosed would be expanded to an approximation of the entire home range
occupied by the individual. Odum and Kuenzlcr ( 1955 ) have presented a
scholarly discussion of this problem, showing that the rate of increase is, on the
average, reduced with each additional record, until it becomes insignificantly
small. For practical purposes Odum and Kuenzler advocated accepting as
complete a minimum home range in which the increase for each additional
record is below the one per cent level. Their procedure was to connect out-
lying points with hnes and measure the enclosed area after each series of ten
records. When such a series of ten added less than ten per cent to the pre-
viously plotted range, they considered addition of further records superfluous.
As demonstrated by Odum and Kuenzler, the number of records required to
achieve this one per cent level varied widely according to the species and also
varied within the species, according to the stage of the nesting cycle and other
factors. In a pair of chipping sparrows that were feeding nestlings, the one
per cent level was obtained by means of only 20 observations. EarUer, in the
nest-building stage, 55 records of the same pair of birds were necessary to
achieve the one per cent level, and the area covered was nearly three times as
large as it was in the nesthng stage.

Using a wood pewee as an example, Odum and Kuenzler (op. cit.: 131)
show the increasing size of minimum home range (actually a territory in this
instance) with the addition of successive tens of records. The one per cent
level was attained after the fourth series of ten records. The polygon outlining
the bird's known range at this stage had nine marginal points (each repre-
sented by a single record). The other records, inside the polygon, tended to
be clustered at a few favored locations. In tliis respect the pewee's pattern of
records resembles those of various other animals, including the small mammals
that are taken in live-traps exclusively.

The wood pewee and chipping sparrow studied by Odum and Kuenzler, are,
of course, almost ideal subjects for such observation. For such active and
vocal small birds that are not especially wary of humans many location records
can be obtained in a short period of observing. In studies of small mammals,
tliousands of trap nights may not jaeld a comparable set of records for any one
individual. In only a few instances was I able to obtain records of individual
movements as complete as the set shown for the pewee by Odum and Kuenzler.
Because most animals tend to return frequently to favored spots, the number
of marginal locations, rather than the total number of records, or even the
total number of points, is the best indication of reliability in the minimiun home
range plotted. While a home range plotted on the basis of only three or four
points is almost sure to be much smaller than the true home range, one
plotted from six ]wints may outhne the range fairly accurately, especially if
»lie points are well spaced along the margin. Home ranges based on seven

Home Ranges of Vertebrates 73

or more marginal points are considered to be sufficiently representative of the
area covered by the animal.

As numbers of marginal points increase with more complete records, the
range plotted generally tended tovi'ard a circular or elliptical shape. If only
marginal points are connected, there are no concavities in the outline of the
polygon representing the home range. Concavities in the outHne of the home
range polygon are permissible when uninhabitable areas extend within it.
For example water may extend into the home range of a terrestrial animal, or
forest may extend into the home range of an animal closely hmited to grassland.
The irregular borderline between forest and grassland resulted in many home
ranges of irregular shape among those that were plotted, both for species
characteristic of forest and those typical of prairie habitat.

Because a large number of records are necessary to plot the home range
of any individual with reasonable completeness, according to this method,
relatively few of the animals recorded are usable. In my study, an alternative
method of determining size of home range was used, which permitted inclusion
of individuals even when only two records were available. This method was
based on the theory that any two locations where an individual was recorded in
its range, were situated at random with respect to each other. Distances
between successive points of record were averaged for all individuals of a
species, or according to sex, age group or season. The average figure obtained
in each instance was considered representative of a home range radius. The
home range is visuaUzed as tending toward a circular shape, and random points
within it will be separated by, on the average, half the diameter of the area.
In an animal the range or territory of which has one or more focal points, where
activity tends to be concentrated, the distances between randomly selected
points will average less than the home range radius reflecting the extent to
which activity is concentrated within one part of the total area.

From the average distance between successive locations recorded, tlie area
covered was computed from the fomiula -m- or the area of a circle. Obviously
this method is subject to various sources of error; the home range nearly always
deviates from a circular shape, and usually some parts of its are used more than
others. The advantages of this method are ( 1 ) that it makes possible a calcula-
tion of approximate size of home range on the basis of a relatively small amount
of data, ( 2 ) that it provides a check on size of area plotted as a minimum home
range from actual points of capture.

Still another method of computing area covered, is applicable specifically to
territorial birds. This method involves measuring the distances between males
that are singing simultaneously and answering each other, in situations where
the habitat is continuous. Under such conditions the territories presumably
have a common boundary. The distance between two singing males may be
considered representative of the distance from one territorial center to another,
the equivalent of a territory's diameter. A possible source of error in tliis
method is that males happening to be in adjacent parts of their territories may
be stimulated to competitive singing in defense of their boundaries, but it
seems doubtful that this was an important factor in any of my records. In
most instances when distances were recorded, singing was general and the
males tended to be rather uniformly spaced. The method provided an im-
portant means of checking the figures obtained from the two methods previously
discussed.

74

University of Kansas Pi^bls., Mus. Nat. Hist.

Fig. 2. Map showing areas where field work was done: (A) the 160-
acre Rockefeller Experimental Tract, and (B) the 59()-acre University of
Kansas Natural History Reservation, showing distribution of weedy fields,
grassland and woodland. The grassland includes prairie, both original and
reestablished, dominated by blucstems and other tall grasses. Former pas-
tures are mostly dominated by awnless brome.

Home Ranges of Vertebrates 75

In the following species accounts, binomials are used for the lands of animals
discussed. Subspecific identities can be determined from the appropriate
reference among the following: Smith, 1956 (reptiles and amphibians);
Tordoff, 1956 (birds); and Hall, 1956 (mammals).

This publication is intended to serve, incidentally, as a list of the vertebrates
recorded from the Reservation. Therefore, all species known from the area
are listed, although for certain kinds so httle information has been obtained
that their inclusion would not be justified otherwise. The present list revises
and extends one published earlier (Fitch, 1952:23-29). The king snake
( Lampropeltis getulus) was included in the earher list on the strength of an
individual found on the county road approximately half a mile south of the
Reservation. However, the habitat there was somewhat diflFerent and the
species still has not been definitely recorded from the Reservation. It is
therefore deleted from the present list. Besides the species added to the
original hst, or deleted from it, certain other species have been found to have
somewhat different status from that originally attributed to them. Domestic
animals are not included in the following lists and species accounts. It is
the poHcy to exclude them from the Reservation; nevertheless, domestic birds
and mammals play some part in the ecology of the area. A colony of the
domestic pigeon (Columba livia) has persisted into 1958 in the old bam on
the Rockefeller Tract, but their effect on the ecology of the area seems slight,
except as they may compete for nest sites with other species that seek an
edificarian habitat. Their foraging is carried on chiefly in cultivated areas
remote from the nest sites. Cats (Felis domesticus) and dogs (Canis fa-
miliaris) from nearby farms cannot be entirely excluded from the area, and
prey upon various kinds of native vertebrates. Livestock, including cattle
{Bos taurus), horses (Equus cahallus), pigs {Sus scrofa) and sheep {Ovis
aries) played an important part in the trends of succession on the area before
it was protected from them. They still stray onto the Reservation from time
to time, and obviously their presence on adjoining farms affects the native fauna.

POPULATION DENSITIES

Over the period of my study every species of vertebrate on the
Reservation has undergone great changes in numbers. For most
species no precise figures for the entire area have been obtained,
even for any one time. The population changes have been actually
measured in relatively few instances.

In most instances the figures listed below represent preliminary
estimates. For convenience in this discussion, I have divided the
species of vertebrates of the Reservation into five main groups as
regards their occurrence. These groupings are entirely arbitrary.
They overlap each other, and many species constitute borderline
cases between two or more groups. Group 1 includes species re-
corded from the Reservation only once or on but few occasions, and
ordinarily not present there. These kinds are of negligible im-
portance in the ecology of the area. Group 2 also consists of species

76 Univtrsity of Kansas Publs., Mus. Nat. Hist.

that ordinarily do not occur on the Reservation, although they ap-
pear there as wanderers or stragglers with some regularity. On tlie
whole, this group too is ecologically unimportant, although in some
instances, as when a great blue heron stops at the pond for a period
of days and eats large numbers of frogs, the fauna may be appre-
ciably aflFected. Group 3 consists of species that reside on the area
( either year round or at one time of year ) but are represented there
by few individuals. In general these species also are unimportant
ecologically because of their small numbers and biomass, but the
larger carnivores and raptors may exert important effects on the
ecosystem even though only a few are present. Group 4 consists
of species which are usually not present on the area, but which
make brief seasonal visitations usually in substantial numbers.
Species of this group are obviously of greater importance in the
ecology of the area than those of the three groups previously men-
tioned. Group 5 includes those species which regularly inhabit
the area (either year round or for part of the year) in substantial
numbers. They are the species which have greatest influence on
the ecology of the area. It is for the Group 5 species, primarily,
that I have attempted to obtain census figures.

In the following lists, the 12 species of birds marked with aster-
isks in Group 1 and Group 2 are those that have been seen flying
over the Reservation but are not known to have stopped there. The
census figures listed for Group 5 have been obtained by various
methods. The only attempt to obtain complete counts of individuals
for the entire Reservation was limited to the birds, and was made
in the week January 29 to February 4, 1954. Numbers of some
kinds of birds were unusually low at the time of this census, and
the counts obtained may be regarded as minimum figures in most
instances. For various kinds of reptiles and mammals the census
figures cited are mostly based upon relatively small study areas. In
some instances these sample aieas are typical of major habitat
divisions on the Reservation, and the figures obtained may be
projected to show an approximation of the numbers on the entire
area. In other instances (collared lizard, six-lined racerunner,
Plains harvest mouse) the small sample areas where high concen-
trations were found were almost the only places where the species
occurred locally. For many of the more secretive species, direct
counts were impracticable, and the figures cited are based on the
ratios of new to marked individuals recorded at various stages in
the course of field work, with due regard for population turnover
through mortality, reproduction, and individual shifts.

Home Ranges of Vertebrates

77

Group 1. (species seldom present)

Black Bullhead

Green Sunfish

Tiger Salamander

Eastern Box Turtle

Painted Turtle

Franklin's Ground Squirrel

Woodchuck

Gray Fox

Pied-billed Grebe

White Pehcan *

Black-crowned Night Heron

American Bittern

Pintail

Goshawk

Rough-legged Hawk

Bald Eagle *

Prairie Falcon *

Sandhill Crane

Virginia Rail

Sora

American Woodcock

Wilson Snipe
Upland Plover
Spotted Sandpiper
Sohtary Sandpiper
Lesser Yellow-legs
Franklin Gull *
Bam Owl
Saw-whet Owl
Yellow-bellied Sapsucker
Scissor-tailed Flycatcher
Yellow-bellied Flycatcher
Olive-sided Flycatcher
Bank Swallow
White-breasted Nuthatch
Red-breasted Nuthatch
Winter Wren
Short-billed Marsh Wren
Ohve-backed Thrush
Gray-cheeked Thrush
Veery

Yellow-throated Vireo
Solitary Vireo
Black and White Warbler
Nashville Warbler
Black-throated Green Warbler
Blackbumian Warbler
Blackpoll Warbler
Oven-bird

Northern Water-thrush
Louisiana Water-thrush
Canada Warbler
American Redstart
Yellow-headed Blackbird *
Orchard Oriole
Scarlet Tanager
Painted Bunting
Grasshopper Sparrow
White-crowned Sparrow
Fox Sparrow
Swamp Sparrow

Fathead Minnow
Great Blue Heron
Green Heron
Canada Goose *
White-fronted Goose *
Snow Goose *
Blue Goose *
Mallard

Green-winged Teal
Blue-winged Teal
Shoveller
Wood Duck

Group 2. (species occasionally

Ring-necked Duck
Lesser Scaup
Sharp-shinned Hawk
Marsh Hawk
Sparrow Hawk
Coot
Killdeer

Black-billed Cuckoo
Nighthawk
Belted Kingfisher
Alder Flycatcher
Least Flycatcher

present )

Rough-winged SwaUow *
Chff Swallow *
Purple Martin *
Mockingbird
Hermit Thrush
Cedar Waxwing
Orange-crowned Warbler
Baltimore Oriole
Vesper Sparrow
Chipping Sparrow

Group 3. ( species with few individuals )

Snapping Turtle
Prairie Skink
Common Water Snake
Smooth Earth Snake
Prairie King Snake
Milk Snake

Slender Flat-headed Snake
Timber Rattlesnake
Plains Pocket Gopher
White-tailed Deer
Coyote

Group 4.

Plains Spadefoot Toad

Eastern Kingbird

House Wren

Catbird

Robin

Golden-crowned Kinglet

Ruby-crovraed Kinglet

Starling

Long-tailed Weasel
Red Fox
Muskrat
Turkey Vulture
Cooper Hawk
Red-tailed Hawk
Red-shouldered Hawk
Broad-winged Hawk
Screech Owl
Homed Owl
Barred Ovvl

( common species absent for most

Warbling Vireo
Tennessee Warbler
Yellow Warbler
Myrtle Warbler
Yellow-throat
Wilson Warbler
Western Meadowlark
Rusty Blackbird

Long-eared Owl
Chuck-will's-widow
Chimney Swift
Ruby-throated Hummingbird
Wood Thrush
Loggerhead Shrike
Long-tailed Chat
Red-winged Blackbird
Blue Grosbeak
Lark Sparrow

of annual cycle)

Bronzed Crackle
Rose-breasted Grosbeak
Purple Finch
Pine Siskin
Leconte Sparrow
Clay-colored Sparrow
White-throated Sparrow
Lincoln Sparrow

78 Uni\'ersity of Kansas Publs., Mus. Nat. Hist.

Group 5. (species of major importance)

American Toad: In both 1955 and 1956, 25 males were caught and marked
in tlie chorus at the pond. From the ratios of marked individuals to others,
it was evident that many more vi'ere present which were not recorded. The
breeding aggregation at the pond represented only a part of the population
on the Reservation. Normally the population probably fluctuates from a low
point of somewhat more tlian 100 to a maximum of many thousands or even
tens of thousands (including recently metamorphosed young).

M^oodhouse's Toad: The number has fluctuated from none to perhaps sev-
eral hundred young.

Cricket Frog: Several hundred adults usually are present at the pond and
the small creek. At times this number may expand to many thousand.

Gray Treefrog: In early summer, breeding aggregations of more than 100
adults normally are present at the pond.

Chorus Frog: Breeding aggregations at the pond have varied from as few
as ten to more than 100. In the breeding season the numbers may increase
to many thousands.

Great Plains Narrow-mouthed Toad: This toad is one of the most numerous
species; its breeding populations normally amount to several hundred and
increase to many thousands, perhaps tens of thousands, in summer after a suc-
cessful reproductive season.

Bullfrog: Numbers of adults and well grown young have fluctuated from
none to probably more than a thousand.

Leopard Frog: Numbers have fluctuated from few or none to probably
several thousand.

Western Box Turtle: Population density of this species is estunated as one
to 13 acres, on the basis of numbers found each year in the northwestern
quarter-section of the Reservation, and the ratio of new to recaptured indi-
viduals.

Collard Lizard: Over the sL\-year period that the colony on the Reserva-
tion survived there were on the average, 3.6 adults in the area occupied at the
quarry, a httle larger than half an acre.

Slender Glass Lizard: The population normally amounts to several per acre
in the more favorable grassland areas.

Six-lined Racerunner: Over a four-year period population density averaged
56 per acre on a half-acre area of favorable habitat, the ditch in the headquar-
ters area.

Ground Skink: The population density on the Reservation is low. Only a
few are seen each season and it seems probable that there is only one to sev-
eral acres of woodland.

Five-lined Skink: Population density was calculated as 67 per acre in an
unusually favorable woodland area of 2)4 acres where this species was most in-
tensively studied.

Great Plains Skink: Population density was calculated as 16 or 17 per acre
on a four-acre area of rocky upper slope where these skinks were intensively
studied.

Brown Snake: Density is probably only one to several acres over most
wooded areas of the Reservation.

Home Ranges of Vertebrates 79

Common Garter Snake: Over the nine-year period of my study numbers
caught annually averaged 23 (11 to 35) and relatively few marked snakes
were recaptured. A population of one to many acres is indicated.

Ring-necked Snake: In 1950, 47 were collected on a study area of a Uttle
more than four acres. Populations exceeding ten per acre probably occiu: over
extensive areas of the Reservation.

Worm Snake: In 1951, 51 worm snakes were collected on a study area of
25* acres, and the ratio of recaptures indicated that this was only a part of the
population. Such high concentrations, exceeding 20 per acre, are limited to
relatively small areas of favorable habitat.

Yellow-bellied Racer: In 1957, 36 were collected in a grassland area of 25
acres, and these represented the majority of the population present. Popvda-
tions of from one to two per acre for grassland habitats are indicated.

Black Rat Snake: The population density for the Reservation is calculated
as perhaps one adult to ten acres.

Bull Snake: The number resident on the Reservation is normally fewer
than ten.

Copperhead: A population of one to several acres is indicated.

Opossum: Population densities of from one in 20 acres to one in 40 acres
are indicated.

Short-tailed Shrew: There may be several short-tailed shrews per acre in
certain areas of favorable habitat in woodland or edge.

Little Short-tailed Shrew: Probably this shrew averages much less than one
per acre in the grassland portions of the Reservation where it occurs.

Mole: No basis is known for estimating numbers; from the abundance of
sign populations exceeding one per acre might be expected, at least for the
wooded parts of the area.

Red Bat: Probably there is one bat to several acres.

Cottontail: In September, 1955, Janes (MS) recorded 33 cottontails on a
21-acre area of woodland.

Fox Squirrel and Gray Squirrel: Packard (1956: 17) found a population
of 1.18 squirrels per acre in the better type of woodland; more than two-thirds
of this total consisted of fox squirrels.

Plains Harvest Mouse: In December, 1957, on a partly barren formerly
pastured 7.1-acre plot of the Rockefeller Tract, 65 plains harvest mice were
caught, indicating a population density of 9.2 per acre, almost the same as for
deer mice on the same plot.

Western Harvest Mouse: In late winter of 1951, on a hilltop grassland plot
of 1.6 acres, dominated by brome grass, an exceptionally high population with
a total of 80 adults was recorded but doubtless some were wanderers and
others lived mostiy outside the plot, having home ranges which merely over-
lapped it slighdy. Populations as high as 50 per acre are rarely attained, but
densities between one per acre and ten per acre are normal for most grassland
areas of the Reservation.

Deer Mouse: In December, 1957, 66 were caught on a partly barren for-
merly pastured 7.1-acre plot on the Rockefeller Tract, indicating a population
density of 9.3 per acre. On most areas where the deer mouse occurs, it is in
lower population densities.

80 University of Kansas Publs., Mus. Nat. Hist.

White-footed Mouse: In early spring of 1951, 16 adult white-footed mice
were caught in a woodland study area of 12)2 acres. Ordinarily the numbers
probably exceed one per acre in edge or woodland habitat.

Hispid Cotton Rat: In November, 1957, 28 cotton rats were caught in a
prairie plot of 3.1 acres on the Rockefeller Tract. The population of nine per
acre represented is regarded as moderately high, but at certain times and
places under favorable conditions it is much exceeded.

Eastern Woodrat: From a high population of several per acre in most
wooded areas in 1948, this rat decreased to a low point of perhaps one per
hundred acres in 1954.

Prairie Vole: Martin (1956: 378) found populations of from 25 to 140 per
acre in the grassland areas where his studies were made.

Pine Vole: The population of the entire Reservation is estimated as con-
sistently less than 100 adults, concentrated in a few highly locahzed colonies.

Norway Rat: For the entire Reservation the population of adults probably
has never exceeded ten.

House Mouse: Population densities of several to many per acre are at times
attained on limited areas, but ordinarily numbers are much lower.

Jumping Mouse: This species is poorly known on the area. It is doubtful
whether population density exceeds one per acre except for brief periods in
unusually favorable habitats.

Raccoon: Several family groups live on the area. Populations as high as
one to 50 acres probably have been attained.

Spotted Skunk: Winter populations somewhat exceeding one to 50 acres are
usual.

Striped Skunk: This species is slightly less numerous than the spotted skunk.
In winter there is somewhat less than one striped skunk to 100 acres.

Bob-tvhite: In the 1954 winter census 25 were recorded, but usually nimi-
bers are somewhat higher.

Mourning Dove: Probably at least 20 pairs breed on the Reservation.

Yellow-biUed Cuckoo: The breeding population probably exceeds 20 pairs.

Whip-poor-will: A breeding population of perhaps ten pairs is present in
summer.

Yellow-shafted Flicker: In the 1954 winter census seven were recorded,
but often the population is much higher.

Red-bellied Woodpecker: In the 1954 wanter census 21 were recorded, prob-
ably a typical number for that time of year.

Red-headed Woodpecker: The resident colony usually includes between 10
and 20 individuals.

Hairy Woodpecker: In the 1954 winter census 13 were recorded; probably
this population is fairly stable.

Downy Woodpecker: In the 1954 winter census, 42 were recorded.

Crested Flycatcher: The normal breeding population is estimated as some-
what more than ten pairs,

Phoebe: The breeding population of the Reservation is usually from three
to seven pairs.

Eastern Wood Pewee: Perhaps as many as 20 pairs nest on the area; no
definite figures are available.

Horned Lark: In 1957 six pairs nested on a 72-acre area of formerly cul-
tivated fields on the Rockefeller Tract.

Home Ranges of Vertebrates 81

Barn Swallow: Two pairs nested at the old bam on the Rockefeller Tract
in 1957. In earlier years many more had nested there.

Blue Jay: In the 1954 winter census 12 were recorded but usually more are

present.

Crow: In the 1954 winter census ten were recorded but this is probably
near the minimum figure. In winter there are, at times, hundreds on the area,
and in summer the breeding population may approach one to ten acres.

Black-capped Chickadee: In the 1954 winter census 214 were recorded, per-
haps a typical figure for that time of year.

Tufted Titmouse: In the 1954 winter census 40 were recorded.

Brown Creeper: In the 1954 winter census two were recorded but usually
a larger number winter on the Reservation.

Carolina Wren: In the 1954 winter census, nine were recorded, probably
a typical number for that time of year.

Brown Thrasher: Probably there are never more than five pairs present in
summer and sometimes none.

Eastern Bluebird: In winter, flocks of as many as 20 bluebirds use the Reser-
vation regularly. In summer there are probably never more than five pairs on

the area.

Blue-gratj Gnatcatcher: It seems probable that more than 20 pairs Uve on the
area, although no census figures are available.

Bell Vireo: With successional changes the number of breeding pairs has
gradually increased since 1952, to perhaps as many as ten pairs.

Red-eyed Vireo: A 1955 census figure of one pair per 2.1 acres on a
wooded north slope may apply for the better woodland habitats throughout
much of the Reservation.

White-eyed Vireo: Usually the breeding population has varied from one to
six pairs.

Kentucky Warbler: Probably not more than five pairs nest on the Reserva-
tion.

English Sparrow: A colony of more than 20 pairs bred in vicinity of the
buildings on the Rockefeller Tract in 1957, but none has successfully nested
on the Reservation.

Eastern Meadowlark: Approximately six pairs nested on the Rockefeller
Tract in 1957. To my knowledge no more than one pair annually has ever
nested on the Reservation.

Cowbird: The breeding population is estimated as the equivalent of at least
20 pairs. The majority of all passerine nests foimd on the area have been
parasitized by cowbirds.

Summer Tanager: At least six pairs breed on the Reservation annually.

Cardinal: In the 1954 winter census 40 were recorded, but usually the
winter population is somewhat higher. The breeding population also probably
exceeds this number nonnally.

Indigo Bunting: Perhaps as many as 30 breeding pairs are present in sum-
mer.

Dickcissel: In recent years the breeding population has amounted to ap-
proximately ten pairs.

Eastern Goldfinch: In the 1954 winter census nine were recorded.

Red-eyed Towhee: The breeding population is estimated at ten pairs or a
little less. The wintering population also normally approximates this number,
but none was recorded in the 1954 census.

82 University of Kansas Publs., Mus. Nat. Hist.

Junco: In the 1954 winter census 41 were recorded, but usually there are
more.

Tree Sparrow: In the 1954 winter census 17 were recorded. Numbers are
variable and sometimes greatly exceed this figure.

Field Sparrow: This is the commonest breeding bird; 58 pairs were re-
corded on a 300-acre area in 1950.

Harris Sparrow: The numbers of these sparrows wintering on the area are
highly variable depending on tlie food supply. In the winter of 1956-57 a
flock of more than 100 lived on the Rockefeller Tract and the adjacent part
of the Reservation, feeding largely on waste grain from a sorghum crop.

Song Sparrow: In the 1954 winter census 18 were recorded, but probably
the winter population is usually somewhat larger.

HABITATS

The habitats of the University of Kansas Natural History Reservation have
been described in some detail by Fitch (1952, 1954), Fitch and McGregor
(1956), Leonard and Goble (1952), and Packard (1956). These include not
only descriptions of the Reservation as a whole, but detailed descriptions of
small areas. The present brief discussion is intended merely to supplement
these earlier descriptions, and to show the chief successional changes that took
place from 1948 to 1957, while the study was in progress.

Fields in bottomland, cultivated until 1949, include Ditch Field and Coon
Field in the small valley of the west-central part of the Reservation, and Oat
Field, Square Field, Com Field, Woods Field and Weed Field in the southern
part. The pioneer vegetation of secondary succession in all these fields at first
consisted chiefly of sunflower (Helianthus annuus) and giant ragweed {Am-
brosia trifida), in a rank growth, with stems as much as two inches in diameter
and plants as high as ten feet. After several seasons the sunflower had almost
completely disappeared and the giant ragweed had become much stunted, and
was absent from parts of the field. Many other weeds, including thistle {Cir-
sium sp.), horseweed (Erigeron canadensis), goldenrod (Solidago sp.), goose-
foot {Chenopodium sp.), wild lettuce (Lactuca scariola), nightshade (Solanum
sp.), spurge (Euphorbia sp.), velvetleaf (Abutilon tJieophrasti) , smartweed
(Polygonum sp.), aster (Aster sp.), Joe-Pye weed (Eupatorium altissimum),
and false boneset (Kuhnia eupatoroides) dominated certain parts of the fields.
Annual grasses, especially foxtail, became increasingly prominent, and dom-
inated extensive patches. Saphngs, especially tliose of American elm ( Ulmus
americanus) , also became increasingly prominent until by 1957 the edges of the
fields bordering woodland had become dense thickets, with trees up to twelve
feet high.

Throughout the woodland, in general, the trend was toward larger size in the
second growth trees, and shading out of the understory layer including not only
saphngs of the chmax species, but species such as red haw (Crataegus mollis),
crab apple (Pyrus ioensis), redbud (Cercis canadensis), dogwood (Cornus
drummondii) and wild plum (Prunus amcricana). Later, in 1953 and 1954,
there was extensive mortality in trees of all sizes, particularly American elm
and black oak (Quercus vclutina), as a result of drought and disease. Large
gaps in the leaf canopy opened up permitting the establishment of various
weedy herbs, notably wild lettuce. The dead trees are in process of disinte-

Home Ranges of Vertebrates

83

gration, producing an abundance of rotting logs and hollow snags, such as has
not existed before since the area first came under control of white immigrants.

While most of the woodland areas were fenced and were protected from
livestock throughout the nineteen forties, the wooded hillsides of "Horse
Woods," "Skink Woods," and "Rat Woods," were connecting strips between
pastiures and were exposed to grazing and browsing until 1949. At that time
the three areas had little underbrush or herbaceous vegetation, and were rela-
tively open as compared with adjoining parts of the woodland over the fence.
By 1957 the reverse was true; the formerly browsed woodlands had dense
thickets of elms, hackberries, and other young trees, up to six feet high, while
in the adjoining areas the underbrush had grown taller and sparser.

Former pastures on the area, including a long flat hilltop from Angle Field
on the southwest to Gate Field on the northeast, Top Field, Quarry Field, Tree
Field and Point Field, also on hilltops, and Picnic Field, House Field, Dike
Field, Gully Field and Cow Field in the two small valleys, have changed even
more than the woodlands. Formerly these fields had only a few trees, which
were of medium to large size, and the awnless brome grass {Bromus inermis)
and bluegrass ( Poa pratensis ) which provided the chief forage were mixed with
many weedy forbs, notably nightshades, spurges, vervains (Verbena sp.),
milkweeds (Asclepias sp.), lespedeza (Lespedeza striata), germander (Teu-
crium canadense), and ironweed (Vernonia baldwini). In succeeding years
since grazing was discontinued, the brome has made up an ever-increasing
proportion of the herbaceous vegetation and all the weedy species have become

Fig. 3. Map of an area in the northwestern quarter of the Reservation,
where field work was most concentrated, showing ten-foot contour intervals,
distribution of woodland (tree pattern) and grassland, roads and buildings,
diversion ditch, hilltop outcrop of rock ledges (stippled), and other details.
The maps in Figs. 4 to 10 and 16 to 24 include parts of this same area.

84 UNi\'ERsn-i- OF Kansas Publs., Mus. Nat. Hist.

scarce. At the same time young trees have sprung up in profusion. These
saplings are unevenly distributed, as in some parts of the fields, especially near
the edges they form dense thickets, while in other areas, farther removed from
seed sources, they are relatively sparse.

An area in the northYv-est comer of the Reservation was long maintained
as bluestem prairie but by 1957, after appro.vimately 20 years of protection
from fire and mowing, encroachment of woody vegetation, including dogwood,
sumac (Rhus glabra), blackberry (Rubus argutus), and saphngs of various
trees, had crowded out so much of the grassland \-cgetation that on the hill
slope the area had assumed the aspect of a thicket. On the adjacent hilltop,
encroachment proceeded much more slowly, and was temporarily halted iii
April, 1956, when a fire spread from neighboring land and burned over this
small hilltop prairie patch. The fire killed most of tlie wocdy vegetation, in-
cluding hundreds of young bo.xelders (Acer negundo). In 1956 and 1957,' big
bluestem (Andropogon gerardi) made up the bulk of tlie vegetation in' this
prairie plot. Otlier kinds of prairie vegetation including many kinds of legumes,
and composites that were abundant on an adjacent plot to the north (on the
RockefeUer Tract) were scarce or absent on the prairie of the Reser%ation.

An upland area under cultivation in the nineteen thirties, and so heavily
eroded that most topsoil was lost, included Road Field and High Field which
were sown to seeds of native grasses in May, 1949. By 1957 the perennial
grasses were well estabhshed and dominated the area causing it to have super-
ficially, the aspect of a prairie, but only a few of tlie grasses and f orbs' typical
of praine were present on this area. Deep gullies cut by torrential rains when
the area was cultivated, have become largely revegetated but there are stiU
barren patches on the steep banks. The stand of perennial grasses is not con-
tinuous but there are intervening weedy patches, with goldenrod and aster
dominating Dry Field and Slope Field constituting the eastern part of this
upland field area were not sown to native grasses, and revegetation has pro-
ceeded slow y. Erosion continues, with vegetation sparse, and many small areas
of bare soil intervening. Goldenrods, aster, lespedeza and three-awn grass
(Artstida oligantJm) are the most prominent plants.

Aquatic and riparian habitats are not well represented on the Reservation.
The pond near the headquarters was variable in extent, depending on weather
trends, but never covered more than an acre. Originally constructed in 1936
this pond had been nearly filled with silt fifteen years later. It was deepened
along the side adjacent to the dam and an outlet pipe was installed in 1953-1954
Although water has been present continuously since then, the level has fluctu-
ated widely. Aquatic vegetation consists chiefly of pondweed (Potamogeton
sp_), naiad (Na,as guadalupensis) , wapato (Sagittaria latifolia), and cattail
(Typha sp.). The willow grove that has overgrown the silted portion of the
pond has a dense understory vegetation with various hydrophytic plants Some
of the wiUow trees have grown to a trunk diameter of six inches, and many
ot them have died and fallen, creating a tangle of dead trunks amid the thick
weedy vegetation.

The intermittent creeks that flow through the two small valleys have cut
deep and narrow gullies. Their bottoms are gravelly. Hydroseral vegetation
is scarce. At the upper ends of the small valleys the creeks emerge from steep
rocky ravines on the wooded hillsides. After heavy rains these rocky stream
beds are filled with torrents of muddy water, but ordinarily they are dry.

Home Ranges of Vertebrates 85

Vegetation adjacent to these channels differs Httle from that found elsewhere
in the woodland, except that sycamore trees tend to be concentrated there.

In the spring of 1957, the Rockefeller Tract, a quarter-section adjoining the
western half of the Reservation on the north (Fig. 2) was acquired by the
University of Kansas as an adjunct to the Reservation itself. Unlike the
Reservation, the Rockefeller Tract is not maintained rigidly as a natural area.
Rather, it is manipulated in various ways with the object of restoring and
maintaining a tall-grass prairie vegetation on the portions that are not wood-
land. Several species of vertebrates ordinarily absent or rare on the Reserva-
tion were common on the Rockefeller Tract and were studied there in 1957.
This 160-acre area had an eight-acre virgin prairie (mowed annually for hay)
in the southwest comer, and had blocks of woodland in the remaining three
corners, but most of it was under cultivation. An 18-acre block in the east-
central portion was overgrazed pasture, dominated by bluegrass but with
many weedy annuals. A small artificial pond in this pastinre had often been
dry, but held water throughout 1957. The northwestern and northeastern
parts were north slopes of oak-hickory woodland somewhat different from any
on the Reservation, with a high proportion of red oak ( Quercus rubra ) and with
linden (Tilia americana) and certain other species not found on the Reserva-
tion. The southwest comer consisted of a ravine and hilltop, with a few large
elms and other trees, and with thickets of underbrush, especially blackberry,
dogwood and young honey locusts. The cultivated parts of the Rockefeller
Tract had been used for growing of milo, oats, wheat, barley and alfalfa for
many years. Soil had become considerably impoverished through erosion and
long continued cultivation. On the edge of the cultivated land the level is one
to two feet lower than that on the adjoining prairie as a result of loss of
topsoil through sheet erosion.

In April, 1957, the pasture and formerly cultivated portions of the Rocke-
feller Tract were disked as preparation for a seedbed, and were sown to
seeds of native grasses, chiefly little bluestem (Andropogon scoparius), with
smaller proportions of big bluestem (^Andropogon gerardi), Indian grass
(Sorghastrum nutans), switch grass (Panicum virgatum), and with some forbs
characteristic of tall-grass prairie. By July, 1957, weedy vegetation, chiefly
sunflower, ragweed (Ambrosia artemisiifolia) , and dogbane (Apocynum
cannabinum), had sprung up, shading the slow-growing perennial grasses. To
reduce competition from these weedy species, the area was mowed. By late
summer, annual grasses, notably foxtail (Setaria sp. ), and crabgrass (Digitaria
sanguinalis) made up a major portion of the vegetation.

As a result of the more open terrain and different vegetation, the Rockefeller
Tract in 1957 provided favorable habitat for the sparrow hawk, kingbird, bam
swallow, horned lark, loggerhead shrike, lark sparrow, English sparrow, chip-
ping sparrow, and a few other kinds of vertebrates that likewise rarely or
never visited the Reservation. Dickcissels, meadowlarks, and redwings were
also common in the open areas. Field sparrows, indigo buntings, and car-
dinals were common along woodland edges. Birds of the woodland were not
known to differ from those of the Reservation. In the pasture of the Rocke-
feller Tract the Plains harvest mouse was common. Otherwise mammals of
this tract were not known to differ from those of the Reservation. Wood-
house's toad, common on the Rockefeller Tract, was the only species of cold-
blooded vertebrate well represented on this tract but not regularly present on
the Reservation.

86 Uni\'ersity of Kansas Publs,, Mus, Nat. Hist.

ACCOUNTS OF SPECIES

Fishes
Pimephales promelas Rafinesque
Fathead Minnow
Status. — Irregular resident and wanderer on area whenever water
supply is adequate.

Habitat. — Lakes, ponds, and slow flowing streams; on Reserva-
tion limited to small creek draining eastern part of area. At maxi-
mum occurs for approximately half a mile along course of this
creek, living chiefly in small pools having clay and gravel bottoms.
Creek channel is within gully 15 feet or more in depth.

Movements. — From 1948 through July 1952 these minnows, and
probably other kinds also, were abundant along the creek's course.
Subsequently, drying progressed in the late summer of 1952, until
no water remained along the creek's channel and all fish had
perished. Throughout the next four years the creek was dry almost
continually and no fish were present on the Reservation. On July
29, 1957, several small adults and many young of various sizes were
found to be present. Frequent heavy rains in May and June, cul-
minating in a downpour of more than six inches near the end of
June had restored the flow of water, creating favorable habitat
conditions and had made possible upstream migration to the area.

These small minnows tend to be nomadic, and heavy summer
rains stimulate them to migrate upstream into small tributaries and
spawn. In the small pools where specimens of minnows were col-
lected, it was noticed that each fish had certain niches or crevices
beneath submerged rocks in which it sought shelter when too
greatly disturbed.

Among many kinds of small fish which might conceivably reach
the Reservation by migrating upstream from the Kansas River to
the heads of the small tributaries, the fathead minnow is the only
one known to have been present in 1957. In future cycles of wet
weather perhaps other kinds of fish will reappear on the area. Nev-
ertheless, it is certain that in the past one hundred years, habitat
conditions for fishes in general have undergone progressive de-
terioration on the area. Before major portions of the drainage sys-
tems were subjected to cultivation, and to heavy grazing by live-
stock, run-off was much less rapid. The two small streams were
then much different in character, meandering, with many pools and

Home Ranges of Vertebrates 87

sloughs, through their partly wooded valleys. The molluscan fauna,
represented by numerous unfossilized shells in the deposited silt
of the creeks, included several pond-living or semi-aquatic species
which have either disappeared from the area altogether, or are
now confined to the one small pond (Fitch and Lokke, 1956: 444).
Severe erosion has occurred. Much topsoil has been washed away,
especially in upland fields, and the creeks have incised their chan-
nels to form gullies that are as much as 20 feet deep, in places. With
each heavy rain there is further gully-cutting, but the rate has been
reduced. Protection of most of the watershed from grazing and
cultivation probably has been instrumental in checking erosion.
The water supply has become uncertain and fluctuating, often
turbid and with occasional flash floods. The fathead minnow seems
better adapted to survive under such unfavorable conditions than
does any other species of fish.

Ictalurus melas (Rafinesque)

Black Bullhead

Status. — Occasional and temporary resident, at least formerly;
none present in recent years.

Habitat. — Ponds, lakes, dams and sloughs, or slow-flowing streams.

Movements. — In 1951 and 1952, schools of young were seen in
the small creek in the eastern part of the Reservation. In 1954 after
exceptionally heavy rains in June, a small group was in the creek
at the west boundary of the Reservation. All these were young less
than an inch long which had moved upstream in schools, after heavy
rains.

Lepomis cyanellus Rafinesque

Green Sunfish

Status. — Reaches area as an occasional wanderer, when water
supply is suflBcient.

Habitat. — Streams with slow moving water, or ponds and lakes.

Movements. — In late June, 1954, several days after heavy rains,
three were collected in a pool at the west edge of the Reservation.
From the behavior of these sunfish it appeared that they were pre-
paring to breed. The heavy rains, following prolonged drought,
had seemed to initiate a breeding cycle, and had caused the fish to
run far upstream into this small tributary.

88 Uni\'ersity of Kansas Publs., Mus. Nat. Hist.

Amphibians
Ambystoma tigrinum Green

Tiger Salamander

Status. — Occasional wanderer; only Uvo found in nine years of
field work.

Habitat. — Both individuals recorded were in funnel traps in wood-
land along hilltop rock ledges, when the surface of the soil was still
damp after rain. One was caught on October 24, 1951, the other on
October 7, 1955.

Movements. — There is no evidence that the tiger salamander
breeds on the Reservation. Both individuals recorded were im-
mature and of approximately the same size, 140 mm. total length.
They probably had travelled considerable distances after meta-
morphosis. The nearest known breeding place was a slough in
the Kansas River Valley, approximately two miles farther south.

Scaphiopus bombifrons Cope
Plains Spadefoot

Status. — Few records have been obtained from the Reservation,
and it is unlikely that this secretive toad is regularly present even
in small numbers. Its voice has never been heard in the choruses
of breeding salientians at the pond.

Habitat. — Breeding choruses of spadefoot toads have been heard
at several places within a two-mile radius south and west of the
Reservation. From the location of these choruses it seems that
the spadefoots occur chiefly in the Kansas River flood plain and the
fairly level land near its margins, although one small pond, on the
tip of a spur of hilltop projecting out into the valley, was used.
Burrows of small mammals are probably an essential feature of the
habitat. Pocket gophers (Geomys bursarius) were abundant near
the places where spadefoot choruses were heard.

Movements. — None was recorded on the Reservation until May
8, 1955, when an adult female was caught in a funnel trap in a ditch
bottom near the headquarters approximately 3000 feet from the
nearest pond where choruses had been heard, and a mile from places
where other choruses had been heard to the south and west. On
July 13, 1956, after a heavy rain, two recently metamorphosed
young were found at points 1900 feet and 3000 feet from the hilltop
pond near the south boundary of the Reservation, where a chorus

Home Ranges of Vertebrates 89

had been heard some nine weeks earher. In the summer of 1957,
ten spadefoots, including both adults and young, were caught in
funnel traps in the western half of the Reservation.

Bufo terrestris Bonaterre
Common American Toad

Status. — Abundant resident.

Habitat. — Rocky situations in open woodland, or near woodland
edge, perhaps provide optimum habitat. The shelter provided by
flat rocks, with damp, loose soil underneath, is an important factor.
This toad is not narrowly Hmited in its habitat requirements, and
probably uses the entire area, but it is seldom found in open fields
with high dense vegetation. It breeds in the pond and, when suffi-
cient water is available, in the two small intermittent creeks, in
marshy situations where the current is normally sluggish.

Movements. — In the breeding pond each trilling male shifts posi-
tion frequently with no attachment to a specific location. Usually
not more than a dozen males were calHng at any one time, and they
tended to be concentrated within an area perhaps 30 or 40 feet in
diameter, but with a few well separated from the main aggregation.
However, from night to night the location of the main aggregation
shifted. An individual was often seen to swim for a distance of sev-
eral yards to take up a new position, and occasionally a marked in-
dividual recaptured after a short interval was found to have crossed
the pond from one shoreline to another. Males that were members
of breeding choruses were recaptured after intervals of 53 days, 31
days, 24 days, 23 days and several shorter periods. It is uncertain
whether these males had left and returned in the intervals.

Occasionally toads were found in their daytime retreats, in hol-
lows beneath flat rocks excavated partly by their own efiPorts. Tenure
of such shelters was found to be brief. For six immature toads
whose shelters were checked from time to time, known occupancy
averaged only 9.5 days. The longest known tenure was one of 26
days. Recorded shifts (in feet) from one shelter to another were:
150, 100, 65, 50, 30, 25. All the toads involved in these records were
immature individuals several weeks or several months old.

Size of home range was suggested by successive captures of seven
individuals all living in the headquarters area. The total of twelve
movements averaged 47 feet, indicating a home range of approxi-
mately .16 acres. This figure is almost certainly too low for a typical

90 Unin-ersity of Kansas Publs., Mus. Nat. Hist.

home range because the resident toads were most likely to be re-
captured when tliey were moving over the sidewalks or lawn ad-
jacent to the buildings, and were least likely to be noticed when
they moved farther away. The longest movement recorded was one
of 150 feet, in tvvo days.

The newly metamorphosed young, like those of other salientians,
disperse rapidly and wander widely in wet weather. On May 30,
1955, swarms of young were noticed at the edge of the water, many
still having tail stubs. Two days later the young were no longer
confined to the water's edge but were abundant on damp ground
near the pond. One had travelled 400 feet from water, but most
were still much nearer the pond. On June 8 one was found on a
wooded hilltop 700 feet from the pond and at an elevation 100
feet higher, and by the following week the young had dispersed to
every habitable situation.

Travel is undertaken at times when the humidity is high and the
soil is damp. The young toads soon become famihar with their
surroundings and each settles within a small area. Marked young
have been recaptured, after periods of days or weeks or even months,
near the original location. One marked on August 9, 1951, when it
was 39/2 mm. in length, was recaptured 150 feet away on April 30,
1952.

Breeding choruses were heard at several farm ponds within a
half-mile radius of the single breeding pond on the Reservation.
Ponds and intermittent creeks are well distributed over the general
area; consequently individuals need not travel more than half a mile
to spawn. One male caught and marked as a well grown young
(51 mm.) on a wooded slope on May 7, 1953, was recaptured in a
breeding chorus at the pond some 1900 feet away, a little more than
a year later. A female marked when she was about half-grown was
recaptured 150 feet away, in the pond, as a breeding adult nearly
22 months later.

Bufo woodhousei Girard

Woodhouse's Toad

Status. — Scarce and irregular resident.

Habitat. — This toad is much more numerous on adjoining closely
grazed pasture and cultivated fields than it is on the Reservation,
where it has been foimd only in grossly disturbed situations, along
the road, on or near the lawn, and at the pond. It would seem that
the woodland, and the dense herbaceous vegetation present in
grassland, render conditions unfavorable for it over most of the
Reservation.

Home Ranges of Vertebrates 91

Movements. — Most records of this species on the area were ob-
tained in 1955, 1956, and 1957. On July 4, 1956, for most of the
day, two were calling at the pond where none had been heard
before, although several other ponds within a mile of the Reserva-
tion were used regularly for breeding. In 1955 an adult female
was seen on September 19, 21, and 23, and on October 10, within
a 50-foot radius about the house. Another large adult female was
recorded near the house on September 24, 1955, and again on
August 21, 1956. Six young were found near the house, in late
August and September 1955, and in July, August and September,
1956.

Acris gryllus Le Conte

Cricket Frog

Status. — Resident, and at times perhaps the most abundant of
aU species of vertebrate animals on the Reservation, although its
numbers vary greatly in response to weather conditions, and time
of year.

Habitat. — Water, either stagnant or running, is the chief require-
ment.

The cricket frog is present about the pond and along the two
small creeks and their tributary gullies, wherever moisture is nor-
mally to be found. Shallow water, either stagnant or with sluggish
current, along muddy banks supporting thick grass, provides the
most favorable type of habitat. The proportion of the area habit-
able at any one time depends upon the humidity and moisture sup-
ply; at times when moisture conditions are favorable there is dis-
persal throughout the woodlands and parts of the open areas. In
time of drought the population dwindles, and becomes restricted
to the remaining limited area with a water supply.

Movements. — Cricket frogs tend to stay in damp places but may
wander extensively along creek beds and pond edges in dry weather,
and almost anywhere in wet weather. Available evidence, from
direct observation on individuals, and from recapture of marked
frogs, seems to show that there is but little tendency to remain
within a familiar area.

In the spring of 1955 I made effort to mark those in a small area
— a strip about 70 feet long along the pond edge at the northwest
comer. At each successive visit most of the frogs taken were new,
unmarked individuals, indicating a high rate of population turn-
over. The numbers of new and recaptured individuals taken on
various dates were as follows:

92 Uni\"ersit\' of Kansas Publs., Mus. Nat. Hist.

April 22 2 males and 5 females marked.

June 12 8 males and 5 females marked; no recaptures from April 22.
June 13 5 males and 5 females caught, all new e.xcept 1 male from June 12.
June 16 2 males and 6 females caught; 1 male and 2 females from June 12.
June 20 3 males and 3 females caught; 1 male and 2 females from June 12,

and 1 male from June 13.
June 21 1 male and 1 female caught, the female from June 20.
July 2 3 males and 8 females caught, including 1 male and 1 female from

June 12 and 2 females from June 20.

The pond's circumference amounted to some 1100 feet and most
of the frogs disappearing from the strip under observation probably
remained somewhere along its borders. However, one adult female
marked on April 11 was recaptured on May 29, 500 feet west of the
pond on damp soil in a ditch bottom.

On many occasions, in rain or mist, cricket frogs have been seen
travelling far from permanent water, in tall grass of fields, in under-
growth of wooded slopes, or crossing roads. Each heavy rain
seems to trigger a mass migration from the normal habitat of marsh,
streamside, or pond. The migrating frogs seem to wander at ran-
dom until, presumably, they perish from lack of moisture as dry
weather again sets in. Only a small percentage would be likely to
find suitable conditions permitting them to survive. In the summer
of 1951 the frequent heavy rains permitted extensive wandering. At
an old quarry 600 feet from the pond and appro.ximately 100 feet
higher, a depression in the limestone, several inches deep and three
feet in diameter nearly always contained water and several cricket
frogs usually could be found in the vicinity.

The frequent dispersal and decimation of the population relieves
overcrowding in the streamside or marshland habitat. Also, it en-
sures that any area of suitable habitat will be found and occupied.
The frogs are able to travel far and rapidly, as they easily cover
two or three feet at a leap. Hence, they are able to seek out remote
and isolated areas of habitat at the times when they are wandering.

The larval and postlarval development is rapid, and the crop of
young cricket frogs produced one year seemingly attains sexual ma-
turity in time to participate in the next breeding season. The repro-
ductive potential is therefore high, with normally an adequate sur-
plus to repopulate all available habitats.

Home Ranges of Vertebrates 93

Hyla versicolor Le Conte
Gray Treefrog

Status. — Common resident.

Habitat. — For breeding evidently requires semi-permanent stag-
nant pools with mud bottoms and weedy vegetation; some occur
in this type of situation throughout summer, but most adults dis-
perse from breeding pool to woodland early in summer. Occurs
throughout woodland preferring situations where trees are large
and where there are low shrubs; least favorable woodland is open
type on dry south slopes, consisting mainly of honey locust and
osage orange. Occasionally found in woodland edge situations.

Movements. — Of 113 treefrogs marked in the course of eight
seasons of field work, only five were ever recaptured. Most of the
frogs marked were adults in breeding aggregations at the pond. One
male marked at the edge of the pond on May 2, 1955, was recap-
tured 30 feet away on June 1. Each of two others recaptured after
shorter intervals had moved approximately 30 feet.

An adult female was found in a niche in the vertical rock face at
the old rock quarry on July 14, 1950. On August 18 she was re-
captured 90 feet farther along the rock exposure. On August 26
she was recaptured a second time at an intermediate location.
Another adult female found on the outside wall of the house on
October 5, 1952, was recaptured there on October 18. The records
of these two frogs suggest that an individual tends to settle in a
limited area, for periods of weeks, at least. The adults are not
commonly seen except at the breeding ponds, and information con-
cerning their movements is difficult to obtain.

Newly metamorphosed treefrogs, unhke chorus frogs, toads, and
narrow-mouthed toads, do not disperse rapidly from the breeding
pond. By the time the treefrogs have metamorphosed, in middle
or late summer, annual vegetation such as smartweed, giant rag-
weed and cattail, has grown tall and rank, providing shade and
maintaining high humidity near the wet ground. The young tree-
frogs live on these broadleaved plants. In several different years
they remained abundant near the pond up until the time when cold
weather arrived.

94 University of Kansas Publs., Mus. Nat. Hist.

Pseudacris nigrita Le Conte
Chorus Frog

Status. — Abundant resident, but with great variation in population
level from year to year; wide fluctuations in numbers are affected
by varying success of breeding season.

Habitat. — Grassland seems to be preferred, but some have been
noted in woodland or at the edge of woodland. This frog is so
secretive in its habits that it is rarely seen, and its abundance
would scarcely be suspected, were it not for the attention attracted
by breeding choruses. Impressions regarding habitat preferences
have been gained chiefly by observations on the dispersal of the
newly metamorphosed young, and the situations where calling
adults have been heard, away from water, just before onset of the
breeding season. Such bits of information indicate a wide choice
of habitats. The frogs seem to be largely subterranean in their
habits. Most of those found were under rocks. Probably burrows
of the prairie vole, and other small mammals, are important in the
chorus frog's ecology.

Movements. — Chorus frogs are active above ground mainly while
rain is falUng, or afterward while soil and vegetation are still wet.
The records are fairly uniformly distributed throughout the year
except for December and January, Excluding those frogs that were
found in or beside water, 38 of the 44 records for adults were in
grassland or open weedy situations, and only six were in woodland.
Similarly, 29 of the 36 records for young were in grassland and
other open situations. Chorus frogs found in woodland were gen-
erally in the more open parts. Most of the frogs found at water
were either breeding adults or newly metamorphosed young. A
seeming exception was an adult male found at the edge of the pond
with swarms of cricket frogs on October 20, 1955.

Chorus frogs found at times other than in rainy weather, or under
extremely humid atmospheric conditions were in well protected
situations, typically under flat rocks. Whether the frogs carry on
all their activities underground for long periods or emerge regu-
larly to forage by night, is uncertain. On the average, they breed
well before other local salientians in spring, sometimes even in
February, but rain is a necessity for migration to the breeding
ponds. In 1956, when no spring rains occurred, breeding was much
delayed at the pond on the Reservation, although the supply of
water was adequate. Throughout April and early May, 1956,
whenever there were warm days with high humidity, one was heard

PLATE 1

Fig. 1. Eastern end oi "House Field" in the headquarters area of the
Reservation, August 4, 1951. The foreground vegetation consists of awn-
less brome grass, interspersed with \'arious weeds, notably vervains, iron-
weed, milkweeds, and ragweeds. The prairie vole, cotton rat, western
harvest mouse, glass lizard, and yellow-bellied racer were abundant in the
area shown. When this picture was taken, herbaceous vegetation was
especially luxuriant as a result of abundant summer rain. The area shown
is included in all the maps, figs. 2 to 24.

Fig. 2. Another view of the eastern end of House Field with dam and
willow grove in background, August 4, 1951. Brush and young trees en-
croaching into the field have created a typical "edge" habitat favorable
to the cottontail, pine vole, cardinal, indigo bunting, and'field. sparrow.

PLATE 2

Fic. 1. Site of old liiiK'stonc qiiariy tjii .soutliu.ucl piojceliug spur
ot liilltop, OctoljtT 13, 1949. Linu-stoiif strata are proiniiiently exposed
with cracks and fissures, pro\iding shelter for many kinds of small
vertebrates. Lc\el n^ky area in foreground has a sparse \egetation of
lespedeza and ragweed. The open, rock\- area was espeeialK- fa\-orable
habitat for various reptiles including the box turtl(>. collared lizard,
racerunner. Great Plains skink, slender-fiat-headcd snake, red milk
snake, prairie king snake. The narrow-mouthed toad, striped skunk,
woodrat, white-footed mouse, bob-white, and broad-winged hawk also

frequented this place.

V

'\

'^•S.

Fk;. 2. Diversion ditch at edge of woodland near
(juarters, Jiuic 21, 1919. The bare soil of tlic banks
provided ia\()ral)lc habitat for the six-lined racerunner.
travelway for coyotes, foxes, opossums, various kinds of snakes, and in wet
weather, amphibians including the spadefoot toad, American toad, cricket
Irog, bulllrog, leopard frog, and narrow-mouthed toad

l^eservation luad-

and rock> bottom

This ditch was a

PLATE 3

Fig. 1. Woodland of a rocky upper slope, dominated by chestnut oak,
July 22, 1950. Many kinds of vertebrates prefer tliis habitat type. Some
species that attained their maximum numbers here were the narrow-
mouthed toad, American toad, worm snake, copperhead, short-tailed shrew,
mole, gray squirrel, and white-footed mouse.

Fig. 2. Hilltop rock ledge in woodland dominated by American elm,
November 7, 1951. The ledge provided shelter for many kinds of animals,
notably the opossum, short-tailed shrew, woodrat, and cottontail. In
autumn various species of snakes including the timber rattlesnake, copper-
head, yellow-bellied racer, black rat snake, king snake, garter snake and
water snake repair to such places to find shelter for hibernation.

PLATE 4

^&fi|k^^.

* *' ».Ai

Fig. 1. Eroded Iiilltop field at end of 6th growing season after sowing na-
tive grass seeds in April, 1949. Side-oats grama to left of gully, and
switch grass to right. Deep gulK ehoked with vegetation and erosion al-
most stopped. U. S. Soil Conserx ation Str\ ice plioto, November 9, 1954.

r??i

ml$<^^'

Fic;. 2. Another |);iit ol miiiic field ( shown in l''ig. 1 ) without iiati\e
grasses, eroded liom c iilti\ation in ihc 19o()s. U. S. Soil (Conservation .Serv-
ice iihoto. \()\cml)cr 9, 1954.

PLATE 5

'^rW^^nnmmm^

l>.

<»4.iT|l «■ ^^

Fig. 1. Pond 200 yards east-northeast of Reservation headciuarters on No-
vember 1, 1951. Cattails, and other hydrophytic vegetation surround the
open water. American toads, narrow-mouthed toads, treefrogs, chorus
frogs, cricket frogs, leopard frogs and bullfrogs bred in abundance here.
Water snakes and snapping turtles preyed upon them. Redwinged black-
birds nested in the marsh.

Fig. 2. Upland prairie in norlhvvesteru part of Reservation, October 13,
1951. Foreground was part of a farm (later purchased as the Rockefeller
Tract) and was subjected to annual mowing and occasional burning. A
rich flora of characteristic prairie plants remains. In background, on Reser-
vation, is prairie area not subjected to burning or mowing for many years,
dominated by big bluestem, but having fewer associated species than the
mowed area. Trees in background are along rock ledge at top of brushy

slope. Photo by Robert Rose.

PLATE 6

Interniiltciit crci'k in (!(ip ;iiill\ in southeastern part ol Reservation,
November 7, 19U). 'I'lie acinalie laiina liere was meaijer, but ialhead
minnows were often present in the ereek. Ameriean toads, erieket frogs,
ehoriis frogs, and leopard frogs bred in the pools. Co\otes, opossums,
raccoons, and skiniks rcgnlarK- hunted along this gully, and used it as a

travelway.

Home Ranges of Vertebrates 95

calling, in a three-inch iron culvert beneath the sidewalk at the
Reservation headquarters. It was not known to have emerged.

Rate of dispersal of the newly metamorphosed young was ob-
served in 1951. On the night of June 6 there was a heavy rain and
on the following morning there were swarms of young at the quarry,
having migrated 200 yards overnight up a wooded slope from the
pond at 100 feet lower elevation. Similar rapid dispersal of young
has been noted on other occasions. Otherwise little is known con-
cerning movements from day to day and over longer periods, be-
cause none of those marked was ever recaptured.

Gastrophryne olivacea (Hallowell)
Great Plains Narrow-mouthed Toad

Status. — Common resident.

Habitat. — Perhaps this species occurs throughout the entire area,
but it prefers rocky situations in grassland or open woods, and is
abundant along hilltop rock ledges, especially where there is open
woodland. Less frequently these toads have been seen or trapped
in open fields of the valleys or hilltops. They are secretive in habits
and nocturnal in movements and finding them is difficult where
there are no flat rocks to turn. In such situations they probably
depend upon burrows of other animals for shelter. Breeding occurs
at the pond and at a rain pool a few hundred feet from the south
boundary of the Reservation. Several times, in summer, after
periods of unusually heavy precipitation, when water had collected
in small weedy depressions in hilltop fields, these toads were heard
calling in them.

Movements. — Freiburg (1951: 384) published findings concern-
ing movements of marked narrow-mouthed toads on the Reserva-
tion. More recently Fitch ( 1955 ) analyzed the findings concerning
movements that have resulted from the capture and marking of
1215 individuals over a six-year period. Only eight percent of the
toads marked were ever recaptured. The mean distance for move-
ment for all individuals recaptured was 72 feet. The secretive
habits of the toads render them difficult to study. Those that
moved farthest from the places where they were marked were
least likely to be recaptured, since most of the work was concen-
trated on small study areas.

Certain individuals were found in the same niche under a flat
rock over periods of days or even weeks, but these individuals were
exceptional. Of the relatively few that were recaptured, most were

2—3117

96 Unutrsitv of Kansas Publs., Mus. Nat. Hist.

not found at the same place on successive occasions. An individual
seems to have no definite permanent home base, but the habits are
sedentary. The toads ordinarily do not venture away from their
protected shelters except when the soil and vegetation are wet,
during or soon after heavy rains.

Of the 59 individuals recaptured after one or more hibernations,
47 were adults, many of which had made one or more round-trip
migrations to the breeding ponds. Most of these toads were re-
captured in the general vicinity of earlier captures, indicating a
homing tendency. Of the toads that were recaptured at greatest
distances ( hundreds of feet ) from the point of release, several were
at the breeding pond or en route to it in heavy rains. One adult
female was caught in the same funnel trap in both 1954 and 1955
while making the trip between the home range and the breeding
pond, suggesting a tendency to follow a definite route on successive
trips to water.

The concept of home range does not apply well to animals having
habits like those of the narrow-mouthed toad, which does not range
regularly over a definite area, but occupies in succession a series of
subterranean niches in the same general vicinity. When weather
conditions are favorable the toad migrates to a nearby niche and
does not necessarily ever return to the one it has left. The niche
occupied provides protection from enemies, and from extremes of
the hostile outside environment; also it provides a regular food
supply (chiefly ants of the genus Cremafogaster).

Narrow-mouthed toads that were recaptured even after intervals
of weeks, months, or years, were most often less than 50 feet from
the site of original capture. For greater distances, up to 400 feet,
there was gradual decrease in the numbers of individuals. Several
individuals recaptured at distances in tlie neighborhood of 400 feet
from an original location were later recorded to have moved back
near the original site. Therefore it seems that some "home ranges"
may have a maximum diameter as great as 400 feet although cer-
tainly the usual diameter is much less.

Rana catesbciana Shaw
Bullfrog

i^tatus. — Resident, with a large population at the pond.

Ilahitat. — The pond, normally having a surface area of about
one-half acre, and adjoining marshy areas of somewhat greater ex-
tent, provide the habitat to which this frog is normally limited on
the Reservation. Although this small area is normally such favor-

Home Ranges of Vertebrates 97

able habitat that it supports hundreds of adult and partly grown
bullfrogs, the population often has been decimated by drying in
late summer.

Movements. — When drying is in progress, dispersal of frogs
occurs mainly at night, or at other times when humidity is high.
At such times the frogs are frequently seen in high grass, often along
the creek bed and sometimes hundreds of feet from it, in the
bottomland valleys. They have seldom been found on wooded hill-
sides. Their dispersal is less random than that of leopard frogs,
and they tend to keep to low ground where the chances of finding
water are better.

Bullfrogs were abundant at the pond in 1952, but in mid-July
the pond dried and all were eliminated either by death from desic-
cation or migration down the creek bed % mile or more to places
where water remained. On the afternoon of July 10 when humidity
was high, a lai-ge adult was found in high grass more than 100
yards from the pond. It did not attempt to escape when disturbed,
but stood high oflF the ground and inflated its body to assume a
threatening posture.

The pond held water in late spring and early summer of 19S3,
On May 18 a bullfrog was calling there. It must have travelled at
least half a mile — farther if it followed the creek bed. Others
reached the pond in the next few weeks but the population re-
mained low and was again eliminated when the pond dried in late
summer. In 1954 the pond did not dry, and by early 1955 several
adults had become established.

In late May and early June, 1952, 24 bullfrogs were marked with
colored tags. A total of 14 recaptures and sight records were re-
corded; in five instances no measurable movement had occurred;
two had moved approximately 10 feet, 3 approximately 20 feet, 2
approximately 30 feet, and one approximately 40 feet. Each indi-
vidual kept to its own small area within the pond. However, over
periods of weeks there was a reshuffling of individuals. Home
ranges, if they existed at all, were ephemeral.

Rana pipiens Schreber
Leopard Frog

Status. — Resident, with high population at pond, and occurring
throughout entire area, population density and extent of area occu-
pied varying greatly according to season and weather conditions.

Habitat. — Largely dependent on stagnant or sluggish water, at
pond and to lesser extent, along two small creeks.

98 UNRTRSiTi' OF Kansas Publs., Mus. Nat. Hist.

In moisture laden vegetation, when humidity is high, after rain
or heavy dew, the population may disperse to every habitat of the
Reservation. The frogs, especially the partly grown juveniles, are
commonly found in woodland throughout the summer.

Movements. — Leopard frogs commence breeding in early spring,
and by early summer the tadpoles are completing their metamor-
phosis. In dry weather large numbers accumulate in the moist
areas along margins of creeks and ponds, but during and just after
heavy rains there is a mass dispersal, with relatively few remaining
behind. The breeding season may continue from early March
until mid-summer or even later, and successive waves of migrants
emanate from the breeding ponds throughout the summer. Pre-
sumably most of them perish without finding a suitable water sup-
ply. However, if drought conditions are not severe, they may sur-
vive for days or weeks, staying in protected shelters under rocks,
logs, or boards, when humidity is low, and moving about at night.
Adults as well as young participate in the mass migrations from
the breeding ponds. Travel is fairly rapid on the overland mi-
grations. On rainy nights the frogs have been seen in the glare
of automobile headlights hopping across roads. Within a few
hours after heavy rains, frogs have been caught in funnel traps on
slopes and hilltops hundreds of yards from tlie pond, where prob-
ably the trip was begun.

No evidence of home range nor tendency to keep within a fa-
miliar area has been found in the leopard frog. Of several dozen
marked and released at different times, none has been recaptured.

Reptiles

Chelydra serpentina Linnaeus

Snapping Turtle

Status. — Moderately common as a resident or transient.

Habitat. — Confined to pond and two small streams on area, so far
as has been observed.

Movements. — Over a seven year period 18 snapping turtles have
been captured and marked on the Reservation but none has ever
been recaptured. Twelve of those marked were hatchlings; the re-
maining six were large adults. Several other large ones were seen
that were not caught. The evidence thus far obtained suggests
that these turtles wander more or less indefinitely following up and
down the small intermittent creeks according to the water supply.

Home Ranges of Vertebrates 99

Terrapene Carolina Linnaeus
Eastern Box Turtle

Status. — This species is tentatively classed as an occasional
transient or resident, on the basis of one record. The Reservation
is slightly outside the known range, vi^hich extends from the east
into nearby Wyandotte and Franklin counties, Kansas. However,
there is an old (and somewhat doubtful) record for Manhattan,
Riley County (Smith, 1956: 137).

Habitat. — Deciduous forest and edge.

Movements. — The only record is of a large adult found by Donna
Fitz Roy on May 29, 1958, 250 feet south-southwest of the Reser-
vation headquarters. It is almost certain that this individual had
recently come to the area; otherwise it would have been seen as
frequently as the other box turtles having ranges in this vicinity.

Terrapene omata Agassiz
Western Box Turtle

Status. — Resident on area, but in sparse population as compared
with numbers in more favorable habitat elsewhere.

Habitat. — Probably this terrestrial turtle occurs throughout the
area but it has not been found in the fallow fields that have rank
growths of weeds so tall as giant ragweed and sunflower. Grazed
pastureland provides the most favorable habitat, woodland next,
open fields with undisturbed prairie vegetation or with brome grass
and weed mixture are less preferred, and the rank weed habitat of
recently fallowed fields seems to be least favorable.

Movements. — Because box turtles were scarce on the Reservation,
the information obtained concerning their movements was meager.
However, in the closely related Terrapene c. Carolina, movements
have been studied more thoroughly than in any other kind of animal
(Stickel, 1950). Probably T. omata is similar to T. Carolina in the
extent and pattern of its movements. Stickel found that in T. Caro-
lina the average range of adult males was 330 feet and that of adult
females, 370 feet; the difference was not statistically significant.
Home ranges are maintained from year to year, but overlap widely
with no indication of territoriality. Within the area of its home
range a turtle seems to wander in an aimless manner, making nu-
merous turns, criss-crossings and detours. However, there seems to
be a tendency to return to certain places and to travel repeatedly
over certain relatively short preferred routes; "One turtle walked

100

University of Kansas Publs., Mus. Nat. Hist.

over a single short stretch of path seven times in eight days, travel-
hng over diverse areas between times." (Stickel, op. cit.: 367).

Fourteen box turtles marked on the Reservation were recaptured
from one to nine times. Seven were adult females, six were adult
males and one was a juvenile of undetermined sex. The time span
involved was within a single season for three; two seasons for four;
three seasons for two; and four, six, and seven seasons for the other
three. In 30 instances distances between successive sites of capture
averaged 325 feet. One exceptionally long movement of 1830 feet
was made by a female in a 53-day interval. This movement prob-
ably represented wandering well outside the usual range, and tliere-

FEET

Fig. 4. Range of an adult female box turtle recorded 17 times from
June 17, 1954, to July 31, 1957.

Fig. 5. Range of an adult female box turtle recorded 11 times from July
11, 1949, to May 25, 1957. The area of activity for this individual probably
is much larger than for the one shown in Fig. 4; in the interval of nearly
eight years the one concerned in Fig. 5 altered her home range extensively.

Home Ranges of Vertebrates 101

fore is best eliminated from the figures used to estimate size of home
range. For the remaining 29 movements, the average was 274, and
there was but httle difference between the sexes ( average for seven
males, 282 feet; for 22 females, 270 feet).

Assuming random movement over the home range (although
Stickel's findings suggest that the movemnts are not entirely ran-
dom), and assuming that captures recorded were within tlie regular
home range of the individual involved (except for that of the in-
dividual that had moved 1830 feet), the radius of the home range,
in the neighborhood of 274 feet, indicates that an area of approxi-
mately 5.4 acres is covered.

One turtle was brought to the Reservation headquarters and re-
leased approximately half a mile from its original location. Nearly
a year later it was found only 210 feet from the point of release.
This single observation suggests that homing tendencies are either
totally lacking or weak in the box turtle.

On several occasions box turtles were equipped with trailers, a
spool of thread rotating on a spindle and unwinding as the turtle
travelled. Such turtles usually travelled several hundred feet in the
course of a day. Several individuals equipped with such trailers
were kept under observation for periods of a week or more by John
M. Legler, who also has made much more extensive studies of the
box turtle's natural history and ecology in grazed pasture habitat
west of Lawrence.

Chrysemys picta Schneider
Painted Turtle

Status. — Recorded just once in fall of 1951, at pond.

Habitat. — The one individual seen was basking in the sunshine
on a floating board in the pond. The species is known to occur in
lakes, large ponds, and sluggish streams where the water supply is
sufiBciently extensive and permanent to provide shelter and to
support an aquatic fauna upon which the turtle depends for food.

Movements. — No definite records of movements have been ob-
tained, but the abrupt appearance of the single adult individual at
the pond, perhaps IM miles from the nearest suitable habitat, indi-
cates that it must have made an overland journey of this distance;
or it might have followed up the creek from a larger stream, which
would have involved an even longer trip.

102 University of Kansas Publs., Mus. Nat. Hist.

Crotaphytus coUaris Say
Collared Lizard

Status. — Permanently resident, 1949 to 1956, but with few indi-
viduals, and limited to small area, at quarry.

Habitat. — Requires open, sunny situations, with loose rocks; habi-
tat represented on Reservation only at old rock quarry.

Movements. — Suitable habitat on the Reservation over the period
of study was limited to an area at an old rock quarry approximately
400 feet long and 100 feet wide. The colony under observation was
small; 56 individuals, in all, were recorded over a seven-year period,
but these represented several successive generations. Although
there was territorial pressure, the colony showed no tendency to
expand into adjacent habitats. They filled the quarry area to the
extent of its carrying capacity, and so far as known, those that were
eliminated were taken by predators without having left the area.
At times when the population was crowded, as a result of suc-
cessful reproduction, incidence of mortality was notably increased
because of intraspecific friction. Recently hatched young, partly
grown young that are attaining sexual maturity, and adult males
that are dominated by others are especially susceptible to elimina-
tion.

Hatchling collared lizards, escaping from the nest burrow in
broods of two to a dozen or more, tend to disperse immediately
and wander for at least a day or two, to distances up to a hundred
feet, or perhaps more; consequently they are not concentrated
where they would compete with each other or where several
at once would be captured by a predator. Hatchlings recently
emerged have been noted to run in a random and aimless manner
when attempting to escape. Others, still less than a week old, have
been noted at each alarm to run to a regular place of shelter, a bur-
row or crevice providing security. Such young obviously had al-
ready become familiar with their immediate surroundings, and in
their day to day activities they tend to stay in home ranges only a
few yards across, consisting of one or several look-out rocks for
perching and basking, and as escape shelter. Over periods of days
or weeks, nevertheless, the juvenile may gradually shift its activi-
ties to a place many rods removed from its original home range.
Also, the size of its home range is gradually increased.

From the time they are about a quarter-grown in bulk (75 to 80
mm. in snout-vent length), males are territorial. On September 30,

Home Ranges of Vertebrates 103

1953, when no adult males were present, the eight young males all
approximately seven weeks old, were almost uniformly spaced
along the 400-foot stretch of ledge at the quarry. Pursuit and
fighting resulted immediately when any one of the young males was
chased into the area dominated by another.

Ordinarily no more than two adult males were present at the
quarry, and they divided the 400-foot stretch of ledge between them.
A break in the outcrop near its middle separated the two territories,
each of which was dominated by a succession of different males.

Territories tended to be linearly arranged along the 400-foot
stretch of the quarry, as the lizards usually stayed close to the rock
outcrop. They perched and travelled mainly along the bare rock
at the top of the ledge and the shallow layer of bare soil adjacent
to it. As a result of quarrying operations some years before, there
was an exposed vertical rock face four to six feet high below the
ledge, and a taluslike accumulation of sloping rock and soil with
weeds and saplings which the lizards also used. The width of the
outcrop and the rocky area below it averaged about twenty feet,
and the lizards' activities ordinarily were confined chiefly to this
width. Occasionally they ventured out on the adjacent flat, almost
barren, limestone area. Grasshoppers, which constituted the liz-
ards' main food supply, were abundant on this flat, but shelter was
lacking, and at any alarm the lizards would rush back to the ledge.

Because of such occasional use of areas outside the usual sphere
of activity, the extent of home ranges or territories cannot be stated
with precision. One female that was recorded more than 100 times
over a six year period was always within an area of approximately
300 feet x 40 feet. She was known to have shifted her headquarters
several times, and the area covered in any daily or weekly period
would have amounted to only a small part of the total area of 12,000
square feet covered in her lifetime. Individuals differ notably in
the extent of their ranges. A female from the same brood as the
one mentioned above, early became established at the opposite end
of the quarry area. She was remarkably sedentary in her habits,
and it is estimated that she spent at least half of her time within
an area of only a square yard. Nearly always she was seen within
a few yards of the same crevice, into which she quickly retired at
any alarm. But to lay her eggs, she moved along the ledge some
150 feet from her home base. The adult and nearly adult females
present at the quarry at any one time usually arranged themselves
in territories that were mutually exclusive, or nearly so. The females

104 University of Kansas Publs., Mus. Nat. Hist.

are much less aggressive than the males, but occasionally pursuit
and attack of one individual by another has been observed. Indi-
viduals of the opposite sex, so far as observed, were never objects
of territorial hostility. Males and females were often associated in
pairs even at times when there was no breeding activity. The males
have territories that average several times as large as those of
females. In August, 1950, the four adult females present occupied
stretches of ledge from west to east, 100 feet, 70 feet, 60 feet and 25
feet long. At times when only one adult male was present, he would
range along the entire 400-foot length of ledge. The adult males
were generally tolerant of partly grown males and, even though the
latter were se.xually mature, did not pursue or attack them. The
small males were wary of the adults and did not challenge their
dominance.

Ophisaurus attenuatus Baird

Slender Glass Lizard

Status. — Common resident.

Habitat. — This species occurs in grassland portions of the area;
also in woodland edge situations and in open woodland or parkland
with a herbaceous ground cover. It was seen only a few times in
1948 and 1949, when the pasture areas were grazed. The lack of
adequate cover under these conditions was unfavorable, and
probably the lizard was able to persist only because of the cover
provided by marginal situations, along edges of fenced woodlands
and edges of thorn thickets where herbaceous vegetation was at
least partly protected from livestock. In later years glass lizards
have been seen more frequently, and a high proportion of those
seen were young, suggesting successful reproduction and steady
increase with amelioration of habitat conditions after grazing was
discontinued and by 1957 they had become moderately abundant.

Movements. — Over a seven-year period a total of 108 glass lizards
have been captured, marked and released, and 31 have been re-
captured from one to eight times. For most, the records extended
over periods of months. Eighteen were caught in only one year,
six over two years, three were caught over a four-year period, and
four over three years.

These lizards are secretive and seem to travel but little, relying
to a large extent on dense cover and concealing coloration to escape
notice of both prey and predators. Most time is spent underground,
or at least beneath the surface mat of dead vegetation. Many glass

Home Ranges of Vertebrates 105

lizards, including all of those that were recaptured, were known
to live in the field within 300 yards of the Reservation headquarters
where my field work was concentrated. Yet seeing a glass lizard
in this area was a rare occurrence. Even when conditions of
temperature and humidity seemed near optimum for them to be in
the open, the area might be traversed several times by a person
without his seeing one.

Glass lizards were usually found in tall grass and were remark-
ably elusive. One that was startled would dart away with rapid
lateral undulations of its body, usually permitting no more than a
glimpse of it. In this initial spurt the lizard rarely traversed more
than ten feet but it was always difficult to relocate, as its dark
brown color blended well with the background of old dead vegeta-
tion of the surface mat. Relying on concealment the lizard might
permit close approach before moving again. Or it might move
away slowly and stealthily with none of the commotion of its initial
spiut. Occasionally in stalking a glass lizard in high grass I have
"flushed" it four or five times in succession before finally catching
or losing it. Working alone I have most often failed to make a
capture, but when accompanied by one or more other persons, I
have often been able to direct their movements in such a way that
the glass lizard was driven toward me, came within easy reach, and
was secured with a sudden grab. This maneuvering involved in
flushing, stalking and capture of a glass lizard was generally within
an area not more than 20 feet in diameter. If not caught within a
minute or two, the glass lizard invariably found shelter and dis-
appeared. In some instances it may have merely wriggled beneath
the surface mat of dead vegetation. In other instances, however,
glass Hzards were seen to escape into burrow entrances of the
prairie vole. The extensive underground runway systems of the
voles may provide the retreats where the glass lizards spend the
greater part of their time when they are not active on the surface.

In 36 instances the intervals between successive captures ranged
from 280 feet to 10 feet, and averaged 88 feet, indicating home
ranges in the neighborhood of .55 acres. The records are not
sufiBciently numerous to show conclusively the differences between
the sexes, and between young and adults, but they do suggest
that such differences exist. For adult males (20) the average was
98 feet, for immature males (11) 90 feet, and for adult females (5)
43 feet. Those that hibernated one or more times between captures
are not included in these figures. For 15 such intervals the average

106 Unhtersity of Kansas Publs., Mus. Nat. Hist.

movement was 282 feet (850 to 20), indicating that there is some
tendency to shift the home ranges over long periods. However,
only one individual had moved more than 500 feet.

Cnemidophorus sexlineatus Linnaeus
Six-lined Racerunner

Status. — Locally abundant on several small areas of Reservation
until 1956; much reduced in numbers since then.

Habitat. — Patches of barren soil seem to be the essential habitat
requirement; therefore the species has been confined to relatively
small areas on the Reservation. Habitat deterioration has pro-
gressed as luxuriant vegetation encroached on sites that formerly
had a sparse vegetation or were largely barren. The quarry for-
merly supported the largest number. A hilltop field formerly cul-
tivated, and heavily eroded, with topsoil washed away and subsoil
dissected by a system of gullies, supported a moderate population
in 1948. At that time the area was partly barren with sparse vege-
tation of bindweed, three-awn grass, and lespedeza. In the fol-
lowing years as grass and weedy vegetation encroached, bare areas
were increasingly restricted to the bottoms and steep sides of the
gullies, and the lizards became correspondingly scarcer. By late
1952 few remained but in the following four years of drought they
partially regained their numbers. A diversion ditch carrying water
from the pond about 1000 feet along the base of a hillside, between
woods and an open field, was usually dry except after heavy rains
when the pond overflowed. In 1952 the dry channel of this ditch
was occupied by a colony of these lizards, which found the bare soil
of the ditch bottom and banks a favorable habitat, and thrived
there for the next five years.

Movements. — The racerunner is far more active than any other
species of reptile on the area. Individuals move about almost con-
stantly when they are active, and are capable of great speed. In
its normal foraging this lizard moves over areas of bare soil with a
brisk, jerky gait that seems to be half running and half walking.
Individuals foraging in this way have often been followed for as
much as 200 feet in the same general direction.

The racerunner shows a strong tendency to stay in a familiar area.
Many of those marked were recaptured frequently, up to as many
as 25 times, and successive captures were nearly always in the
same general area. On the Reservation the individual areas of the
lizards tended to be long and narrow, probably because barren
ground occurred in strips along the hilltop ledges, gully banks,

Home Ranges of Vertebrates

107

ditch and road. In a more typical habitat of sand dune or open field,
a home range probably would be more nearly circular or oval. For
the entire group a total of 290 movements between successive cap-
tures were recorded and the average movement was 141 feet, but
with notable disparity between the sexes, and between young and
adults. For 102 movements of females the distance was greatest.

Fig. 6. Records of an adult male six-lined racerunner captured
17 times from June 19, 1952, to July 13, 1953. The records for this
individual and most others on the Reservation tend to be linearly
arranged where there are strips of barren soil — along banks and
bottom of a diversion ditch in this instance. Such disturbed situa-
tions provide the final refugia as revegetation proceeds on the

Reservation.

171.3 feet; for 188 movements of males average was 125 feet, and
42 movements of young averaged 99 feet.

The young were of both sexes and various sizes, from hatchlings
to subadults. Observations on hatchhngs indicate that at first they
stay within an area of only a few yards, and that as a lizard grows,

becoming fleeter of foot and
more familiar with its sur-
roundings, the sphere of ac-
tivity gradually enlarges to the
size of the adult's home range.
The partly barren area of
the ditch banks and bottom,
providing favorable foraging
grounds for racerunners, aver-
aged approximately 25 feet in
width. If the lizard wandered
at random along the length of
the territory, it is calculated
that any two records of an
individual would, on the aver-
age, be separated by a distance
35 per cent of the range's full length. On this basis the figures
already cited would represent home ranges of .25 acre for all race-

FiG. 7. Records of an adult female
glass lizard captured eight times in the
period May 30, 1956, to August 27,
1957. The area covered was approxi-
mately 125 feet in greatest diameter.
For male glass hzards ranges recorded
were often somewhat larger.

108 University of Kansas Publs., Mus. Nat. Hist.

runners, .28 acre for adult females, .20 acre for adult males, and
.16 acre for young.

Frequently, both on the Reservation and elsewhere, racerunners
have been seen in locations familiar to me, where none had been
observed previously. Obviously, at times they leave the areas
habitually frequented and make movements of an exploratory na-
ture. Exceptionally long movements recorded for the marked in-
dividuals recaptured in the course of my field work were 1100, 700,
and 600 feet (and back again) for adult males; 900, 750, and 750
feet ( and back again ) for adult females, and 650 feet for a young
male. The previously mentioned hilltop quarry, and the ditch in
the field near the headquarters of the Reservation were separated
by a wooded slope 600 feet across at its narrowest. The woods
were thick, with tangled undergrowth, and seemed entirely un-
tenable as habitat for the racerunners. Nevertheless, several that
were first caught and marked at the quarry were later recaptured
along the ditch bank. Also, on several occasions, lizards that were
not individually identified, were seen in the woods. Most of these
were on a man-made trail where ground vegetation was sparse and
trampled, making running easier for them. Probably this trail
served as an avenue of travel and permitted the racerunners to
cross the wooded slope more easily than they could have otherwise.

Having large feet and powerful claws, racerunners are efficient
burrowers, and excavate underground retreats into which they re-
tire at night and in periods of inclement weather. On many occa-
sions racerunners have been found in small nest cavities beneath
flat rocks. Such cavities do not constitute permanent home bases, as
an individual has rarely been found more than once in any one nest
cavity.

The temporary nest cavities seem to be defended. On several
occasions lizards, even young less than one-third grown, have been
seen to dart out of burrows in pursuit of others that had started to
enter, and chase the intruder for several yards. Chasing has been
noted frequently, but it is usually sexual pursuit or masculine hos-
tility not involving the defense of a definite territory. Individuals
overlap extensively in their ranges. In favorable habitat a given
situation may be used by several or many individuals of both sexes.

Home Ranges of Vertebrates 109

Lygosoma laterale ( Say )
Ground Skink

Status. — Uncommon resident.

Habitat. — ^Woodland and woodland edge are preferred; the rec-
ords of this species on the Reservation are perhaps too few to de-
fine its habitat requirements with precision, but seem to indicate
that it occurs in all types of woodland, and ordinarily does not in-
vade open fields beyond the shade of adjacent woodland. It is
usually found in ground litter of fallen leaves and has been found
most often in situations dominated by oak or hickory. One was at
a board pile in the edge of a field of brome grass a few feet from the
edge of hickory woods. Several were taken at a rock pile near the
pond, adjacent to woodland. Several were found in grass in a
parklike situation beneath large elms near the headquarters. One
was found in bluestem prairie about 40 feet from trees and brush
near its border.

Movements. — No records of movements were obtained; of a
dozen individuals marked and released none has been recaptured.

Eumeces fasciatus Linnaeus
Five-lined Skink

Status. — Resident throughout the wooded parts of the Reserva-
tion.

Habitat. — Occurs in all types of woodland but in varying abun-
dance; most numerous in hilltop ledge situations in open woods, and
least numerous on heavily wooded north slopes where thick canopy
permits Httie penetration of sunlight to forest floor. Flat rocks for
shelter are an important requirement and where they are scarce or
absent skinks are correspondingly scarce.

Movements. — In a recent publication (Fitch, 1954:99-115) I have
discussed in some detail the data concerning movements of five-lined
skinks accumulated over a four-year period. Several hundred skinks
were marked and released, mostly on study areas of only a few
acres. A total of 323 marked skinks were recaptured, many after
one or more hibernation periods. None was captured frequently.
Seven captures were the maximum recorded for any individual, and

110 Unintrsity of Kansas Publs., Mus. Nat. Hist.

these were spread over a period of nearly four years. In almost
every instance of recapture of a marked skink, the actual distance of
travel and the time, frequency and motivation were unknown.
Although the records were inadequate to show the home range of
any one individual, they were sufficiently numerous to show the
trends according to sex, age, and lapse of time. Five-lined skinks
are secretive, living where there is abundant shelter in the form of
leaf litter, decaying wood and rocks. They stay under cover most
of the time, and when travelling, they move furtively, always keep-
ing near cover. The home range usually centers about a decaying
log or comparable shelter, and the size and shape of the range is
altered accordingly. Home ranges averaging approximately 90 feet
in greatest diameter are indicated both for adult males and young.
These skinks seem to have no regular home base, but use any con-
venient shelter within the area when they are inactive, or escaping
from enemies. The adult females, however, have regular nest bur-
rows in which they usually stay, except for occasional brief emer-
gences to find food, from the time they become pregnant in early
May until after the eggs hatch in July or August. On their brief
forays for food these nesting females usually venture only a few
yards from their burrows. Although they occasionally leave the nest
burrow and eggs to forage, they then range much less widely than
at other times. Several male skinks that were recaptured repeatedly
were shown to have moved back and forth within areas 200 to 275
feet in greatest diameters, which constituted their home ranges.
Others were limited to much smaller areas. A rock pile of approxi-
mately 70 x 30 feet harbored many skinks of both sexes and various
sizes because food and shelter were abundant there. The rock pile
was bordered on three sides by areas unsuitable as habitat, and on
the fourth side the habitat also was much less favorable than in the
rock pile itself. The rock pile or parts of it constituted the entire
home range for all the skinks present there. These skinks are not
territorial but males are hostile and fight on sight in the breeding
season.

Eumeces obsoletus Baird and Girard

Great Plains Skink

Status. — Confined to relatively small parts of area where its
habitat is present; there moderately common.

Habitat. — This large skink typically lives in open grassy situa-
tions with large flat rocks. Hundreds of captures have been re-
corded, mostly within four small areas totalling only a few acres

Home Ranges of Vertebrates 111

in extent. Perhaps not much more than one percent of the area is
regularly occupied, and the habitable area steadily diminished as
ecological succession proceeded with woody vegetation steadily
encroaching. Under original conditions, with the area mostly
prairie, the hilltop rock ledges several miles long, must have sup-
ported a large population of these skinks. By 1955 they dwindled
to four separate reHct colonies, one at the quarry, and the other
three along stretches of rock ledge and adjacent woodland that
were grazed by livestock until 1949 (the remainder of the rock
ledge and woodland was fenced and protected from livestock
beginning in the 1930's).

Movements. — "The usual concept of a home range does not apply
well to the Great Plains skink despite its tendency to remain within
a small area. One skink may spend many days in its small burrow
system under the same rock, or else if it does emerge at all it makes
only short trips and then returns. But such shelters and burrow
systems are temporary, generally occupied for only a few days, or,
at the most, a few weeks. Then the skink moves on to another
site, often only a few yards away. Later as it shifts to still other
successive locations, it ordinarily does not move farther in the same
direction, but tends to keep within a small area, probably using the
same travel routes and depending on the same landmarks. The
Tiome range' encompassing the area to which normal movements
are confined, is, however, ill-defined, because it is covered infre-
quently and has no definite boundaries, and its size, shape and
position are ephemeral." (Fitch, 1955: 69.)

Great Plains skinks were much less numerous than five-lined
skinks on the Reservation, but, for individuals, records of recapture
were more frequent. Over a four-year period 634 captures were
recorded for 208 individuals. In general, individuals tended to
stay within areas more than 50 feet but less than 150 feet in greatest
diameter. Males are more vagile than females and adults are more
vagile than young. For twelve adult males each recorded in at least
four locations, the average distance between most remote points
of capture was 125 feet; for six adult females and seven juveniles
comparable figures were 75 feet and 60 feet respectively. All these
were selected individuals that moved back and forth without making
long shifts. For 226 recorded movements of marked individuals,
between successive captures, mean distances were as follows:
adult males (80), 70 feet; adult females (51), 31 feet; young (95),
39 feet. The higher figure for adult males is influenced by the

112 UNiNTRSiTi' OF Kaxsas Publs., Mus. Nat. Hist.

fact that they are more inclined to shift. Eliminating the few
movements of more than 200 feet, which are obviously shifts in
most instances, average movements are: adult males, 51 feet; adult
females, 19 feet; young, 30 feet. On this basis it is estimated that
home ranges are .2 acre or less in adult males and of even smaller
size in the females and young. Not included in these figures are
the 45 movements in the intervals that included one or more hiber-
nation periods, 121 feet for 16 adult males, 119 feet for 10 adult
females, and 81 feet for 19 young. These figures suggest that the
lapse in day to day routine and the seasonal change in habitat
associated with hibernation often induce individuals to shift to new
home ranges. These skinks often do shift away from an original
home range to a new area more or less remote from it. Over period-;
of months or years many of the skinks on the study areas turned
up at locations hundreds of feet from the areas encompassed by a
series of early capture records. After such a long movement a
skink never returned to an original home range. The longest
record of movement was one of 500 feet ( 830 feet if the skink kept
to the usual rock ledge habitat) in an individual whose records
extended over 47/2 months.

On many occasions skinks seemed to appear abruptly on study
areas that had long been under intensive observation. After an
original capture some such individuals were often recorded regu-
larly over periods of weeks or months, indicating that they had
settled in the new location after immigrating from a more or less
remote location.

Unlike most other kinds of reptiles occurring on the Reservation,
Great Plains skinks dig well in soil that is loose and moist, and
they spend most of their time in the burrows that they have ex-
cavated. The burrows may be several inches or several feet in length
and usually they are under rocks. A skink may stay in such a bur-
row for periods of days, then may emerge, wander briefly, and begm
a new burrow a few yards away. In the breeding season, May and
June, adults are often found in pairs in their burrows imder large,
flat rocks. Such burrow systems may be thought of as temporary
small breeding territories because they are never occupied by more
than the pair of adults, and the male will viciously attack intruding
males.

Although these skinks seemed to be disappearing from the Reser-
vation by 1955, they made surprising gains in numbers in 1956 and
1957, and even appeared in several areas where they had not been
recorded previously.

Home Ranges of Vertebrates 113

Eumeces septentrionalis Baird

Prairie Skink

Status. — Only three recorded on Reservation, perhaps survivors
of relict colony.

Habitat. — All three skinks were found on lower slopes of hillsides
having southern exposures, in grassy places sparsely wooded with
osage orange, honey locust, dogwood and haw, grazed by livestock
until 1949. As compared with most other places in northeastern
Kansas where the species has been collected, these sites differed in
having encroaching woody vegetation, and more rank herbaceous
cover. Discontinuance of grazing is resulting in the elimination of
this sldnk's habitat.

Movements. — In the summer of 1953 one was caught twice, at
stations 100 feet apart in funnel traps set in a ditch between wood-
land and grassland. Breckenridge's ( 1943 ) study of marked indi-
viduals of a local colony of these skinks suggests that the species is
at least as sedentary in habits as is E. jasciatus or E. obsoletiis.

Natrix sipedon Linnaeus

Common Water Snake

Status. — Uncommon, and probably irregular, resident, its number
influenced by uncertain water supply.

Habitat. — Mainly in vicinity of permanent water supply, either
stagnant or running; in spring and autumn has been found along
hilltop rock ledges far from water, and evidently seeks such situa-
tions to hibernate.

Movements. — Of 23 water snakes marked and released, four have
been recaptured. A large adult female caught twice in 1949 was re-
captured in 1952 and 1953. All captures were at the pond or near
it, with the exception of the second 1949 record, which was some
450 feet farther west along a diversion ditch. Of two young females
recaptured, one was taken in early September and again the follow-
ing May in the same rock pile near the pond. The other was recap-
tured after a three-month interval less than 100 feet from the original
location, beside the pond. A juvenile was recorded three times
within a 15-foot radius, at the Reservation headquarters in Septem-
ber 1953. The records of these several marked individuals that were
recaptured demonstrate that individuals may live for periods of
months or years where there is a limited and isolated water supply.
The other records suggest that individuals occasionally wander long

114 Univ'ersity of Kansas Publs., Mus. Nat. Hist.

distances to find such habitat. Stickel and Cope (1947:129) re-
corded one recaptured at a distance of 380 feet after a lapse of two
years.

Storeria dekayi Holbrook

Brown Snake

Status. — Uncommon resident; actually may be more numerous
than some of larger snakes that are considered moderately common,
but is seldom seen because of small size, secretive habits, inconspic-
uous dull coloration, and preference for situations with dense ground
cover.

Habitat. — Woodland in damp, well-shaded situations where leaf
canopy is dense; also in wet situations near stagnant or running
water.

Brown snakes are most commonly found in late March or April,
under flat rocks, often associated with ring-necked snakes and worm
snakes.

Movements. — No information has been obtained; none of several
brown snakes marked and released has been recaptured.

Thamnophis sirtalis Linnaeus
Common Garter Snake
Status. — Common resident.

Habitat. — Damp situations wherever its prey, earthworms and
amphibians, are available. The pond and the two small creeks
provide the most preferred habitat, but this garter snake also ranges
far and wide through moist fields and woodlands. Garter snakes
have been caught in greatest numbers in October, in funnel traps
set along hilltop rock ledges, which are preferred hibernation sites.
The ledges chosen are those in warm and dry situations, with south-
facing exposures.

Movements. — A total of 150 garter snakes had been marked and
released on the area and 22 had been recaptured by the end of
1956. Twelve of these records were of snakes recaptured after
one or more hibernation periods. In eleven instances the distances
between successive capture locations averaged 688 feet (777 feet
for seven males and 532 feet for four females) indicating home
range areas in the neighborhood of 34 acres (or 43 acres in males
and 20 acres in females, if the differences between the sexes in this
small sample are representative). Movements to or from hilltop
rock ledges, where hibernation dens usually are located, were not

Home Ranges of Vertebrates 115

included in the computation of average movement between cap-
tures. In most instances travel to or from the hibernation den
must have taken the snake well beyond the limits of the usual
range. The longest movement recorded was one of 2300 feet, in a
female first caught as a juvenile at a hilltop ledge on October 25,
1950. An adult male recorded at a hilltop ledge twice in Novem-
ber was recaptured 1040 feet from there in a bottomland meadow
the following May. No garter snake was recaptured in the same
denning area after an intervening summer, although in several
instances a snake was recorded more than once in a denning area
in the same autumn, or in autumn and again in spring. From the
trend of the records, it seems that garter snakes have some attach-
ment to definite areas, but tend to range more widely than most
other snakes, and, over periods of months or years may wander
into new areas. Of 97 records on the area, 49 were at hilltop ledges,
30 were in the vicinity of water including several at a rock pile
near the pond which was also used for hibernation, 13 were in
grassland and five were in woodland. Double or multiple catches
were made in funnel traps much more often than might have been
expected from chance. The catch might consist of three males
(once), two males and a female (twice), two males (twice), two
females (twice), male and female (five times). Most of these
catches were made along hibernation ledges, and it seems probable
that trailing by scent of one snake by another was responsible for
the double or multiple captures. This idea is borne out by the fact
that when a garter snake was removed from a trap on one day, a
second was often caught on the day following.

Carpenter (1952:250) studying a population of this species in
Michigan found the average "activity range" to be about 600 feet
in length and 150 feet in width, thus having an area of 2.07 acres.

Haldea valeriae (Baird and Girard)

Smooth Earth Snake

Status. — Rare and seldom seen (only two records) because of
secretive habits, but probably resident on Reservation in small
relict population.

Habitat. — This small, burrowing snake lives chiefly in deciduous
forest. Here, in the northwestern corner of its range, it seems to
prefer open woodland with rocks and brush but not a continuous
leaf canopy. The species is known from only five localities in
Kansas (Smith, 1956:287), these clustered in or near Douglas
County, where the Reservation is situated.

116 Universiti- of Kansas Publs., Mus. Nat. Hist.

Diadophis punctatus Linnaeus

Eastern Ring-necked Snake

Status. — Abundant resident; at times this snake probably outnum-
bers, in individuals, all other kinds of vertebrates on the Reserva-
tion.

Habitat. — Optimum habitat seems to be in open woodland with
abundant flat rocks. Ring-necked snakes do not often bask in direct
sunlight, but regulate their body temperatures by maintaining con-
tact with the undersides of rocks warmed by sunshine. They are
largely subterranean in habits. They are relatively scarce in wood-
land with a continuous leaf canopy shading the ground. In meadow
habitat, where rocks are absent, the snakes are difficult to find, but
have been seen on many occasions crawling through tlie grass or
crossing roads. On several occasions they have been found in
underground tunnels of the prairie vole. Several have been found
in the humus and litter beneath the stick houses of woodrats.

Movements. — The ring-necked snake is probably the most abun-
dant reptile of the Reservation, but because of its secretive tenden-
cies, attempts to study it were remarkably improductive. A total of
333 have been marked and released, mostly in 1950 and 1951, but
only 11 recaptures for ten individuals have been recorded. Most
of the individuals marked were in small areas of favorable habitat
where intensive field studies were in progress, which should have
greatly increased the chances of recoveries. However, of the ten
individuals recaptured, seven were taken after intervals of less than
a month, and perhaps had not had time to wander far from the
original location. An eighth was a juvenile caught and marked
shortly before going into hibernation (October 2) and recaptured
the following spring at the same place soon after emergence (April
26). Of these eight, two had moved 60 feet, one 30 feet, and the
remaining five had made no measurable movement. An adult male
caught and marked on April 26, 1950, was recaptured 90 feet away
on May 1, 1951. An adult male marked on April 27, 1951, was
recaptured 350 feet away on May 22, 1954.

These small snakes may not have regular home ranges as they
seem to wander widely. Each year in April and May, ring-necked
snakes were found in large numbers in certain rocky places that
were evidently optimum habitat at that time of year or served as
hibernation sites. Most of the individuals marked were those found
in such places, which had an almost complete turnover of the

Home Ranges of Vertebrates 117

population from year to year. Evidence of wandering was obtained
frequently. The grassy field where the Reservation headquarters
are situated was much less favorable habitat than the rocky slope
where the snakes were more often found. However, on many occa-
sions, when temperature and humidity were favorable, the snakes
were seen crossing the road or crawling tlirough high grass. Strips
of tarpaper were placed on the ground in many places in the field.
When weatlier was moderately cool and soil was moist, ring-necked
snakes usually could be found under these strips, but none of these
was ever recaptured.

Carphophis amoenus Say
Worm Snake

Status. — Common resident.

Habitat. — This snake occurs throughout the wooded portions of
the Reservation and along woodland edges. Most individuals have
been found in open rocky woods. Nearly all were uncovered in
loose soil beneath large flat rocks. These snakes have been found
chiefly in the rock-strewn portions of the woodland, but perhaps
are not actually concentrated in such situations to the extent that
my records would suggest, since it would be far more difficult to
find them in the absence of rock cover. Isolated large rocks in wood-
land often have worm snakes hiding under them. Loose damp soil
is a necessity. The snakes are found mainly in April and May, and
are little in evidence later in the season, when evidently they retreat
deep underground. In periods of rainy weather when the soil is
unusually damp, they may be found occasionally in the types of
situations frequented in spring, while in periods of dry weather,
they cannot be found at all.

Movements. — Worm snakes usually stay beneath the surface of
the soil. They are more specialized for fossorial life than any other
species of reptile occurring in the area. The head is small and flat-
tened, with snout spade-like, lower jaw counter-sunk, eyes minute,
and cephalic plates reduced. The neck is especially thick and mus-
cular, adapted for driving the head through soft soil. I have never
found a worm snake traveUing above ground in the open. Such
travel is indicated by occasional individuals caught in pitfalls or
funnel traps, but probably they emerge only under cover of dark-
ness.

Of 137 worm snakes marked and released, nine have been recap-
tured. One was recaptured four times, another twice and the others

118 UNiNTRSiTi' OF Kansas Publs., Mus. Nat. Hist.

only once each. Seven were recaptured only in the same season they
were marked; the other t\vo were recaptured in the following season
after lapses of 12 months and IO/2 months. A large adult female had
moved 410 feet between May 19 and June 7, 1951, but all other in-
dividuals were much more sedentary. An adult male had moved 50
feet between May 30, 1949, and April 14, 1950. An adult female had
also moved 50 feet between April 30 and May 25, 1951, but on May
29, May 31, and September 14, she was found within five feet of
the second location. An adult male had moved 20 feet in 38 days.
Each of the other individuals either was recaptured under the same
rock where the snake was captured originally or had moved no more
than five feet. Span of records between first and last captures for
these individuals were: 352, 5 (witli three captures), 5, 4, and
2 days.

The trends of these meager data seem to suggest that worm snakes
have no home range in the usual sense, but tend to stay for long
periods in the soil at the same place, moving mainly in vertical
planes in response to changes in temperature and moisture. Oc-
casionally individuals may make relatively long shifts, perhaps wan-
dering at random, and then may settle in new locations. There is no
evidence that the snakes ever return from such wanderings.

Coluber constrictor Linnaeus
Yellow-bellied Racer
Status. — Common resident.

Habitat. — Occurs throughout the entire area with seasonal shifts.

In summer it ranges over grassy fields, especially those of the two
small valleys, also in bluestem prairie, along woodland edge thick-
ets, and in open woodland where some herbaceous ground cover
is present. In fall most of the population or all of it resorts to hill-
top ledges for hibernation, after moving across wooded areas to
reach these sites, in some instances.

Movements. — By July 15, 1955, a total of 337 yellow-bellied racers
had been marked and released on the Reservation and 74 had been
recaptured, mostly after substantial intervals. Forty-two were
recaptured after lapse of a year or more. Four was the maximum
number of captures for any one individual.

In the area of the Reservation, yellow-bellied racers spend most
of the active season in prairie, pasture, or hay field habitat or oc-
casionally can be found in glades of open woodland. However,

Home Ranges of Vertebrates 119

in autumn there is a general shift to hilltop rock outcrops, where
clefts and fissures provide dens for hibernation. After moving to
the rock outcrops the population is much more concentrated than
it is on the summer range. As a result, most of the individuals re-
captured (48 of the 74 total) were taken only along hilltop rock
outcrops. These were classed as follows: 8 adult males, 11 adult
females, 17 yearling males, 7 yearling females, 4 hatchling males,
and one hatchling female. Although these groups are too small for
statistical comparison, the trends do not appear to be significantly
different. Excluding several that shifted from one hilltop ledge to
another distant one, the mean movement for 48 was 210 feet, rang-
ing from 0 to 875. For the 26 recaptured after intervals of a year
or more, the mean distance was 174 feet. Individuals may return
year after year to the same part of the hilltop rock outcrop. After
55 months a large female was found 300 feet from the original site
of capture. Another was found at 390 feet after 43 months.

Eleven individuals were caught both at a hilltop rock outcrop,
where presumably they had come to hibernate, and in a bottom-
land valley where presumably they spent the summer, providing
some idea of the distance of travel to and from the hibernation
ledges. Distances ranged from 500 to 1360 feet and averaged 770.
The tendency seems to be for the racers to seek the nearest suitable
stretch of ledge rather than to make an unnecessarily long trip to
one that is farther away. Hilltop fields near or adjacent to the
ledges where snakes hibernated also provided suitable habitat.
Two individuals were caught both at a ledge and in a hilltop field,
and they had moved distances of 230 feet and 1350 feet.

Twenty-three individuals were recorded and then recaptured
away from outcrops in the areas where, presumably, they live during
the summer. One of these was caught as a hatchling and recaptured
19 months later at a distance of 1500 feet. This long movement
probably represents one or several shifts rather than normal use of
a home range. Another hatchling male had moved 470 feet in 18
months. The remaining 21 movements were all of adults. For 13
records of males the average was 567 feet, indicating an average
home range of 23 acres, and for nine records of females the average
was 367, indicating home ranges of 9.7 acres.

Eight of the individuals captured at hilltop ledges had made
relatively long movements, probably having shifted permanently
away from the original site of capture. These movements were:

120

University of Kansas Publs., Mus. Nat. Hist.

Fig. 8. (A) Capture locations of an adult male yellow-bellied racer
(6 records, October 21, 1953, to July 15, 1957); and (B) capture locations of
an adult female. In each instance summer records were in a grassy bottom-
land field, but fall records were at hilltop rock ledges where the snakes came
to hibernate after moving 138 to 200 yards up a wooded slope.

3150 feet in six months, hatchling female; 3000 feet in two months,
yearling female; 2950 feet in 31 months, adult female; 2900 feet in
three months, yearling male; 2700 feet in two months, subadult
female; 2430 feet in two months, adult female; 2250 feet in twelve
months, yearling female; 1250 feet in six months, yearling male.
Because field work was mostly limited to a square mile area, and
was concentrated to a large extent on relatively small parts of it,
the chances of recapturing individuals that made long movements
were poor. The number of long movements actually recorded
therefore seems high. Possibly much longer movements may be
made occasionally. Much more information is needed concerning
the vagility of the species. With the information available at
present it can be said that individuals have a strong tendency to
stay within the same home range, and to traverse an area that may
be several times the diameter of the home range to return to the
same hibernation shelter each autumn, but that some, of both
sexes and various ages, may shift permanently for half a mile or
more beyond the original range of activities.

Home Ranges of Vertebrates 121

Elaphe obsoleta Say
Black Rat Snake

Status. — Moderately common resident, evidently utilizing all
parts of area.

Habitat. — Tliis snake is characteristic of deciduous forests; on
the Reservation it has been found most often along hilltop rock
ledges. It is notorious as a climber and raider of birds' nests. It
often has been found along woodland edge and even in open fields.
The open field situations are utilized chiefly in summer, when
there is high grass.

Movements. — Of 159 black rat snakes marked and released on
the Reservation 22 were recaptured. Many of those marked were
small young which probably had a short life expectancy and, in
some instances, perhaps had not yet established themselves in
definite areas. Some adults were caught in outlying areas that
were seldom visited in the course of routine field work, and the
opportunities for recapturing them were limited. For the adults
captured within the area where field work was concentrated, the
incidence of recaptures was relatively high. Span of records, in
months, for the 22 individuals were: 49, 48, 48, 44, 42, 32, 25, 24,
22, 21, 20, 20, 17, 15, 12, 12, 12, 10, 8, 1, 1, 1, and 0. From the
trend of these records it is evident that adults and well grown young
remain in familiar areas for periods of years, or perhaps for life.
Greatest distances recorded as covered by black rat snakes were:
2200 feet (adult male in 44 months), 1680 feet (adult male in 49
months), 1650 feet (adult male in 21 months), 1600 feet (hatchling
in 32 months), 1560 feet (adult male in 42 months), 1450 feet (half-
grown male in 24 months), 1200 feet (adult male in 24 months),
1100 feet (hatchling male in 22 months). Thirteen others had
moved distances of less than 1000 feet.

Several of those that made long movements were recaptured
later, back nearer the original capture point, suggesting that they
were still within a home range. For example, an adult male first
recorded on October 5, 1949, was recaptured 1900 feet away on
June 12, 1950. On April 21, 1953, he was captured a third time 2200
feet from the second location but only 300 feet from the first location.
Another adult male was caught on July 16, 1950, September 19,
1950, October 3, 1950, May 15, 1951, June 12, 1952, June 17, 1954,

122

UNTVERSITi' OF KANSAS PuBLS., MUS. NaT. HiST.

and July 19, 1954, The second and third captures were at approxi-
mately the same place on a hilltop ledge more than 1000 feet from
the other six capture points, which were all within a hundred-foot
radius near the Reservation headquarters. Another adult male
caught on June 6, 1950, June 2, 1952, April 27, 1953, May 9, 1954,
June 6, 1954, and July 27, 1954, was found at four places in the
headquarters field, over a distance of 1380 feet, and shifted back and
forth between this field and a hilltop ledge three difiFerent times,
covering distances of 850 to 1350 feet in doing so.

Fig. 9. Capture locations of an adult female black rat snake recorded
eight times in a six-year period.

For the total of 38 movements between successive captures, the
average distance was 817 feet. Stickel and Cope (1947: 129) re-
corded ten movements of black rat snakes, ranging from 130 feet
to 1760 feet and averaging 767, on the Patuxent Research Refuge
in Maryland. Marked disparity between the sexes in vagility was
evident in the trend of my records. For the 29 movements of males
the average was 897 feet, indicating a home range of 57.6 acres,
while for nine movements of females the average distance was 559
feet, indicating a home range of 17.9 acres.

Pituophis catenifer Blainville
Bullsnake

Status. — A few individuals are permanent residents on the Res-
ervation, but others from adjacent cultivated land resort to the
area to find hibernation shelters along hilltop rock ledges.

Home Ranges of Vertebrates 123

Habitat. — Bullsnakes live primarily in grassland but also in open
woodland. Fields of alfalfa and other hay crops in nearby farm-
lands provide a habitat more favorable than any to be found on
the Reservation itself, perhaps because of the abundance in them
of pocket gophers, which constitute the chief food of the bullsnake.
On the Reservation bullsnakes have been found from time to time
in brome grass fields and also have been found in summer in open
rocky woods on south-facing slopes. The majority of those seen
were individuals caught in funnel traps along hibernation ledges
in October.

Movements. — Eighteen bullsnakes have been marked and re-
leased on the area. Of the 23 captures 17 were made in late Sep-
tember or October, or in early spring, along rock ledges where pre-
sumably the snakes had come to hibernate. One large female was
caught on September 22, 1951, crossing a county road at the edge
of an alfalfa field. Nine days later she was caught 2700 feet away
at a hilltop rock ledge on the Reservation. Another adult female
was caught at sites along the ledge 310 feet apart on October 17,
1950, and September 1, 1953.

Some of the records suggest that these snakes trail each other by
scent to the hibernation dens. On "Sugarloaf," a hilltop overlook-
ing an expanse of cultivated fields to the west, three large adults
were caught within a few days in the autumn of 1950. In October
of 1949, 1950 and 1951, an adult female was found along a 200-foot
stretch of ledge where evidently she returned each year to hibernate.
She was found 420 feet farther along the same ledge in June, indi-
cating that her summer range also extended into this same area.

Imler (1945:273) recaptured 12 marked bullsnakes in Nebraska
and found that 11 had moved distances less than 300 yards, while
the twelfth had moved IM miles. Fitch (1949:556) working with
the subspecies catenifer in California, recorded movements of less
than 1000 feet in 25 of the 28 snakes recaptured.

Lampropeltis calligaster Harlan
Prairie King Snake

Status. — Scarce, but seems to be regular resident on Reservation.

Habitat. — Open grassy situations with flat rocks seem to be pre-
ferred. Several have been trapped in fall along old rock walls, and
rock ledges in thick woods of oak, hickory and elm. Several have
been found in brushy pastures of brome and bluegrass.

Movements. — Over a seven-year period, 16 have been marked

124 Unrtrsit^' of Kansas Publs., Mus. Nat. Hist.

and released on tlie Reservation, but none has ever been recap-
tured. The secretive and partly subterranean habits would account,
in part, for this lack of success, but it also seems probable that the
snakes wander widely.

Stickel and Cope (1947:129) recorded two recaptures of this
species (subspecies rhomhomaculata); one was recaptured at the
same place after one day; the other had moved 600 feet in six weeks.

Lampropeltis doliata Linnaeus

Milk Snake

Status. — Rare, probably with small permanent population tend-
ing to be localized in few small areas that satisfy habitat require-
ments.

Habitat. — Grassy, rock-strewn areas, usually in open woodland or
at woodland edge.

Movements. — Over an eight-year period ten milk snakes have
been marked and released, and two have been recaptured. A small
juvenile marked and released on June 16, 1951, was found again
after 31 days, beneath the same piece of rusty sheet metal where
it was originally discovered. A half-grown individual marked and
released on May 26, 1955, was recaptured 12^ months later at a dis-
tance of 1320 feet from the original location, and had grown to
small adult size.

Tantilla gracilis Baird and Girard
Slender Flat-headed Snake

Status. — Rare and seldom seen (only three records) because of
secretive habits, but probably resident in small relict populations.

Habitat. — The three found on the Reser\'ation and a few found
at other places in northeastern Kansas were all on sparsely wooded,
rocky hilltop edges of southward exposure. Of those found on the
Reservation, one was at the old quarry, the other two were found in
1950 and 1956 at almost the same spot of "Rat Ledge."

Agkistrodon contortrix Linnaeus
Copperhead
Status. — Common resident.

Habitat. — Living chiefly in woodland, the copperhead is one of
the species most characteristic of the deciduous forest climax. It
ranges throughout all parts of the woodland, but with some change
in preference according to weather and season. Rocky upper slopes

Home Ranges of Vertebrates 125

near hilltop ledges in woods of chestnut, oak, hickory and elm per-
haps provide tlie optimum habitat, and provide hibernation sites
where there is some concentration in fall.

Shelter is provided by crevices in the ledge, and by loose flat rocks.
The rocks selected for hiding places are often massive slabs several
inches thick and several feet wdde. In summer the snakes undergo
some dispersal from these ledges throughout the woodland, wood-
land edge, creekside and roadside thickets, grassy fields, both in the
bottom lands and on hilltops, and even to hay fields adjacent to
the Reservation.

Movements. — Copperheads are sluggish by nature and seem to
obtain their prey by ambush rather than by active search. Their
movements are therefore usually short and leisurely, and are gov-
erned to a large extent by changes in temperature and humidity
during the daily cycle. The fact that activity is largely nocturnal
makes observation difficult.

A total of 94 individuals caught and marked on the area have
been recaptured, after intervals of from a few days to five years.
Two (both adult females) were released at distances from their
capture points and they had made unusually long movements 2870
feet and 600 feet when recaptured after intervals of four months
and six months. The remaining 92 individuals averaged a distance
of 455 feet between capture points. The significance of this aver-
age figure is lessened by the fact that trends differed notably be-
tween the sexes and in different age groups. For 31 adult males
the average of 717 feet is approximately twice that in the 25 fe-
males— 358 feet. Probably difference in vagility between the sexes
is even greater than these figures suggest, because of the 11 males
that had moved less than 100 feet nine were recaptured after rela-
tively short intervals of about a month, or less. Restlessness and
frequent activity caused these males to be recaptured in traps or
found in the open before they had moved far from the sites of orig-
inal captures. However, of the six females that had moved less
than 100 feet only three were captured in intervals of less than a
monLh; others were recaptured after intervals of 45 months,
21 months, 20 months, 11 months, one month, and one month. For
36 other copperheads that were marked as young of various sizes,
and recaptured, average distance was 298 feet. Thus it seems that
adults of either sex are somewhat more vagile than the young.
Among the juveniles, females had moved somewhat farther on the
average than had males.

Difference in vagility between the sexes in adults is best demon-

126 Untversity of KaxNsas Publs., Mus. Nat. Hist.

strated by eliminating from consideration all those individuals re-
captured after less than six months. For the 15 males distances
ranged from 205 to 3040 feet and averaged 1323; for the 15 females
distances ranged from 0 to 1950 feet and averaged 292.

Of the 25 adult females two had made outstandingly long move-
ments: 1950 feet and 1350 feet. These probably represent shifts
from one home range to another, and therefore should not be in-
cluded in a computation of home range size. The remaining 23
movements are rather evenly distributed from zero up to 500 feet
(with one of 655 feet), and they average 246 feet. Most probable
home range diameter is, then, approximately 500 feet, representing
an area of roughly 6.5 acres.

Sedentary nature of the adult females is demonstrated also by the
frequency with which individuals were recaptured as compared
wdth males and young. Of the individuals captured three times,
seven were adult (or subadult) females, one was an adult male,
two were immature females, and one was an immature male. Of
individuals caught four times, two were adult females and one was
a half-grown male. The only individual caught five times was an
adult female. She was caught on June 27, 1951, July 17, 1951,
August 3, 1951, August 6, 1953, and September 26, 1953, all within
a one hundred-foot diameter.

The nature of long-time movements in adult males remains some-
what obscure. Of the 31 recaptured, 8 made movements of 1000
feet or more as follows: 3040 feet in 12 months, 2750 feet in 12
months, 2250 feet in 32 months, 2250 feet in 26 months, 2250 feet in
11 months, 1300 feet in 12 months, 1120 feet in 8 months, and
1000 feet in 19 months. On the other hand prolonged occupancy
of a relatively small area is suggested by the following records:
700 feet in 57 months, 650 feet in 21 months, 300 feet in 17 months,
270 feet in 39 months, 210 feet in 20 months, 205 feet in 37 months,
90 feet in 6 months, 60 feet in 6 months.

Available observations suggest that newborn copperheads are
remarkably sedentary and that they extend their spheres of activity
slowly over periods of weeks, as they gradually become familiar
with their surroundings. On September 17, 1948, in a deep crevice
of a limestone outcrop I found shed skins of several young copper-
heads entwined in the same mass of sticks, dry leaves, and cobwebs.
Obviously these were snakes of the same litter, which had all shed
about the same time, before they dispersed. According to my own
findings and those of Gloyd (1934:600) young born in captivity

Home Ranges of Vertebrates 127

most often do not shed their skins for a number of days after birth.
On September 24, 1950, a Utter of young copperheads born in
captivity was released at the spot where the female was originally
found. On the afternoon of September 26 all seven were found
again at the same place. The day was cloudy and humid, about
72° F., and all the young were in tight circular resting coils in the
open. Four were within a ten-inch radius just where they had
been dumped from the sack 48 hours earlier, under a skunkbush
( Rhus trilobata ) . Some two hours later, when the temperature was
falling rapidly, the spot was checked again. One snake had disap-
peared and another had begun to move toward shelter of the rock
cleft. On September 28 four of the litter were again found at the
same place, but a Httle more scattered, all within a ten-foot space.
On September 30 five of the litter were found at the same place, and
four were found again on the following day. Two were found
there again on October 3 after a minimum night temperature of
approximately 40° F.

Gloyd (1934:592) on three occasions found associations of gravid
females, and found other reports in the literature. He speculated
that the gravid females tend to assemble in places where conditions
provide favorable shelter. On August 3, 1950, I turned a flat rock
two feet in diameter and two to three inches thick and found two
gravid females. They were coiled in a nestlike depression which
had the damp earth of its sides packed and smooth as if the snakes
had been there for a long time. While I was catching these two
a third gravid female was heard gliding through the grass toward
the rock, and it also was captured.

I found a second aggregation on August 9, 1950, about 400 feet
from where the first group was found, under a large flat rock that
was exposed to sunshine for several hours during the middle of the
day. Four gravid females were lying in resting coils in contact
with each other. When uncovered, they began to move about
restlessly but seemed reluctant to leave the depression. While I
was catching three, the fourth escaped into thick brush nearby. On
the following day the fourth female was found within a few feet of
the rock where the group had been, in a resting coil on the sticks
of an old woodrat house beneath a bush and partly in the sunshine.

On the Reservation copperheads are too scarce to be seen fre-
quently, and rarely is more than one found in a day, even though
intensive search is made, and many rocks suitable for shelter are
turned. Shelter is abundant, and similar conditions of temperature

3—3117

128 UNi\'ERSi'n' OF Kansas Fuels., Mus. Nat. Hist.

and humidity can be found under hundreds of different rocks within
a small area. Therefore it seems that the aggregations of females
found are not due to chance meetings at sites where physical con-
ditions are favorable but that the females have a definite affinity
for each other causing them to seek others and remain associated
v.ith them, in suitable shelters, for extended periods.

Crotalus horridus Linnaeus
Timber Rattlesnake

Status. — Rare resident.

Habitat. — This rattlesnake prefers deciduous forest, especially
along hilltop rock outcrops in thick woods. Individuals range
widely and may temporarily invade other habitats. One was
found in brome grass at the edge of a field a few yards from edge
of the woods, near the pond. One was killed in a hayfield adjoining
the Reservation. The few records on the Reservation are well
scattered.

Movements. — On September 25, 1950, an adult female was caught
at a hilltop rock ledge. On October 4 she was caught only 40 feet
from the first location. In the period September 30 to October 17,
seven newborn young were caught within 50 feet of the female's
second location. One young was caught at the same place on
September 30 and October 7; two others each moved approximately
10 feet between October 7 and October 13, and another moved
approximately 10 feet between October 13 and October 17. These
young were obviously litter mates. For each only a "prebutton"
was recorded in the nine captures made up to October 13. On the
night of October 15 one underwent its first molt while confined in
a sack, and the one recaptured on October 17 also had molted.
In mid-October, an estimated three weeks after their birth, the'e
young had scarcely begun to disperse and they were probably still
near their birth place when they hibernated.

A half-grown female was caught on October 14 and October 26.
1952, at the same place on a hilltop rock outcrop. Tliese several
recaptures near the original location, after short intervals show little
regarding extent of home range, but emphasize the sluggish nature
of rattlesnakes, which ordinarily hunt by awaiting the approach
of their prey in strategic locations. After approximately 3^2 years
an adult female was recaptured on the same ledge 280 feet from the
original location. This is the only instance of a rattlesnake re-
captured near the same place after a long interval, and it seems

Home Ranges of Vertebrates 129

clear that these snakes wander widely. A large adult male marked
in the autumn of 1948 was reported killed in a hay field more than
half a mile away in the early summer of 1950. It was recognized
by the wire marker attached to one rattle. On June 2, 1953, a
large adult male was marked in a hilltop woods, and on October
16, 1954, it was recaptured at a ledge 2100 feet away. Another
large male was marked in November, 1950, and almost five years
later its recently cast skin was found 2940 feet from the original
location. The fact that these individuals made movements among
the longest for any snakes of the hundreds recaptured over a nine-
year period, and that none of the 26 other rattlesnakes marked was
ever recaptured after intervals of more than a few days, indicate
that timber rattlesnakes travel extensively. The wandering tendency
probably has been an important factor in reducing the population
of rattlesnakes locally and throughout much of the range. Accord-
ing to written and verbal testimony of longtime residents in Douglas
County, rattlesnakes were abundant on the area that is now the
Reservation in the late eighteen hundreds, and even up into the
nineteen twenties. Under modem conditions, Mdth heavy motor
traffic on county roads, each rattlesnake in its wanderings is liable
sooner or later to be run over and killed. Copperheads, being less
vagile, tend to stay on the rough wooded slopes where habitat
conditions are most favorable and therefore remain common locally,
holding their own against encroaching civilization even within the
city limits of Kansas City and other urban communities.

Mammals

Didelphis marsupialis Linnaeus

Opossum

Status. — Common resident.

Habitat. — Primarily woodland, woodland edge, and along mar-
gins of pond and creeks; also, to lesser extent, utilizes open fields.

Movements. — Home range and movements of the opossum on
the Reservation have already been discussed in a recent publication
(Fitch and Sandidge, 1953). To summarize the findings of this
study: Individual opossums have home ranges of irregular size
and shape over which the animal roams more or less at random in
the course of its nocturnal prowling. Home ranges tend to be
circular or oval, with an average radius somewhere in the neigh-
borhood of 800 feet. There is no territoriality and several or many
individuals may use the same area simultaneously. Two or perhaps

ISO University of Kansas Publs., Mus. Nat. Hist.

more may even use the same den. The animal may shift frequently
from one den to another. Home range boundaries are fluid, and
the ranges are constantly altered, partly in opportunistic exploitation
of new food sources that become available, and also perhaps as a
result of innate wandering tendency. The tendency to wander and
shift the range is much more apparent in the male, and normally
results in complete replacement of the individuals on any small
area within a year. After attaining independent status, young males
are especially likely to shift to new areas, while young females are
likely to remain on or near the area occupied by the mother. In
actual size of range no difference is discernible between males and
females or between old adults and yearlings.

Blarina brevicauda (Say)
Short-tailed Shrew

Status. — One of commoner small mammals of Reservation.

Habitat. — ^This shrew occupies most of the area, occurring in a
variety of habitat subdivisions. Woodland is its main habitat. It
has been taken most often in wire funnel-traps set for reptiles along
hilltop rock ledges. Often it has been taken in rainy weather in
mouse traps set in brome grass fields, especially near woodland
edge. Also it has been taken at the quarry rock ledge, the rock
pile near the pond, and in brush along fence rows at field edges.

Movements. — Although shrews are fairly common on the Reser-
vation, the information regarding their movements remains meager
because of the lack of a satisfactory method of catching tliem alive
in good condition. All captures were made in traps set for rodents,
reptiles or amphibians. Because the shrews were always caught
in relatively small numbers, it was not practical to run trap lines
for them specifically. The shrews, being much more delicate than
the other animals trapped, were subject to heavy mortality while in
the traps, and their numbers would have been decimated on the
study areas but for the fact that they entered the traps so seldom.
The box traps set for small mammals were normally tended at 24
hour intervals, but shrews caught usually exhausted the food supply
and starved within this time. Pitfalls and funnel traps set for am-
phibians and reptiles were checked less frequently since these ani-
mals were not harmed by going unfed for periods of days. The
shrews caught by chance in traps of these types usually died before
they were found.

Home Ranges of Vertebrates 131

When grids of one hundred or more mouse traps were maintained,
capture of one or more short-tailed shrews could be expected on
days when there was heavy rainfall. Such captures were made
both by day and by night. Occasional captures were made on days
following a heavy rain, when the ground was still soaked, but in
most instances the shrew was caught while rain was still falling.
Similarly, in the pitfalls and funnel traps, which were unbaited
except with insects and other small animals that happened to wan-
der into them, captures of shrews were almost invariably made in
heavy rains. During dry weather the same trap lines might be
maintained for weeks, or even months, without a single capture of
a short-tailed shrew. The implication is that, ordinarily, these
shrews venture forth on the soil surface only in wet weather, and
spend the remainder of their time under shelter in burrows and
runways. When there is snow on the ground, they are perhaps more
inclined to wander than at other times. On several occasions their
tracks have been noticed in snow on the frozen surface of the pond,
extending for 100 feet or more away from cover, in a meandering
course.

Although shrews dig burrows and make runways of their own,
the tunnels and runways of voles seem to play an important part
in their ecology. On various occasions the tracks of the shrews or
the animals themselves have been seen entering burrows of voles.

Whether shrews are to any extent territorial has not been de-
termined, but they are quarrelsome and pugnacious. On various
occasions their high, trilling squeaks have been heard in dense
vegetation, but usually the sound ceased abruptly when attempt
was made to locate the source. On November 3, 1951, a shrew was
heard squeaking in a thicket, and then was glimpsed thrashing and
darting about among dead leaves on the ground, but its supposed
antagonist was not seen. After a few seconds the shrew darted
into a burrow entrance which appeared to be that of a pine vole.

Although the shrew is known to be an important predator on
voles, it may not be able to cope with an adult pine vole. On April
18, 1951, while checking a trap line, I heard the shrill squeaking of
a shrew and traced it to a live-trap some 50 feet away. The shrew
was in the trap huddled against the door. A large pine vole was
also in the trap, and it was standing menacingly, with its head held
high, at the entrance to the nest box. From time to time the vole
would dart forward and strike the shrew a sharp body blow, forcing
it back, and eliciting the protesting squeaks.

132 University of Kansas Publs., Mus. Nat. Hist,

Only 14 of the short-tailed shrews marked were recaptured. Two
were each caught five times, two others were each caught four
times, one was caught three times and the remaining nine were each
recaptured just once. Records of these are summarized below: 570
feet in 18 days, 2 captures (adult female); 490 feet in 215 days, 2
captures; 340 feet in 97 days, 4 captures (twice near the same place
in December, 100 yards away in February, and had moved back to
near the original location in March); 180 feet in 18 days, 4 cap-
tures (first and second capture sites were 150 feet apart, the third
was the same as the second, and the fourth was back near the start-
ing point ) ; 95 feet in 8 months ( first four captures within a 40-foot
space and a 17-day period). The other nine shrews had moved
distances of 75 feet, 60 feet, 60 feet, 32 feet, 30 feet, 22 feet, 20 feet,
0 feet, and 0 feet in periods of from one day to three months. From
the trend of these few records it appears that shrews temporarily
at least may carry on their activities in small areas of not much
more than 100 feet across, but that they may also shift their activi-
ties for distances of hundreds of feet and return.

The two longest movements probably represent shifts beyond
the limits of a home range. The remaining 12 movements probably
are all within home ranges, as indicated by return movements of
300 feet and 150 feet. These 12 movements average 110 feet, indi-
cating home ranges of .87 acres. Manville (1949:65) recorded 38
movements of short-tailed shrews, which averaged 91.5 feet (30
to 315). He found no significant difference between the sexes in
the 13 males and 15 females that he was able to sex with certainty.
Burt (1940:50) concluded, on the basis of movements made by
twelve shrews that he had marked and recaptured in Michigan that
a normal home range is probably about 50 yards in diameter and
covers about .4 acre.

Cryptotis parva (Say)
Little Short-tailed Shrew

Status. — Common resident.

Habitat. — This species inhabits grassland and open woods. Most
of the little short-tailed shrews recorded have been taken in pitfalls
as they are small enough to squeeze out of most of the live-traps
used, and are little attracted by bait. Most habitats have not been
thoroughly sampled for these shrews and pitfalls are limited to a
small part of the area. The shrews have been caught most often
along the dry, exposed ledge at the quarry, and many have been
caught at the ledge at "Skink Woods," in open grassy woodland.

Home Ranges of Vertebrates 133

Others have been found in various parts of the woodland. Several
were caught in mouse traps in open fields.

Movements. — Little short-tailed shrews seem to be somewhat less
common than Blarina on the Reservation. At any rate they are
caught in traps set for small mammals much less frequently. Be-
cause of their minute size they are sometimes able to enter and leave
such traps without tripping them, and are also able to squeeze
through small openings to escape. The majority of those caught in
pitfalls were found dead, and those that were marked and released
were never recaptured.

Captures of Cryptotis in pitfalls, like those of Blarina, were almost
invariably made in heavy rains. At other times the shrews seem to
stay beneath ground litter. One shrew caught in a mouse trap had
killed and eaten a harvest mouse larger than itself, which was in the
trap with it.

On several occasions two adults of the little short-tailed shrew,
male and female, were caught together in the same pitfall trap. In
these instances it seemed probable that the pair had been travelling
together when they were caught, although this could not be defi-
nitely determined.

Scalopus aquaticus (Linnaeus)
Eastern Mole
Status. — Common resident.

Habitat. — Most typically, woodland, but also present in parklands,
pasture, prairie, and fallow fields.

Movements. — No suitable live-trap was available for the capture
of moles; therefore nothing was learned concerning the movements
of individuals. Moles were rarely seen, and the only basis for judg-
ing their activity was provided by the mounds and ridges of earth
thrown up by them. Activity was noted to differ greatly from time
to time and from place to place. During times of drought, as during
most of 1953, and for shorter periods in other years, mole workings
were seldom seen. However, after heavy rains had softened the soil,
there was always a surge of activity and evidence of burrowing was
conspicuous. For example, on February 15, 1954, there was a rain
of 1.35 inches, and within three days many new mounds and ridges
had appeared. Groups of such workings were concentrated, each
in a small area, and each presumably originating from a different
mole. Within a 20-foot radius of a large elm in the headquarters
area, the ground was criss-crossed with a network of mole ridges.

134 Unr'ersity of Kansas Publs., Mus. Nat. Hist.

Branches extended out in several directions and one was traced for
more than 100 feet.

After prolonged drought, moles were noticeably less common
than at other times. Difficulty in digging through the hard dry soil
presumably restricted their movements and limited their food sup-
ply.

In the spring of 1957 after soaking rains, new mole ridges were
noted to be numerous along the edges of formerly cultivated fields
on the Rockefeller Tract. Several runways that were followed ex-
tended out into tlie fields for 200 feet or more in meandering courses,
as if the moles were finding some kind of subterranean insect prey.
No deep burrows or mounds were noted in the fields, and obviously
the moles were not living in the fields, but entered them temporarily
from adjacent woodland. Brumwell (1951:208) mentioned similar
exploratory tunneling in recently deposited sandbars along the Mis-
souri River, with digging concentrated about patches of smartweed,
sedge, grass, or willows where insect prey was abundant.

Lasiurus borealis (Muller)

Red Bat

Status. — Common resident in summer; evidently migrates to more
southern regions in winter.

Habitat. — Primarily woodland; often noticed flying at dusk near
woodland edge or through parklike open stands of large elms, or
near pond.

Movements. — Red bats often were observed at dusk flying over
a regular beat. Most often the bat was 30 to 50 feet above ground
and kept to a fairly straight course, in the open but in the vicinity of
trees. For example, on July 27, 1951, at twilight, one was coursing
back and forth over an open stretch about 100 feet long, between
large elms just east of the headquarters area. On other occasions,
the beat covered was somewhat longer and less regular.

On August 18, 1949, a red bat was found hanging on a bush three
feet from the ground, at a hilltop rock ledge in thick woods.

Sylvilagus floridanus (J. A. Allen)
Eastern Cottontail
Status. — Abundant resident.

Habitat. — The cottontail uses every part of the Reservation. It
is seen frequently in both grassland and woodland. Actually it is
concentrated along the line of contact or transition from woodland
to grassland, in "edge" habitat. Blackberry thickets provide the

Home Ranges of Vertebrates 135

most preferred type of refuge, and the rabbits are especially numer-
ous where such thickets are well developed. Other types of shelter
that are used are: thorny clumps of gooseberry; rock crevices along
ledges; deserted and partly collapsed old houses of woodrats; dead
brush from crowns of fallen trees; grass clumps in open fields;
among dead leaves in woodland. Under cover of darkness the rab-
bits forage out from these shelters and may visit relatively exposed
situations. In summer, when fields are covered with high grass,
rabbits are much less closely hmited to situations with woody vege-
tation for protection, and may shift their headquarters out into the
fields.

Movements. — From September 1954 to March 1956 Donald W.
Janes carried on a program of live-trapping and marking cotton-
tails on part of the Reservation. A total of 89 were caught and
marked; 54 were recorded repeatedly and yielded data permitting
computation of home range areas. These cottontails were equipped
with colored markers attached to their ears, serving for sight iden-
tification, and many were trailed from the stain left in the snow
after the feet had been soaked in Hquid dye of a bright color. The
data were hence far more complete than they would have been if
based on live-trapping alone. Janes computed the average home
range as 8.34 acres (2.42 to 12.62). For males the average was 8.92
acres, and for females 7.76. Janes found that when records of an
individual were suflBciently numerous to form a pattern with nine
marginal points, the area encompassed was fairly representative of
the entire home range, and was ordinarily increased little or none
by additional records. Computation of the home range area from
distances between successive (random) points of capture, assumed
to represent a home range diameter, yielded a figure of 8.40 acres,
approximately the same as the 8.34 acres obtained from actual
measurements. In several cottontails Janes noted increase of the
home range area in summer five to fifteen per cent beyond the size
of the area covered in Mdnter.

Young cottontails that are old enough to forage increase their
ranges gradually. For one female Janes recorded the following:
age three weeks (July 1954), range approximately .25 acre; age 11
weeks, range 1.5 acres; age 23 weeks, range 8 acres; age 75 weeks,
range 11 acres.

In Michigan Haugen (1942:366) found that adult females oc-
cupied an average home range of 14.0 acres during the winter, and
that the size increased to 22.5 acres during the breeding season. He
found that adult males wandered much more extensively.

136 UxrvERSiTY OF Kansas Publs., Mus. Nat. Hist.

Spermophilus franklinii (Sabine)
Franklin's Ground Squirrel
Status. — Rare wanderer.

Habitat. — Situations with tall grass or other rank herbaceous
vegetation are preferred.

Movements. — On June 27, 1958, Robert L. Packard saw one on
the Rockefeller Tract east of the buildings in a field having low
grass and weedy vegetation. It escaped into woodland and could
not be found again on subsequent days, when search was made.

Marmota monax (Linnaeus)
Woodchuck

Status. — Uncommon transient.

Habitat. — Woodchucks have been observed at well-scattered
points, both in woodland and in grassland. In foraging, the species
prefers situations with lush grass, at edges of fields near thickets of
woodland edge or fence rows. It has not been seen in eroded field
areas having scant weedy vegetation.

Movements. — Over the nine-year period covered by my work on
the Reservation woodchucks were noted appro.ximately a dozen
times. However, no two records were in the same place, and
none was asrociated with a burrow. In each instance the animal
was a wanderer. No permanently occupied burrows were known
nearer than a mile to the Reservation. The woodchucks noted on
the area were all adults and were mostly in brushy and wooded
situations. One found on April 18, 1951, was in a small elm tree
20 feet above the ground. It remained "frozen" motionless as
persons walked beneath it and tossed sticks at it. On June 13, 1951,
one was seen three times at about the same place on the road
between an alfalfa field and a brushy hillside.

Sciunis carolinensis Gmelin

Gray Squirrel
Status. — Moderately common resident.

Habitat. — Dense woodland of mesic aspect, typically oak-hickory,
where trees are large and have a thick leaf canopy.

Movements. — Packard (1956: 29) presented figures showing the
average movements of the squirrels marked by him and recap-
tured, both on the Reservation and on the Dillon Farm some
eleven miles south, classified according to sex and age groups.

Home Ranges of Vertebrates 137

Packard kindly made available his original records for the Reser-
vation, For 26 intervals between successive captures in gray
squirrels on the Reservation the average distance was 513 feet.
However, it is questionable whether all movements recorded were
within the home range of the animals. Of four movements of
more than 1000 feet recorded (1350, 1250, 1110, 1110), each of
three was made in a single day, supporting the supposition that it
was within the individual's normal range. For ten movements of
males the average of 626 feet was higher than the average of 472
feet for the 15 movements of females.

Packard (op. cit. :117) found a population of nine gray squirrels
and 15 fox squirrels on a 27.1-acre tract of woodland where his
study was concentrated. No interspecific strife was observed.
Obviously the gray squirrel does not maintain large individual
territories, although the nest and its surroundings are defended
against other individuals.

Sciurus niger Linnaeus
Fox Squirrel

Status. — Abundant resident throughout wooded portions of area.

Habitat. — Woodlands of all types are utilized, most notably
where there are large trees of oak and hickory, and especially wal-
nut. These squirrels sometimes live in parklike situations where
trees are well scattered. Even isolated lone trees that are produc-
tive of food are visited regularly. Some such trees may be 100
yards or more from the edge of woodland. Extensive treeless
areas are avoided.

Movements. — Packard (op. cit. -.28) presented figures concerning
the average movements of fox squirrels and gray squirrels trapped
by him on the Reservation. These figures are well adapted to show
diflFerences between the species, and within a species according
to age and sex. However, they give little idea of the normal move-
ments of an individual, because diflFerent kinds of movements are
combined. Packard has kindly permitted me to examine and re-
arrange his original data. Fifty-five movements are rather evenly
distributed for distances up to 1000 feet. Ten longer movements
(1600 and 1600 feet for adult males; 3400, 2175, 1550 and 1050 feet
for adult females; 2850, 2175, 2050 and 1450 feet for young males )
seem to represent shifts of range, in most instances at least, and
they are therefore excluded from computations of home range. For
the 55 movements of less than 1000 feet, the average is 436 feet.

138 University of Kansas Publs., Mus. Nat. Hist.

representing a home range 13.7 acres in area. This figure may be
representative for the species in a woodland habitat. Significant
differences in the average movement were evident according to
age and sex, as follows: 22 adult males, 450 feet (14.5 acres); 14
adult females, 364 feet ( 9.5 acres ) ; 11 young males, 500 feet ( 18.0
acres); 8 young females, 439 feet (13.7 acres).

Geomys bursarius (Shaw)
Plains Pocket Gopher

Status. — Occasional on the Reservation; abundant on adjacent
cultivated land.

Habitat. — Locally this gopher is found mainly in hay fields, espe-
cially those that are used for growing alfalfa. The population of
an extensive agricultural area of the Kansas River flood plain was
largely eliminated in the flood of July, 1951, and the area has been
re-occupied gradually. Gently sloping land along the edges of
the valley, like the farm lands adjoining the Reservation on the
west and south, provided retreat where a population was able to
survive the flood. Individual gophers have shifted onto the Res-
ervation from time to time. Their workings have been found in
bottomland of each of the two small valleys, in brome grass fields.

Movements. — When field work was begun on the Reservation in
1948, the whole area was searched thoroughly for sign of gopher,
but none was found. Subsequently, on several occasions gopher
mounds appeared in situations remote from any known colonies, and
the individual (or individuals) involved were knowii to have re-
mained in the same small area for periods of months. Isolated
individuals, such as those that appeared on the Reservation, offered
unusual opportunity for determining the extent of a burrow system
and the changes that occurred in it with time. Ordinarily, where
gophers occur the burrow systems of individuals overlap and inter-
communicate so extensively that the limits of any one system
cannot readily be determined.

On the Reservation gopher mounds were found first in September,
1948. Several were noticed on a gentle slope in the pasture a short
distance north of the headquarters area. On October 12, a week
after a heavy rain, there were many more mounds at this place;
22 fresh ones were counted, all within a space of 40 feet. On Octo-
ber 25 there were many new mounds within the same area. The

Home Ranges of Vertebrates 139

most remote were 50 feet apart, with freshly deposited mounds
at each edge and at several intermediate locations. By this time
the loose earth of the mounds covered a substantial portion of the
area encompassed by the burrow system. By November 2, a linear
series of mounds extended out from the former limits of the system,
up the gentle slope some 50 feet to a ditch bank. For the next 17
months this burrow system was continuously occupied, and new
mounds were tlirown up from time to time, all either within the
limits of the area occupied in September and October, 1948, or
within a few yards of its edges. This burrow system was in a
former pasture dominated by brome grass, 1450 feet from the near-
est boundary of the reservation and hundreds of yards farther
from any area that was permanently occupied by gophers. In early
October, 1949, a second group of mounds was noticed in the same
field 950 feet farther east southeast. The extent of this burrow
system was checked from time to time and in the next three months
it was steadily extended north northwest 160 feet up a gentle slope
to the edge of woodland. From the trend of the lines of mounds,
it was evident that several times in the course of extending its
burrows up the slope, the gopher had divided its efiForts between
two more or less parallel or diverging branches.

In the summer of 1953 a group of large mounds was noticed
beneath a locust grove in the large pasture in the southeastern part
of the Reservation. This burrow system may have been present
for a long time before it was noticed. On December 21, 1953, most
of the mounds were within a span of 40 feet, but several fresh
mounds were more than 100 feet from the main group. Soil was
deep and rich at this place, and the mounds were exceptionally
large; some were estimated to contain as much as 100 pounds of
earth. Throughout the winter, the burrow system was almost co-
extensive with a grove of locusts and osage orange, about two
dozen trees in all, covering an area approximately 150 by 100 feet.
For most of the next winter the burrow system was limited to the
same area, but by April 1 a linear series of 35 new mounds extended
the system for 100 yards east southeast, in an almost straight line.
This temporary extension of the burrow system was probably soon
abandoned, but activity continued in the burrow at least into the
spring of 1958, indicating more than 4/2 years of continuous occu-
pancy.

In each instance when a gopher appeared on the Reservation,

140 University of Kansas Publs., Mus. Nat. Hist.

the animal probably had wandered on the surface for hundreds of
yards, from an established colony. An immature male, probably
a wanderer, was found dead on the road near the Reservation
headquarters. It appeared to have been attacked by a predator.

Reitlirodontomys montanus (Baird)

Plains Harvest Mouse

Status. — Uncommon resident.

Habitat. — This mouse is confined to grassland, especially where
the grass is short. The habitat requirements are more restrictive
than those of the western harvest mouse, which is common and
widely distributed on the area. One Plains harvest mouse was
caught at the headquarters, where grass had been cut several times
annually. Others were caught in the large hilltop field where
prairie grasses had been restored, but their records were concen-
trated at a few trap stations where grass was short. In the spring
of 1957 a colony was discovered in the blue-grass pasture on the
Rockefeller Tract. A few weeks later, when this pasture, no longer
grazed, had become overgrovm with high weeds, none could be
caught in live-traps.

Movements. — Twenty-five of the Plains harvest mice recorded
were recaptured only at the site of the original record. One of these
was recaptured four tim.es; three, three times; five, twice; the others
only once each. Twenty-nine movements were recorded, and aver-
aged 83 feet, a figure remarkably close to that obtained for the west-
em harvest mouse, with more than twenty times as many records.
Therefore, I conclude that in this species also, the home range aver-
ages approximately half an acre.

Most captures were made on the Rockefeller Tract in November
and December 1957, especially in the more barren parts of the pas-
ture, where vegetation was chiefly lespedeza and three-awn grass.
Because of minute size and delicate touch, these mice were often
able to enter the traps without springing them, and after traps had
been set for periods of days, individuals or pairs might take over
as residences the nest boxes attached to some of the traps. The har-
vest mice released from live-traps were easily followed in their open
habitat, but attempts to trail them to a hole or nest were invariably
unsuccessful. The mouse followed wandered slowly and furtively
in an irregular course, often stopping to crouch in a depression at-
tempting to conceal itself, but never entering a burrow.

Home Ranges of Vertebrates 141

Reithrodontomys megalotis (Baird)
Western Harvest Mouse

Status. — Abundant resident.

Habitat. — Grassland of various types.

Especially high concentrations were present along the weedy mar-
gins of the pond where there were dense mats of rice cutgrass
(Leersia onjzoides) mixed with other grasses and forbs; and in tall-
grass prairie. In the latter there were more harvest mice in mixed
stands and those of switch grass, Indian grass and side-oats grama
than there were in stands of little bluestem or big bluestem. In
fallow fields, patches of foxtail grass supported colonies. When the
Reservation was first established, in 1948, the pastured areas, con-
sisting of brome grass with a mixture of many weedy species, were
closely grazed, and did not provide suitable habitat for harvest mice.
In subsequent years, after discontinuance of grazing, the rank vege-
tation provided a favorable habitat, and by late 1949 all these for-
merly pastured areas were well populated with harvest mice. The
mice continued to thrive in 1950 and 1951, but subsequently as var-
ious kinds of herbaceous plants were crowded out, and succession
progressed toward a pure stand of brome, the numbers of harvest
mice dwindled to a low level.

Movements. — Harvest mice were trapped in large numbers in
several habitats, and a large amount of data was obtained concern-
ing their movements from day to day and over longer periods. In
the months of June, July, August and September harvest mice were
trapped in much smaller numbers than at other times of year. They
seemed to be less attracted to bait in summer, probably because of
the abundance of preferred natural foods. The June-to-September
period therefore constitutes a gap in the records of many individuals
that were marked and recaptured over periods of months.

Harvest mice that were released from traps most often hid in
dense ground vegetation, but sometimes they moved oflF in unhurried
fashion, seemingly unconcerned by the presence of an observer.
Occasionally such a mouse was followed for 100 feet or more, but
none was ever followed to its nest. Such mice progressed through
tall grass by alternately walking and making long bounds, gripping
the stems in all four feet, and more often cHmbing than walking on
the ground surface.

One harvest mouse was trapped 16 times, and many others were
caught five times or more, permitting some idea of the extent of

142 University of Kansas Publs., Mus. Nat. Hist.

their normal movements. For one the span of records covered 20
months, and for a dozen others the records covered a year or more.
No movements of hundreds of yards, from one study area to an-
other, were recorded. It seemed that each individual was restricted
to a limited area and there was but little tendency to wander. For
indixiduals that were caught many times there was a dispropor-
tionately large number of captures at one or two stations and other
captures were more scattered at several other sites. Outlying sta-
tions were most often represented by just one capture each.

For 106 harvest mice each caught at four or more locations, the
"minimum home ranges" (areas encompassed by the capture sites)
averaged .29 acres. For the 30 that were each captured at four lo-
cations, the average area was .16 acres; for 26 each captured at five
locations the average area was .29 acres, and for those captured at
seven locations, .24 acres. For 18 each caught at more than seven
locations, the average area was .53 acres. Corroborative evidence
of home ranges approximately half an acre in extent was obtained
from many other individuals recaptured too few times to permit
plotting their separate ranges. For 327 intervals between successive
captures in males, the distance averaged 85 feet, indicating home
ranges of approximately .52 acres. For 261 intervals between suc-
cessive captures in females, the distance averaged 84 feet, indicating
that there is no significant difference betAveen the sexes. In each in-
stance those movements of more than 300 feet (5.5 per cent in
males and 5.1 per cent in females) were arbitrarily eliminated from
the computation, as such long movements were considered to rep-
resent shifts in home range or wandering beyond the usual bound-
aries. The relatively few movements recorded in sinnmer did not
differ in their trend from the year-around movements, and it seems
that the same home range may have been maintained tliroughout
the year.

The extent of movement does not seem to differ much in males
and females. For 37 females each caught at four or more locations
the "minimum home ranges" plotted averaged .26 acres, while for
67 males the comparable figure was .29.

On many occasions nests of harvest mice have been found,
mostly on the surface of the soil in dense vegetation. Some were
partly underground in shallow depressions or in cracks of eroded
banks. Others were fixed in plant stems above the ground surface,
and one was three feet above ground in a dense osage orange
thicket. The harvest mice found in such nests were mostly litters
of young, but pairs of adults also have been found occupying nests.

Home Ranges of Vertebrates 143

The nest found above ground in the osage orange was occupied by
five harvest mice that seemed to be of adult size but may have all
belonged to the same litter. On many other occasions, in walking
through grass, I have flushed harvest mice without being able to
find a nest. Individuals may spend a good deal of time away from
their nests resting or feeding in sheltered places. Each individual
probably has a nest to which it returns regularly. Those kept in
captivity always made nests of cotton, shredded grass, or any other
available material.

Live-trapping records seem to indicate complete lack of terri-
toriality in harvest mice. The home ranges plotted show extensive
overlap with several or many adults of both sexes using the same
area. Frequently more than one harvest mouse was caught in a
trap. Most often the mice caught together consisted of pairs, adult
male and female. In many instances the females of such pairs were
not in breeding condition. Therefore it seems that the members of
a pair live together more or less permanently in the same nest and
forage together. Not all multiple captures involved a pair; some
consisted of a female and her well-grown young, and other captures
were of two adults of the same sex, either male or female. On many
occasions mixed groups of adults have been caged together without
any fighting, and they were generally tolerant of other individuals.

Peromyscus maniculatus (Wagner)
Deer Mouse

Status. — Moderately common resident in more open situations.

Habitat. — Grassland, especially where grass cover is low and
sparse because of heavy grazing, or erosion, or poor soil, or combina-
tions of these factors and others.

As compared with the white-footed mouse this species occurs in
much lower populations on the area. Habitat separation between
the two species is almost complete; overlapping occurs only in
grossly disturbed situations where vegetation is in an early serai
stage: margin of the pond, talus at base of rock quarry, and along
cut banks of gullies. Most captures of deer mice were made in a
field which had been stripped of sod, eroded and covered with a
thin stand of lespedeza and poverty grass, which was gradually
being replaced by brome grass. None was ever caught in the
woods.

Movements. — Deer mice were persistent repeaters where live-
traps were available to them. One was caught 43 times, and eleven

144 University of Kansas Publs., Mus. Nat. Hist.

were caught 12 times or more. Those individuals caught frequently
were usually taken time after time at one or two stations, and were
caught much less frequently at outlying stations. The female caught
43 times was taken 12 times in 6 different months at the most
favored station, and 5 times in 3 different months at a second sta-
tion 75 feet from the first. She was taken 4 times at another station
and 3 times at each of two others and twice at one station. At each
of the remaining fourteen stations she was caught only once.

One individual was caught seven times in succession at the same
station, and was not caught elsewhere, although the station was
within a 50-foot trap grid. Another individual was caught eight
times in succession at the same place, but subsequently it made
shifts of 50, 100, 50 and 70 feet. Of the individuals recaptured, 52
had too few records to show extent of home ranges. For these,
however, 149 recaptures do show something of the average trend
of movements. In 43.3 per cent of the recaptures the animal was at
the same station as at its preceding capture; in 18.8 per cent the
animal had moved from 10 to 50 feet; in 20.1 per cent it had moved
from 51 to 100 feet; in 12.4 per cent it had moved from 101 to 200
feet, and in 5.4 per cent it had moved more than 200 feet. The
longest movement recorded was 525 feet, and other notably long
movements were: 500, 350, 310, 300, and 275 feet.

"Minimum home ranges" were plotted, by drawing a line around
all capture stations ( except those involving obvious shifts ) for 18 in-
dividuals. The average area was .74 acres, but with a wide range,
from 2.24 to .27 acres. The two with only four capture stations
had the smallest areas, .27 and .28 acres; the eight with six or seven
stations each had areas from .34 to .50 acres ( average .39 ) , and those
with 8 or more stations ranged from .53 to 2.24 acres, averaging .96.
Only the one individual covered more than an acre. If this one
notably large area is excluded, the average for the seven others
caught eight or more times is .78 acres. Corroborative evidence con-
cerning size of home range is provided by a total of 84 intervals be-
tween successive points of capture. These intervals ( excluding two
that were exceptionally long and were thought to represent shifts
in range) averaged 107 feet, representing areas of .8 acres. The
ranges of males are, on the average, larger than those of females;
for 43 movements of males the average was 120 feet, indicating a
home range of one acre, while for 41 movements of females the aver-
age was 92 feet indicating a home range of .6 acres.

On many occasions deer mice released from live-traps were
trailed to hiding places which may or may not have been their

Home Ranges of Vertebrates 145

regular home bases. Several times individuals trapped near the
edge of a road ran rapidly along the open area of the wheel track
for distances up to 200 feet, turning off sharply into tliick grass in
a manner suggesting that they were following routes familiar to
them and frequently travelled. Released individuals, unless startled
by sudden movements of the observer, usually moved slowly and
cautiously slinking through dense vegetation, and occasionally cross-
ing open areas with a series of brisk bounds.

Deer mice were caught at all seasons, but records tended to be
concentrated in cold weather, November dirough March. Pre-
sumably preferred natural foods available in quantity during the
warmer months caused the mice to be less tempted then by trap
bait. Seasonal change in food supply and cover may result in
small scale shifts in range. One adult female, caught four times
within a 75-foot radius in February and March, was caught in May
at a station 1000 feet from her original range. This was by far the
longest movement recorded. In several other instances much
shorter movements were recorded to stations that seemed to be out-
side the main home range of the animal involved.

Blair (1942:29) estimated that home ranges of adult male prairie
deer mice in Michigan average .77 acre and those of adult females
.63 acre. He found that the woodland deer mouse (subspecies
gracilis) in northern Michigan has a home range considerably larger
than that of the prairie subspecies. He calculated average home
ranges of 2.31 ±: .27 acre for adult males and 1.39 ±: .16 acre for
adult females. Manville (1949: 14) reported home ranges for P. m.
gracilis in northern Michigan, of .11 acre to .31 acre in adult males
and .12 acre to .25 acre in adult females. Manville's calculations
were based on a grid of live-traps spaced at ten yard intervals, and
he considered each trap to cover effectively and to represent the
100-square yard area nearest it. Trap stations where an animal
was not recorded were not considered to be within the home range
and their areas were not included even though they were sur-
rounded by other capture sites. Blair's calculations included such
areas and therefore the home ranges indicated were larger. Never-
theless, the possibility was mentioned by Blair, and by Dice and
Howard (1951:6) that the method used would result in too small
an estimate of home-range area, because each individual tends to
be trapped each night before he has travelled from his homesite
to the outer limits of his range. Dice and Howard presented many
instances of deer mice that, in the course of their normal activities,
moved distances considerably greater than would be expected if

146 University of Kansas Publs., Mus. Nat. Hist.

their movements were confined to a 2.29 acre area (the maximum
home range area recorded by Blair ) . Dice and Howard found that
dispersal occurs in many young deer mice when litters are nearing
maturity, with a mean dispersal distance of 188 feet for females and
339 feet for males.

Peromyscus leucopus (Rafinesque)
White-footed Mouse

Status. — Abundant resident throughout woodlands and woodland
edge situations of the Reservation.

Habitat. — Though present in all types of woodland this mouse
occurs in varying populations depending on availability of shelter,
and on the food, especially at critical seasons. Relatively few have
been taken in deep woods having a thick canopy. In open woods
having a ground layer of grass and other herbaceous vegetation, the
mouse tends to be more numerous and probably attains highest
populations in woodland edge habitat, and in parkland situations.
Coralberry bushes, bearing abundant crops of berries, where tliey
are not too heavily shaded, provide one of the main food sources.
Some favorite types of shelter used as nesting sites are: holes and
crevices in hilltop rock ledges; spaces beneath loose flat rocks;
hollows of dead logs, stumps and snags; stick heaps of deserted
woodrat houses and low squirrel houses; and root tangles of over-
hanging banks. This mouse usually does not venture more than a
few yards out into open fields, but has been trapped, hundreds of
feet from woodland, along brushy fence rows, and in patches of
sumac ( Rhus glabra ) .

Movements. — White-footed mice were caught in almost every
kind of trap line that was maintained on the Reservation. In
autumn they were often caught in wire funnel traps set for snakes
along hilltop rock ledges. The traps set for reptiles were not
checked daily and the incidence of mortality was high among the
mice trapped. Traps set in trees and bushes, for birds, at times
when ground was snow-covered, caught white-footed mice con-
sistently if left overnight. Lines of mouse traps baited with grain,
caught white-footed mice in a variety of habitat situations, but
these mice were rarely taken in open fields more than a few yards
from the edges.

In extremely cold weather these mice rarely were caught. Sud-
den warming after such cold spells may induce the mice to emerge
and forage in daylight, providing unusually favorable opportunity
for observing them. On the afternoon of January 3, 1952, after two

Home Ranges of Vertebrates 147

days and nights of temperatures mostly below 15° F., weather had
warmed to slightly more than 40° F., the sun was shining and the
two-foot snow was melting. Several sets of recent tracks of the mice
were noted. Each set followed a meandering and aimless course.
One set was followed to a distance of more than 100 feet from the
starting place, the mouse having travelled several times as far in
actual distance. Another set led to a clump of coralberry bushes
where the mouse itself was found, feeding on the seeds two feet
above the surface of the snow, which was littered with tlie dis-
carded hulls of the berries. The mouse dropped onto the snow
when I approached, but after moving only a few inches, it resumed
its search for food, burrowing in the snow to reach a honey locust
pod on which it began to feed. In snow, trails are often short and
direct, leading from one shelter to another, sometimes only a few
yards apart. After snow has been on the ground for several days,
such a route may become worn to a well-beaten trail with numerous
sets of tracks superimposed. White-footed mice released from
live-traps often returned to shelter by an indirect but convenient
and presumably familiar route. For example, one followed be-
neath one edge of a log, turned sharply to run up an inclined stick,
through a bush along horizontal branches, and then passed into
the side of a rotten stump. On December 23, 1953, I noticed
tracks of a mouse in snow on a wooded slope. The mouse had
made many trips of approximately 100 feet up and down the
slope, tending to follow the same route, but often deviating from it
for part of the distance, so that there was one main trail with sev-
eral parallel to it, frequently merging and separating.

Each mouse maintains a home base, a shelter where it is in-
sulated from extremes of weather and relatively safe from predators.
Dry leaves, straw and fur may be brought to the nest cavity.
Unless there are dependent young, material is scanty in the summer,
but may be abundant in cold weather. The mouse may use several
such shelters, alternating between them. In late July, 1951, mice
of this species were found in unusually large numbers beneath
large flat rocks on wooded slopes. Excessive rainfall in the pre-
ceding month probably had saturated many of the nests in bur-
rows, rotten logs and stumps, rendering them untenable. On many
occasions, females with young have been found in nests under flat
rocks. Pairs of adults have been found together in nests at all times
of year, and have often been caught together in traps of various
types.

Territoriality seems to be only weakly developed; several or many

148 Unr'ersity of Kansas Publs., Mus. Nat. Hist.

white-footed mice inhabit the same area. A trap set beside a
deserted woodrat house caught five adult mice simultaneously.
Probably all were occupants of the rat house.

Of the white-footed mice live-trapped and marked, 249 were
recaptured one or more times providing information concerning
movements. Each of 16 mice was caught a dozen times or more,
over periods of up to 15 months. The longest distances recorded
were, in feet: 1760, 850, 660, 620, 600, 580, 520, 480, 440, 420, 380,
330, 330, 330, 330, 320, 320, 305, 300. These few relatively long
movements involved only 2.1 per cent of the total. Even after
periods of months any individual mouse was usually caught within
the same small area. Numbers of records representing various
distances were as follows: 0-20 feet, 449; 21-40 feet, 139; 41-60 feet,
77; 61-80 feet, 46; 81-100 feet, 46; 101-120 feet, 24; 121-140 feet, 26;
141-160 feet, 17; 161-180 feet, 12; 181-200 feet, 12; 201-220 feet, 7;
221-240 feet, 5; 241-260 feet, 6; 261-280 feet, 5; 281-300 feet, 3.

Twenty individuals each caught at six or more different places
provided information concerning size of home range. Minimum
home ranges were plotted by drawing lines on a map between
capture points and measuring with a planimeter the enclosed area.
The areas ranged in size from .03 up to .36 acre (average .16 acre).
Better information concerning the usual extent of home range is
available from the 637 intervals between successive points of
capture. These intervals averaged 74.4 feet, indicating home
ranges of approximately .4 acre (.35 for 341 movements of males
and .42 for 295 movements of females).

Burt (1940: 26) found an average "minimum home range" of
.27 acre for 58 adult males and .21 acre for 65 adult females. In my
study a large adult male was caught at six places linearly arranged
within a 425-foot stretch along the edge of a gully in an open field.
The gully was fringed with trees and bushes, and presumably the
mouse kept to the gully and the adjacent thicket, within a sp.ice
only a few yards wide. On one occasion when the mouse was
released it was followed for more than 100 feet. It stopped once
beneath a flat rock where quantities of discarded seed hulls showed
that the shelter had been frequently used for feeding, and finally
took refuge in a deep vertical crack where soil of the cut bank
was sloughing away. Social wasps (Polistcs sp. ) were wintering
in large numbers in this and other cracks of the gully bank, and
quantities of their discarded remains showed that small predators,
probably Peromyscus, fed upon them frequently. In an adult

Home Ranges of Vertebrates 149

female captured at nine different places the records were also
linearly arranged and they encompassed a distance of approximately
320 feet. Other home ranges plotted tended toward triangular,
circular, or oval shapes. The data suggest that males and females
differ little or none in extent of area covered; that the individual
tends to stay in the same home range throughout the year but
makes minor seasonal adjustments according to changing avail-
ability of food and shelter. Records were obtained in greater
numbers in cold weather when bait seemed more attractive.

Sigmodon hispidus Say and Ord
Hispid Cotton Rat

Status. — Common resident at times, but subject to such drastic
population reductions that at other times it virtually disappears from
the Reservation. Probably in each instance small breeding stock
has remained, although for periods of months none was noted.

Habitat. — Grassland, in situations with rank weedy vegetation,
as at edge of pond, and near brushy margins of fields.

Movements. — Cotton rats are diurnal, but are rarely seen because
of their tendency to keep to situations where dense grass and other
vegetation provide cover. Because of their timidity and rapid
movements, they were usually only glimpsed as they darted through
the grass. Information concerning their movements was obtained
almost entirely from individuals marked and retrapped. In the
course of their feeding and trampling, cotton rats make runways
through the grass that are less well defined than the runways of the
prairie vole. In the course of their normal movements, cotton rats
seem to be much less closely confined to runways than are voles. A
cotton rat released from a live-trap, or flushed, dashes through the
grass in an almost straight course, with long bounds, often crash-
ing into grass stems and other obstacles. After it is lost to view its
course may be followed for many yards by the sound and the waving
grass stems, as it runs without seeking out a runway, making much
more rapid progress than a vole would make in its runway.

In cotton rats trap habit was often strongly developed. Each of
several individuals was caught several times in the same trap witli-
out visiting any of several nearby traps. Some individuals were
caught repeatedly at traps that were well separated, but were
caught only occasionally or not at all in traps that were more or less
intermediate in position.

Most records of cotton rats on the Reservation were obtained in

150

UisnvERSiTY OF Kansas Publs., Mus. Nat. Hist.

the summer and autumn of 1951, and in the following winter and
early spring of 1952. By the end of April, 1952, few remained.

Many of those recorded were originally caught as partly grown
young. After its initial capture a cotton rat might be recaptured
almost daily, when the trap line was operated, over periods of
weeks. The maximum number of captures recorded for any indi-
vidual was 46. Many of the rats were caught in only one, two or
three locations; each of 42 others was caught four or more times
(average 11.7). In each of these instances the records were scat-
tered and seemed to indicate the general size and position of the
home range in each instance. These records show that the home
range is small, and that its size varies widely. One adult male had
ranged over an area of 2.03 acres (8 captures in 23 days) and
another had ranged over 1.16 acres (20 captures in 43 days). An
adult female had ranged over 1.11 acres (with 46 captures in 295
days ) .

Table 1. — Average Sizes of Minimum Home Ranges in Cotton Rats,
Showing Increasing Size with Number of Marginal Points in Minimum

Home Range.

Number of marginal points. . .

Average minimum home range
Number in sample

3

4

5

6

.14

.16

.29

.72

10

20

10

6

,72
4

Minimum home ranges of 50 cotton rats were plotted. A trend
toward larger areas in those with more numerous records was evi-
dent, as shown in Table 1. Hayne (1949:14) questioned whether
the apparent levelling off in size of minimum home range, with fur-
ther captures, after a rapid increase with the first few captures, can
be assumed to show accurately the size of the actual home range.
He suggested that the tendency toward such levelling off might be
due to the mechanics of trapping instead. His suggestion is particu-
larly apropos for such small mammals as the cotton rat which
quickly acquire a trap habit and an addiction for the bait. An in-
dividual's preoccupation with the traps and bait available nearest
its "home base" or "activity center" might on most occasions prevent
it from reaching the outlying parts of its home range where it could
be caught in other traps. Disregarding this unmeasurable factor,
the following average distances between successive captures and

Home Ranges of Vertebrates 151

home ranges were indicated for the different categories of cotton
rats that I trapped.

Adult and nearly adult males ( 129 ) 95 feet ( .65 acre )

Adult and nearly adult females ( 67 ) . . 79 feet ( .45 acre )

Young males (77) 62 feet ( .26 acre)

Young females (73) 55 feet ( .22 acre)

Neotoma floridana ( Ord )
Eastern Woodrat

Status. — Woodrats occur in varying abundance, chiefly in wood-
land areas. From a peak population of several per acre in favorable
tracts of woodland in 1947, woodrats dwindled to only a few indi-
viduals on the entire 590-acre Reservation in 1953 and 1954. Ex-
tremes of weather in the winters of 1947-1948 and 1948-1949 were
probably responsible in part for this drastic reduction. By 1958
numbers had increased to a level somewhat intermediate between
the high point of 1947 and the low point of 1953.

Habitat. — Primarily woodland, but extending also into various
marginal situations. Since 1949 the reduced surviving population
has been confined mainly to hilltop rock outcrops in woodland edge.
Those outcrops with south exposures are favored. Most houses
are built where a loose boulder, a deep crevice, or a tree or large
shrub along the outcrop provides support and additional protection.
Projecting spurs of the hilltop where the outcropping ledge of ex-
posed limestone is especially prominent and several feet thick were
used more than parts of the hilltop where the outcropping was less
prominent. Osage orange trees were often used as house sites, the
house being built either at the base (especially in those where the
trunk forked at the ground level or near it) or in crotches a few
feet off the ground. Other favorite situations included hollow trees
and logs, blackberry thickets, exposed tree roots along cut-banks
of gullies, among dense, interlaced branches of tops of fallen trees,
in brush and weed thickets along fence rows and at the edge of
woodland. As the population dwindled, such sites were gradually
deserted, until the rats were almost confined to hilltop rock ledges.

Movements. — Each woodrat is normally the possessor of one or
more stick houses, shelters within which its nests are located. Such
a home base is the focal point for the rat's activities. The rat may
maintain two or several such shelters at well separated points, es-
pecially if it is a dominant individual, or if the population is low

152 University of Kansas Publs., Mus. Nat. Hist.

and effective competition is lacking. When an individual controls
several houses, it alternates bet^veen them, using each in succession
for periods of days or weeks. In sorties from the house, the rat
ordinarily keeps to beaten trails, and its activity is most concentrated
within a 30-foot radius. Most foraging for food is less than 75 feet
in any direction from the house. Because the woodrat usually
chooses a residence where there is dense low woody vegetation and
feeds upon such abundant materials as the foliage and bark, the
food supply within its small range is adequate. The home range
cannot be effectively measured as a tu'o-dimensional area because
of the rat's climbing tendencies. Much of the food is obtained in
trees and bushes, and travelways are often above the ground level,
typically along logs and horizontal or gently inclined branches.
Over periods of months, rats often shift into new areas, deserting
their former houses and home ranges permanently. Atlhough va-
rious factors may motivate such shifts, search for a mate is probably
the most common cause. Males wander farther and more frequently
than females, and large old males are more inclined to shift than are
younger individuals. The longest shift recorded was 1080 feet for
an adult male; 650 feet was the maximum recorded for a female.

Woodrats are intolerant of each other, and ordinarily when an in-
dividual occupies a stick house, otliers are excluded. Pairs are as-
sociated only during a brief period when the female is in oestrus.
Ordinarily an intruder is quickly routed when it ventures into an
occupied house. The house itself may be thought of as constituting
a small territory. The area about the house, in the rat's usual for-
aging range, is likewise defended. However there is no evidence
that territorial boundaries are sharply defined. Intruders may often
venture within the territory undetected since each rat spends a
major portion of its time in the nest.

Along an osage orange hedge row which offered favorable habitat
conditions, a series of twelve occupied woodrat houses were linearly
arranged at almost regular intervals averaging approximately 60 feet
( minimum 42 feet ) , but with three longer gaps which probably rep-
resented unoccupied potential nest sites (Fitch and Rainey, 1956:
522).

Microtus ochrogaster (Wagner)

Prairie Vole

Status. — Abundant resident in grassland.
Habitat. — Mainly fields dominated by brome.
In this habitat type the voles thrive best where the grass is rank
and lush; they tend to be correspondingly scarce where grass is short

Home Ranges of Vertebrates 153

and sparse. Consequently they are less numerous in hilltop fields
where soil is shallower and drier than in the bottomland fields. In
time of drought in the bottomland fields they tended to retreat to
marginal situations, such as edge of the road, and disturbed soil of
ditch banks where the vegetation was most luxuriant. Coralberry
thickets at the edges of fields provided favorable habitat. In some
instances weedy fallow fields were occupied, after several successive
crops of weeds had produced a protective cover of horizontal stems.
Voles were found in such old-field situations where there was
scarcely any grass but they were more numerous where there were
patches of foxtail grass forming a ground mat. In almost pure
stands of big bluestem and other perennial tall grasses the voles were
relatively scarce.

Movements. — The prairie vole, like other members of its genus,
makes runways through or beneath herbaceous vegetation on the
soil surface, and excavates subterranean tunnels. The activities
of an individual are largely within these surface and underground
runway systems. The pattern of surface runways especially is in
a state of constant change as older trails are abandoned and new
ones are established. Those runways leading to the most used
burrow entrances are well worn and are much more permanent than
the runways in outlying areas. The voles are somewhat social in
habits, and ordinarily a small group or colony inhabits each system
of burrows and runways. Such a group may include several adults
of both sexes, as well as young of various sizes. Well situated
burrow systems may be occupied over long periods of years by
successive generations, although individual occupancy is only a few
months at most. Unless the population is unusually low, each
colony is in contact with others, the anastomosing surface runways
merging in such a way that no boundaries can be identified. The
normal range of an individual may include two burrow systems, and
the vole may alternate frequently from one to the other. Over periods
of weeks or months a vole may gradually alter its range, withdraw-
ing from one portion and extending its activities in another until
it occupies a range entirely separate from that in which it lived
originally. More rarely, an individual venturing farther than
usual may become disoriented, losing contact with its home range,
and after a period of wandering may settle in a new location. In a
few instances, such shifts, involving distances of hundreds of feet,
were recorded in marked voles.

For the prairie vole more data have been obtained concerning
movements than for all other kinds of mammals combined. The

154

Univ'ersity of Kansas Publs., Mus. Nat. Hist.

excessive bulk of these data have presented a serious obstacle to
their analysis. Thorough analysis has therefore been postponed. It
may be expected that a detailed study of the available records will
show the extent of differences between the sexes, and differences
arising because of season, vegetation type, and age of individual
in the normal range, and the frequency of shifts. Even without
such analysis, the trends are evident.

The female vole usually leaves her young in the nest when she
emerges to forage, but occasionally, on short expeditions, she may
drag the entire litter behind her, each young clinging to a teat.
When the young are approximately nine days old, their eyes open
and thenceforth they become more active, emerging from the nest
burrow either on their own or following the female, and gradually
extending their areas of surface activity, from a radius of only a
few inches at first, as they become familiar with their surroundings.
The young may remain after they have grown to adults, continuing
to use the same burrow system and surface runways which the
female parent occupied. However, there is some tendency for
dispersal in those that have attained the age of independence. Over
periods of years many prairie voles have been caught in funnel

T^

A

'.'rc'- .

FEET

200

#•

Fig. 10. Typical home ranges of several small mammals: (A) adult male
western harvest mouse recorded seven times from November 16 1950,
to August 12, 1951; (B) adult female western harvest mouse recorded five
times from April 21, 1953, to September 10, 1953; (C) male hispid cotton
rat recorded 20 times from July 24, 1951, to September 5, 1951, both as a
juvenile and as an adult; (D) adult female prairie vole captured 52 times
from August 3, 1954, to June 30, 1955; (E) adult male pine vole captured
65 times from March 7, 1951, to March 20, 1952. Note small sizes of ranges

in the voles.

Home Ranges of Vertebrates 155

traps placed for snakes in a woodland habitat where the voles
ordinarily do not occur. Nearly all the voles so trapped were
within the weight range 20 to 30 grams, a little more than half-
grown, and they included a disproportionately high ratio of males.
As a result of extensive live-trapping on the Reservation, Martin
(1956:381) found that in both males and females the mean size of
home range was .09 acre. He found that over periods of months
the total area encompassed by a vole's movements expanded, to a
maximum of .26 acre in nine months.

Microtus pinetorum (Le Conte)
Pine Vole

Status. — Common resident, localized in small colonies where hab-
itat conditions are favorable, and subject to drastic fluctuations in
population density.

Habitat. — Chiefly parklike areas of woodland edge, where ground
is usually shaded and has dense, low herbaceous vegetation. In
the bottomland field where the house was located, a strip several
hundred yards long and 50 to 150 feet wide, at the edge of wood-
land, was occupied by the voles. This was in an area shaded by
large elms. There was a ground mat of muhly grass and bluegrass,
with patches of coralberry, and with giant ragweed and various
other coarse weeds. These voles have been taken also along hill-
top edges at scattered locations. They have never been taken
either in deep woods or in open fields more than a few yards from
trees.

Movements. — Each pine vole ordinarily stays within an estab-
lished system of runways and burrows. Territoriality seems to be
entirely lacking. A small group including several adults of both
sexes and young of various sizes usually occupies each burrow
system. Presumably such social groups live together in the same
nest chamber. Often two or even three adults are caught simul-
taneously in the same live-trap. Frequently such associations
involve two males or two females. There is complete lack of hos-
tility between such voles, both while they are in the traps, and when
they are kept in cages.

The runway systems of these voles are situated mainly along the
line of contact between woodland and pasture. The majority of
records obtained in the present study were within the 1000-foot
strip of woodland edge in the small valley where the Reservation
headquarters are located.

156 University of Kansas Publs., Mus. Nat. Hist.

Runways pass through and beneath the ground htter of dead
vegetation, usually a mixture of old leaves and remains of herba-
ceous vegetation. Runways in current use are so well roofed over
that their presence may be unsuspected. Runways may pass to
greater depths by imperceptible stages, and the runway system may
include extensive timnels three to four inches beneath the surface,
or occasionally deeper. Usually the tunnels pass through mats of
roots, making excavation difficult, either for a predator or for an
investigator. Abandoned burrows of moles (Scalopus aquaticus)
are often taken over by die voles and incorporated in a runway
system.

In general, both the underground tunnels and the runways in
surface litter consist of one main axis and numerous lateral branches,
most of which have blind ends. Occasionally, however, the sec-
ondary runways loop back to join the main tunnel, or anastomose in
a complex pattern. The runway system has no well defined limits.
Whereas the main runway in the vicinity of the nest is heavily used
and well worn, outlying ends of the main runway and numerous
branches may be faintly defined, with but little evidence of use.

Presumably the home range of each pine vole is more or less co-
extensive with the runway system it occupies. It travels mainly on
well established runways, but some of those used earlier may be
abandoned, and replaced by other extensions elsewhere as season
and food sources undergo changes. Short exploratory trips away
from the runway sytems are made frequently, and the route taken
on such a trip may be retraced until it is incorporated in the run-
way system.

Seventy per cent of die recapture records were within 30 feet of
the vole's last capture site. It seems that most of an individual
vole's foraging is confined to a 30-foot radius, and within this small
area is confined to the established runway system or its immediate
vicinity.

Pine voles were more difficult to catch than were most of the
other small mammals occurring on the Reservation. Over a seven-
year period approximately a half dozen individuals have been
caught in wire funnel traps set for reptiles along hilltop rock out-
crops. In these situations no signs of activity of the voles were
noted and the voles caught were all more than half-grown but less
than full adult size. I suspect that they were recently independent
young dispersing from the parents' runway systems and invading
unoccupied situations. Other voles live-trapped and marked and

Home Ranges of Vertebrates

157

released while they were still less than half-growii have been known
to stay in the same location and use the same nmway system until
adult size was attained and afterward. Dispersal may involve only
a small percentage of the young, or may occur only under unusual
conditions.

Although the majority of recorded movements were short, marked
pine voles occasionally were retrapped hundreds of feet from an
earlier location. In every instance these individuals were trapped
in well established runway systems. In a large proportion of such
instances the same vole was soon retrapped back at its original
location. Just why and how such relatively long movements are
occasionally made is not altogether clear. It does seem evident
that the vole remembers its route and returns by it. Runway sys-
tems that are a hundred feet or more apart and appear to be well
separated may be connected by little used outlying runways fa-
cilitating visiting between them. There is little or no difference
between the sexes in vagility. Of the 62 males and 69 females for
which movements were recorded, nine males and ten females had
made relatively long movements of 200 feet or more. Several of
them shifted repeatedly, some as many as six times. Even though
individuals may from time to time visit and explore a neighboring
runway system, the latter does not constitute a part of the regular
home range as it is not used regularly. Ordinarily, shifts in home
range are gradual, and usually they involve use of a succession of
nest burrows within an area familiar to the animal.

Table 2. — Recorded MovE^rENTs of Pine Voles, Grouped According to

Distance.

Distances

Distances

between

Percentage

between

Percentage

recorded

of total

recorded

of total

capture

movements

capture

movements

sites in feet

sites in feet

0to5

35.5

96 to 105

1.3

6 to 15

23.5

106 to 115

.3

16 to 25

11.9

116 to 125

.7

26 to 35

7.3

126 to 135

.6

36 to 45

6.2

136 to 145

.3

46 to 55

3.2

146 to 155

.5

56 to 65

2.4

156 to 165

0

66 to 75

2.2

166 to 175

.1

76 to 85

1.3

176 to 185

0

86 to 95

.5

186 to 195

.4

200 or more

1.5

158 University of Kansas Publs., Mus. Nat. Hist.

Burt (1940:51) stated that the normal home range of the species
(subspecies scalapsoicles) is about one-fourth of an acre. This
figure was based on recapture records of 17 individuals which indi-
cated an average range-diameter of 38 yards.

Ondatra zibethicus (Linnaeus)

Muskrat

Status. — A permanent population is lacking because of insuffi-
cient aquatic habitat. Nomadic individuals stray over the area from
time to time and some of these have established themselves at the
pond and resided there for periods of months when conditions re-
mained favorable.

Habitat. — Marshland is required; the shallow pond with marshy
margins, and with stands of Typha kitifolia and Tijpha angustifolia,
provides favorable habitat when the water supply is sufficient, but
on such a limited scale that only a few individuals can be supported.

Movements. — The small amount of information gathered con-
cerning muskrats on the Reservation serves to illustrate: 1. long
movements by immigrants, and 2. repeated occupancy of a small
and isolated area of favorable habitat by such wanderers.

The pond, when filled to capacity, covers approximately an acre.
On the upstream side a silt flat slopes off gradually to the pond bot-
tom, and supports a dense stand of cattails that provide food for
muskrats. In the spring of 1948 an old, deserted muskrat house
was noticed in the pond, and evidently there was at least brief oc-
cupancy in 1949 and again in 1950. In the spring of 1951 trails and
cuttings were noticed, but there were no houses and the animals
must have been living in bank burrows. High water may have
driven these rats from their burrows and exposed them to preda-
tion. In any case no more sign was seen until November, when two
adults were present. Occupancy, perhaps by these same two, con-
tinued through June 1952. At this time the pond was rapidly dry-
ing and the water was so shallow over most of it that the rats were
unable to submerge completely in swimming. In late June the
stand of cattails showed heavy use by the rats, but with lowering of
the water level this food source had become inaccessible to the rats
unless they exposed themselves crossing the open mud flats to reach
it. The rats had, in part at least, turned to a variety of herbaceous
vegetation growing on the pond dike nearer their burrows. On
July 5 the fresh half-eaten carcass of an adult female was found
on the dike a few yards from the burrows. Judging from the man-

Home Ranges of Vertebrates 159

ner in which the carcass had been attacked, a red-tailed hawk was
the most probable predator. On July 9, at 4:00 P. M. another adult
was foraging for green vegetation on the dike near the spot where
the female had been killed. It was unselective, gathering Japanese
chess {Bromus japonicus), prickly lettuce, thistle, and giant rag-
weed, and carrying them back to the burrow. It made half a dozen
trips while I watched, coming within a few feet each time without
seeming to notice my presence. On July 14 remains, probably of
this same individual, were found on the dike. The remains con-
sisted of scraps of entrails and fragments of skin, and a scat nearby
imphcated a fox or coyote as the predator.

On March 25, muskrat remains were found beside the small creek
in the east part of the Reservation. The carcass had been partly
skinned out and the bones picked, in a manner suggesting the red-
tailed hawk as the predator.

The muskrats that appear on the Reservation from time to time
are wanderers that reach the area from the Kansas River Valley.
The nearest known colony, in an old slough cut off by a shift of the
river channel, was a little more than two miles air line from the
places where the animals were found on the Reservation. In most
instances, wanderers probably follow up stream courses.

Rattus norvegicus (Berkenhout)
Norway Rat

Status. — Irregular resident, reaching the area from nearby farms.

Habitat. — Most typically the Norway rat lives in urban situations
and about barns, corncribs and occasionally along creeks adjacent
to grain fields. A quantity of com ruined in the 1951 flood was
dumped on the edge of the Reservation, and several months later
rats were found to be living in the vicinity. They may have been
transported there with the loads of corn, or may have been attracted
there from a nearby farm. They were living in rock clefts that had
been occupied by woodrats until a short time before. Also, several
Norway rats were trapped about the laboratory building approxi-
mately 900 feet from the com pile and probably twice as far from
any other colony. In this situation they were subsisting largely on
natural foods, perhaps supplemented by occasional access to the
grain used as bait.

Movements. — The Norway rats that lived beneath the laboratory
building at the Reservation headquarters restricted most of their
activities to a radius of a few yards. There were several burrow

4—3117

160 UNivTRSiri' OF Kansas Publs., Mus. Nat. Hist.

entrances beside the foundations of the building, and there were
trails leading from these burrows into nearby clumps of high weeds
and dense patches of foxtail grass (Sctaria viridis). The rats seemed
to be living on this vegetation and on waste grain. The animals of
this colony may have originated from those at the pile of waste corn
900 feet north on the hilltop, or they may have made their way up
a small creek from cornfields adjoining the Reservation to the west;
in the latter case they must have travelled at least half a mile.

Mus musculus Linnaeus
House Mouse

Status. — Common resident in vicinity of buildings, and other fa-
vorable situations.

Habitat. — Most numerous and persistently present in buildings.

Elsewhere the house mouse prefers disturbed situations with
rank weedy growth, as on silty soil at pond margin, and among
roadside weeds. It may occur in brome fields in the same habitat
occupied by the prairie vole, harvest mouse, and cotton rat. It has
been taken also in wooded hilltop rock ledge situations where the
white-footed mouse is the most common small rodent.

Movements. — House mice were not trapped with any degree of
regularity. In certain locations they were caught frequently over
periods of days and weeks. In view of the low incidence of cap-
tures, the frequency of double captures was especially high, sug-
gesting that these mice often live and travel in family groups.
Because of their agility, house mice often escaped as the investigator
opened the live-trap to remove them.

Occasionally in unusually cold weather of winter, house mice
Were found dead in a variety of situations. It seemed they had died
of exposure, while searching for food in open situations away from
shelter. With the advent of cold weather each autumn, house
mice became increasingly troublesome in a small building at the
Reservation headquarters. Often a dozen or more adults have been
trapped within a few days, after tlie population of nearby areas had
converged on this building to find shelter from the cold. On one
occasion an adult male live-trapped in the building was carried in a
box 860 feet west on the road and released. On the following day
it was re-trapped in the building, demonstrating homing ability.

Twenty-five of the house mice live-trapped, marked, and released
were subsequently recaptured a total of 36 times. Movements
ranged up to 300 feet. Two-thirds of the total were within the range

Home Ranges of Vertebrates 161

20 feet to 100 feet. Movements, 33 in all, averaged 73.6 feet. Several
of the mice were marked and recaptured on a quarter-acre area
that was intensively live-trapped with a grid of traps at 20-foot
intervals. All the movements recorded on this trap grid were
relatively short, and they lower the average. Exclusive of the
movements recorded on this quarter-acre grid, only eleven move-
ments were recorded and these averaged 143.6 feet indicating a
home range of 1.48 acres. The minimum home range of an adult
male plotted from five successive captures was measured as .44
acre.

Zapus hudsonius (Zimmermann)

Meadow Jumping Mouse

Status. — Moderately common resident.

Habitat. — Mainly high grass, such as in bluestem prairie or
brome, especially near woodland edge, where coralberry and
various weedy plants occur; also in woodland, at least along its
edges.

Jumping mice have been caught in open woods of honey locust
and elm on south facing slopes, where a ground layer of muhly
grass and low brush was present.

Movements. — In the course of live-trapping for small mammals,
jumping mice proved to be particularly elusive — so much so that
scarcely any information on movements was obtained from re-
capture of marked individuals. Most captures in mouse traps have
been made in late April and early May, soon after emergence from
hibernation. At other times the mice seem not to be interested in
bait. However, more than a dozen have been caught in unbaited
wire funnel traps set for reptiles along hilltop rock outcrops in
October. Traps set in the same situations at other times of year
have not caught jumping mice; therefore it seems that some of these
mice move to the outcrops from adjacent grassland to find places
favorable for hibernation. A young male jumping mouse trapped
on April 24, 1952, had perhaps recently emerged from hibernation.
He was re-trapped 100 feet farther south on April 29, and again 50
feet farther in the same direction on April 30. This is the only indi-
vidual that has been re-trapped.

Blair (1940:244) on the basis of live-trapping in southern Mich-
igan, arrived at a figure of .89 =h .11 acre as the average home range
of a male jumping mouse, and .92 ± .11 acre as the home range of
a female. Manville (1949:69) recorded an adult female on sue-

162 University of Kansas Publs., Mus. Nat. Hist.

cessive days in live-traps set 180 feet apart. Quimby (1951:87)
recorded an average home range of .38 acre (.19 to .87) for four
females, and .43 acre (.14 to 1.10) for five males, near Itasca, Min-
nesota. Corresponding figures for a larger series obtained near
Centerville, Minnesota, were 1.57 ± .22 of an acre for 17 females
and 2.70 rt .50 of an acre for nine males. Quimby attributed tlie
much higher figures at Centerville to the more open habitat there,
with more scattered food and cover. He concluded that the range
df an individual jumping mouse is relatively unstable.

Canis latrans Say

Coyote

Status. — Frequent transient and irregular resident.

In the summer and fall of 1948 a pair of coyotes probably denned
on the area and made it their headquarters, and from time to time
others may have lived on the area temporarily. However, it is
believed that most often the coyotes on the Reservation were merely
crossing the area.

Habitat. — The entire area of the Reservation is used by coyotes.
In daytime they tend to keep to brush and thick woods, and have
been flushed from such situations on a few occasions. While trav-
elling or hunting at night they seem to prefer grassland.

Movements. — Several travelways were present on the Reserva-
tion, One followed the middle of the long narrow hilltop field,
leading from the northeastern part of the Reservation almost to
the southwest corner of the section. A similar path crossed the
large bottomland field in the southeastern part of the area. When
the intermittent creek in the eastern part of the area was dry, the
bottom of the ravine, walled by steep banks on either side, was
used as a travelway. The coyotes that left sign on the Reservation
mostly came from rough, wooded areas in the sections to the north
in Jefferson County, and were travelling to or from the edge of the
Kansas River Valley. Of 118 scats collected on the Reservation
and analyzed, several contained remains of animals — such as sheep,
pig, domestic chicken, and jack rabbit — that must have been found
elsewhere, demonstrating the wide range of the individual coyote.
Several times coyote tracks in fresh snow were followed a mile or
more, and led beyond the boundaries of the Reservation, bearing
out the supposition that the individual animals range widely.

Packs of hounds sometimes crossed the Reservation, baying in
pursuit of quarry, which was assumed to be a coyote in most in-

Home Ranges of Vertebrates 163

stances, although raccoons, deer, and other animals were sometimes
chased. The route followed by the pursued animal could be traced
by the baying of the dogs. On one such occasion the owners were
intercepted as they were attempting to retrieve the dogs, and they
claimed that the chase had begun more than three miles northeast.

In 1948, when the newly created Reservation was still used for
pasturing livestock, tracks and scats of coyotes were noticed in
much greater numbers than at any time subsequently. Scats col-
lected in 1948 contained a high proportion of carrion from the car-
casses of cattle, whereas in later years when hvestock were no
longer present on the area, rabbits and cotton rats made up most
of the food ( Fitch and Packard, 1955 ) .

Young and Jackson (1951:59) state that the runway used by a
group of coyotes may be no more than ten miles in length, and
that it may be used throughout the life span of the individual. As-
suming such a route to be circular, and bounding the edge of the
home range, the range would have a radius of approximately 1.59
miles, and would cover 7.85 square miles.

Vulpes fulva (Desmarest)
Red Fox

Status. — Irregular resident; sign seen less frequently than that of
coyote.

Habitat. — In the winter of 1951-1952 one lived in the northwest-
ern part of the Reservation, having a den in a cleft of the rock ledge
on a southwest slope in oak woodland. Its hunting was done
mainly in the vicinity of the pond, and in the field near the house.

Movements. — The fox mentioned above was tracked to its den,
in a rock cleft along the hilltop outcrop, when snow was on the
ground. Subsequently in the winter and spring the den was
checked frequently, and it always showed evidence of recent use.
Whenever the ground was snow-covered, tracks were seen in great
abundance about the upper end of the small valley where the pond
was situated, approximately 700 feet southeast from the den. The
fox regularly ranged at least 500 feet farther in this direction, and
at least 2000 feet south from the den site. Although it probably
covered at least half a square mile, its hunting tended to be con-
centrated in a much smaller area. On March 5, 1952, following
the tracks in the snow, I found where the fox had approached a
cottontail crouching in a clump of big bluestem. The cottontail,
flushing from the clump, had dodged once, avoiding the fox's rush,

164 Unhtrsity of Kansas Publs., Mus. Nat. Hist.

and had escaped to the nearby brushy slope, with the fox in pursuit.
In the winters of 1954-55 and 1957-58 fox tracks were again
abundant in the same area, and the rock cleft den showed signs of
occupancy.

Urocyon cinereoargenteus (Schreber)
Gray Fox
Status. — Uncommon resident or transient.
Habitat. — Woodland or brushy areas.

Movements. — On April 7, 1958, Phillip Ogilvie saw one in the
northwestern acre of the Rockefeller Tract. On September 7, 1958,
one (possibly the same) was seen to cross a county road headed
east onto the Reservation 100 yards north of the entrance gate.

Procyon lotor ( Linnaeus )
Raccoon
Status. — Common resident.

Habitat. — The raccoon is primarily a woodland inhabitant and
it is notorious for its habit of foraging along edges of streams or
ponds. On the Reservation this tendency was well defined, and
sign was most frequently noticed at the pond, and along the creek
flowing through the eastern part of the area. However tracks and
other sign, and experience in trapping raccoons suggested that these
animals range throughout all parts of the area, including grass-
land, woodland, hilltops, and stream courses.

Movements. — Raccoons range over relatively large areas as com-
pared with the smaller mammals on the Reservation. Few indi-
viduals were live-trapped and the meager data available do not
show the extent of home range. Raccoons probably use every part
of the Reservation but change their activities according to seasonal
shifts in food sources. Wide expanses of open fields are generally
avoided.

In 1948 corn was cultivated on several bottomland fields of the
Reservation. Tracks and dropping of raccoons were conspicuous
along the margins of these fields adjacent to woodland. It was
evident that the animals were utilizing waste corn as a food supply.

The two small creeks on the Reservation were favorite routes of
travel for raccoons, whose tracks could be found in wet streamside
soil at all seasons. In autumn each year the pond on the Reserva-
tion either went dry entirely, or underwent partial drying, leaving
mud flats uncovered. Leopard frogs, cricket frogs, and bullfrogs
were then partly exposed to predation by raccoons, and the pond

Home Ranges of Vertebrates 165

became a focal point for raccoon activity. In late September, 1948,
the pond had dried completely and there were deep cracks in the
mud. These cracks and the dense stands of cattails provided refuge
for swarms of young leopard frogs and cricket frogs, and raccoon
tracks were especially numerous. In April, 1954, frogs and toads
had begun breeding when there was scarcely more than a puddle
of water in the bed of the pond. The water was in the center of
an area of open mud with no protective vegetation. Amphibians
therefore had no adequate escape shelter. Under these conditions
the raccoons made heavy inroads on the breeding populations of
American toads and chorus frogs. A large sycamore with a hollow
at the base was used for a den in 1953 and perhaps in other years.
Sign of raccoon was especially concentrated at the pond in autumn
at the time of drying, when swarms of newly metamorphosed frogs
and toads were leaving the water, and in spring when breeding
aggregations were present.

Sign of raccoon also was often noticed along hilltop rock out-
crops. Deep crevices in the rock provided the best den sites, as
large hollow trees were scarce on the area. On May 7, 1952, hounds
were heard baying, coming from the west probably off the Reserva-
tion. Quarry was chased in an almost straight course approximately
1/3 mile to a hilltop rock crevice where it escaped. An hour later,
investigating, I found that the dogs had done much digging but
had not been able to reach the raccoon except to tear a few tufts
of hair from its partly exposed rump. The raccoon was an immature
one.

The raccoon's tendency to shift to any readily available food
source was demonstrated on many occasions when these animals
formed the habit of following trap lines and systematically raiding
them, eating either the bait, the trapped animals, or both. Because
of these activities raccoons became a major pest to persons attempt-
ing to study populations of small mammals or reptiles. They would
follow along the lines of mouse traps, arranged in a grid with a 50-
foot interval, and would overturn each trap and eat the grain
beneath it. On a few occasions sign indicated that the voles or
other small mammals in the traps likewise were eaten. The rac-
coons showed dexterity in removing the covers of the metal nest
boxes attached to the wdre live-traps. The nest boxes were con-
structed with the metal sprung so that these covers snapped into
place, and ordinarily the operator removed them with pliers. Once
raiding had begun on a trap line, it usually continued wdth increas-
ing severity until trapping had to be abandoned. In April, 1954,
after such raids had forced abandonment of mouse trapping near

166 University of Kansas Publs., Mus. Nat. Hist.

the Reservation headquarters, the family of raccoons raided the
bait supply kept on the porch of a building, and tliey learned to
unlatch and remove the heavy metal top of a large garbage can
in which grain was stored. The raccoons also frequently raided
wire funnel traps set along hilltop outcrops, breaking open the
traps and eating the insects, reptiles or mammals caught in them.

For most of the year, at least, raccoons seem to travel in family
groups. On September 19, 1955, at 1:00 A. M. after a brief shower,
a group of four or more young raccoons moving along a nearby
dry creek bed, attracted my attention by their peculiar low trilling
calls. These calls served to keep members of the group informed
of the others' whereabouts as they moved along in the same direc-
tion strung out over a distance of 100 feet or more. When the
group had travelled approximately 100 yards down the creek
channel, they came to a road where I was standing. One, coming
within a few feet, suddenly took alarm at my presence and climbed
a tree, where it stayed for five minutes, frequently answering the
calls of other members of the group. Two others appeared, one
keeping close behind the other. They were attracted by the calls
of the treed individual, and almost brushed against me, then shied
away upon noticing my presence.

Stuewer (1943:226) stated that the normal feeding range in-
cludes distances up to one mile from the home base but that rac-
coons will travel even farther, if necessary, to obtain a preferred
food. Whitney and Underwood (1952:118) state that a hungry
raccoon may travel several miles, in a meandering course, in one
night, and they cite an instance of a female and her three young
that travelled 2/2 miles to a cornfield for their nightly foraging. Sum-
marizing 459 records, for 87 animals, Stuewer found the average
range to be 503 acres for adult males (maximum 2012 acres), 268
acres for adult females and juvenile males, and 111 acres for juve-
nile females.

Mustela frenata Lichtenstein

Long-tailed Weasel

Status. — Scarce and seldom seen; it is not known whether a perma-
nent populations is present.

Habitat. — No definite preference has been noted; probably it
ranges over the entire area. One was seen in March hunting in the
daytime at the edge of a fallow field overgrown with giant ragweed
and sunflower. Two were caught at different times and places but
in similar situations in wire funnel traps set for snakes along hilltop

Home Ranges of Vertebrates 167

rock ledges in oak-hickory woods. Another was seen in this same
type of habitat.

Movements. — No information has been obtained. The fact that
no weasel or sign has been seen more than once in any one place
seems to suggest that these secretive small predators wander
extensively.

Spilogale putorius (Linnaeus)
Spotted Skunk

Status. — Common resident.

Habitat. — Records of the spotted skunk on the Reservation are
so well scattered in both woodland and grassland situations, that
no definite preference for any habitat subdivision is detectable.

Movements. — In 1950 during the colder months a line of 50 or
more live-traps were kept in operation and were moved from time
to time. These traps were set primarily for opossums, but spotted
skunks were caught frequently. In the course of a year 18 indi-
viduals were caught within an area of approximately half a square
mile. All of these were marked by toe-clipping and released, but
the data from recaptures are so scanty that little can be learned
from them concerning movements. Only two marked skunks were
recaptured. One of these was caught at the same place on Janu-
ary 3, January 15, and March 27. An adult female was trapped and
marked on October 16, 1950; she was recaptured on March 8, 1951,
at a point 900 feet southeast, and on April 4, 1951, she was caught
approximately 3300 feet south of the original location. These rec-
ords, together with the fact that most individuals marked were not
recaptured at all, indicate that an individual skunk ranges over
an area considerably larger than the area live-trapped.

Crabb (1948:232) in southeastern Iowa found that in winter a
spotted skunk has a home range of about .25 square mile with
two or three separate dens. He found that in spring the range was
similar to that of late winter for the females, but the male's range
was greatly expanded, to an area of two to four square miles.

Mephitis mephitis (Schreber)
Striped Skunk
Status. — Common resident.

Habitat. — This skunk prefers grassland, especially in woodland
edge situations, for hunting its prey. The captures recorded ( mostly
in wire live-traps set for opossums) were nearly all in grassland
within a few yards of woodland edge, or near margins of streams or

168 University of Kansas Publs., Mus. Nat. Hist.

the pond. So far as has been observed dens are usually situated
in rock clefts along hilltop ledges in woodland.

Movements. — Eleven striped skunks were live-trapped in a six-
months period and within an area of approximately one-third of a
square mile. Only two of the eleven were recaptured. One of these
was caught at the same place on March 30 and April 4, but was
caught at a second location 750 feet away on April 8. Another
skunk, caught within a few yards of a currently used den on No-
vember 26, 1949, was recaptured 1700 feet away on April 12, 1950.
Most captures of skunks were made along edges of fields of brome
grass, formerly pastured. Several captures were made along the
channel of a small creek draining the eastern half of the Reserva-
tion. Skunks released from live-traps did not, like most other ani-
mals, make for the nearest shelter, but struck out with seeming
fixity of purpose and slight deviation from a straight line course, on
relatively long treks, with the home burrow as the obvious objec-
tive. Several times attempt was made to follow such skunks, but
usually they eluded pursuit by scrambling through thickets or
tangles. One such individual, caught at the head of a small valley,
travelled 700 feet uphill in its homeward dash to shelter in a fissure
at the hilltop rock outcrop.

Dama virginiana (Boddaert)
White-tailed Deer
Status. — Irregular resident.

Habitat. — Chiefly woodland, especially where there is an under-
story of dense brush; on many occasions seen crossing open fields,
or feeding along their edges.

Movements. — Twelve occurrences of deer were recorded from
August 18, 1949, to May 25, 1951. All these records probably per-
tained to the same two individuals, which were usually seen to-
gether, a doe and her yearling fawn. However, none was seen from
October, 1949, to August, 1950. From May, 1951, to March, 1956,
no deer or sign of them were recorded on the Reservation, but
subsequently in 1956 and 1957 at least one individual was present
on the area and was recorded frequently.

The records of 1949 through 1951 were well distributed over an
area of a little more than a quarter of a square mile, rhomboidal in
shape and including a narrow hilltop field, the wooded slopes on
either side of it, and the meadows at the heads of two small valleys
approximately half a mile apart. The area seemed to provide favor-

Home Ranges of Vertebrates 169

able habitat conditions. It included approximately two miles of
woodland edge thickets, and the meadow portions were being in-
vaded by many kinds of woody vegetation. The area was just over
I mile in greatest diameter from the southwestern edge to the north-
eastern, and § mile from the southeastern limit to the northwestern.
This entire area was not covered uniformly by the deer. The two
northwesternmost records were near the Reservation headquarters.
Deer or their sign would have been readily noticed in this vicinity
and it seems obvious that the animals rarely ventured so far in this
direction. In their normal daily activity the animals must have been
Hmited to a smaller home range, from which they wandered occa-
sionally for a variety of reasons.

The deer seen on August 18, 1949, was in the center of a hilltop
field late in the afternoon. It seemed nervous but was little alarmed
by the nearness of the observer. It stood watching curiously, from
time to time giving a snort, or walking a few steps in one direction
or another. A few minutes before this deer was seen, coyotes were
heard yapping excitedly a few hundred yards farther east, and this
disturbance probably had caused the deer to move into the open.
It finally moved off into the woods on the opposite side of the field
from that where the coyotes had been heard. On May 9, 1951, near
the central part of the Reservation, tracks of a running deer and a
running coyote were seen. Probably the coyote had been chasing
the deer, but their courses could be followed for only a few yards
because of the rocky nature of the soil. In late November, 1950,
after the pond had dried, deer tracks were extremely numerous in
the muddy bottom. One or more deer were visiting the area fre-
quently to feed upon the succulent vegetation.

In the period of this study the Reservation offered habitat condi-
tions favorable to deer. Dense cover was abundant, with a variety
of shrubby and herbaceous vegetation for food, freedom from dis-
turbance by humans and from competition with livestock. The
590-acre area would undoubtedly support a large number of deer.
The fact that the species still, in 1957, has not become well estab-
lished in the area can be attributed almost entirely to domestic
dogs. In the course of a year many individual dogs hunt on the
area, usually in twos or threes, or sometimes in larger packs. Some
have been recognized as residents of nearby farms; others perhaps
travel many miles to reach the area. Their quarry includes rac-
coons, coyotes, opossums, squirrels, and rabbits. Hunting is limited
almost entirely to the colder half of the year. In summer, because
of the abundance of insect pests, dense thorny vegetation, heat and

170 Uni\'ersity of Kansas Publs., Mus. Nat. Hist.

humidity, the dogs' hunting activity is much reduced. Were it not
for this respite during late pregnancy and the period when small
young require care, deer probably would be unable to survive at all
in this part of eastern Kansas. As it is, the population is kept
sparse and scattered. Deer are still sufficiently uncommon that
notice of them by farmers or other individuals are frequently re-
ported in local newspapers. In such accounts there is often mention
of chasing by dogs, and such incidents as the chasing of a nearly
exhausted deer into a farmyard or into the edge of town are com-
mon. If chasing by feral and free-ranging dogs could be partially
controlled, a rapid rise in the deer population of eastern Kansas
might be expected.

Birds

Podilymbus podiceps (Linnaeus)

Pied-billed Grebe

Status. — Occasional migrant.

Habitat. — Sizable bodies of water are required for existence of a
permanent population.

Movements. — On at least one occasion a pied-billed grebe was
seen in the pond at the Reservation, and on several other occasions
lone individuals have been seen on a small pond several hundred
yards south of the Reservation. In each instance the grebe was
present on only one day. No specific dates were recorded.

Pelecanus erythrorhynchos Gmelin
White Pehcan
Status. — Uncommon migrant.

Habitat. — Large lakes, or bays and inlets of the sea coast.
Movements.— On an afternoon in late May, 1949, pelicans in a
large flock were seen flying north, high over the Reservation. On
June 11, 1953, at 8:45 A. M., a flock of approximately 25 were seen
flying slightly above the level of the tree tops, headed north and
a little west.

Ardea herodias Linnaeus

Great Blue Heron

Status. — Occasional visitant, especially in April and May.

Habitat. — Normally limited to vicinity of lakes, ponds and
streams.

Movements. — These large herons pass over the Reservation fre-
quently but irregularly, usually without stopping. They have been

Home Ranges of Vertebrates .171

seen or heard passing over most often around dusk. Usually such
herons were flying high, on what appeared to be long overland
trips. From time to time a lone individual or a pair has stopped
temporarily at the pond, or at one of the small intermittent creeks
on the area, attracted by an abundant food supply of frogs and
tadpoles. In most instances the leopard frog or the cricket frog,
or the tadpoles and young of the bullfrog seemed to be the chief
prey. In some instances a heron has remained for a few hours,
but failed to return after completing its foraging for the day. In
other instances, what appeared to be the same individual returned
day after day to exploit an easily available food source, though
leaving each evening for a distant roost.

Specific dates when great blue herons were recorded from the
Reservation were the following: 1949: September 19, November 24;
1950: April 9, April 13 (one), July 23 (one); 1951: April 14 (one),
September 14; 1952: March 30 (one), May 22 (one), October 18
(one); 1953: May 19 (two); 1954: April 19 (four), May 28 (heard);
1955: June 10 (one), June 11 (one); 1956: April 18 (two), April 26,
May 1 (one). May 15 (one). May 30 (one); 1957: April 7 (three),
April 30 (one). May 6 (two), October 19 (one), October 26 (one).

According to a survey of nesting colonies in Kansas, made by
Andrews and Stephens (1956:295), the nearest rookeries are ap-
proximately 45 miles west, near Maple Hill, Wabaunsee County,
and 35 and 40 miles southwest, near Scranton and Burlingame,
Osage County. Since most of the records obtained on the Reserva-
tion fell within the nesting season, the status of the herons involved
is doubtful. Some may have been non-breeding adults, either resi-
dents of the general area or wanderers. Some may have been nest-
ing apart from the rookeries, at places within a few miles of the
Reservation. However, Andrews and Stephens obtained no records
of nesting colonies in the two easternmost tiers of Kansas counties,
including Douglas County.

Butorides virescens (Linnaeus)
Green Heron
Status. — Common summer visitant.

Habitat. — Normally limited to vicinity of lakes, ponds and streams
in or near woodland.

Movements. — These herons have frequently been attracted to
the pond on the Reservation, and occasionally have come to the
two small intermittent creeks. Frogs constitute the chief food
supply attracting them to these places. The cricket frog is the kind

172 University of Kansas Publs., Mus, Nat. Hist.

most utilized. On various occasions the herons have been seen
strolling about on mud near the pond edge where these small frogs
were extraordinarily abundant. Dates when green herons were
recorded at the Reservation were as follows: 1950: July 13 and 15;
1951: April 25 and 27, xMay 2, June 22, August 6; 1952: May 3, 23,
24, 28, and 30; 1953: May 9; 1954: May 6; 1955: April 27 and 28,
May 8, July 4 and 19; 1956: April 14, 25, 26, 27, 28, and 30, May 1,
4 and 13; 1957: April 30 and May 3.

In some instances the green herons that came to the pond prob-
ably stayed there continuously for several days, or even longer
periods. In 1956, especially, a pair was seen regularly during the
latter half of April and early May. They seemed to be preparing
to nest tliere, but left suddenly after heavy rains on May 7, 8, and 9.

Several of tlie green herons that appeared in late summer were
juveniles. On July 19, 1955, in late afternoon, one perched on a
treetop 200 yards from the pond for more than an hour, calling at
intervals of approximately five minutes.

Nycticorax nycticorax ( Linnaeus )

Black-crowned Night Heron

Status. — Occasional transient.

Habitat. — Swampland is the preferred habitat. On several occa-
sions migrant night herons have been flushed from the willow
grove or the pond edge on the Reservation.

Botaurus lentiginosus (Rackett)
American Bittern
Status. — Occasional transient.
Habitat. — Marshland.

Movements. — Only one record — an individual flushed from the
pond on May 9, 1953.

Branta canadensis (Linnaeus)

Canada Goose

Status. — Uncommon migrant.

Habitat. — Marshlands.

Movements. — These geese have been seen only as migrants pass-
ing over the area in spring. On April 14, 1951, at 8:30 A. M., a
flock of approximately 40 were seen flying low, headed due north.
On March 19, 1952, at 8:30 A. M., a flock of eleven were seen flying

Home Ranges of Vebtebrates 173

north. On April 4, 1955, at 7 A. M., a solitary individual was
flying low, headed north.

Anser albifrons (Scopoli)

White-fronted Goose

Status. — Rare migrant.
Habitat. — M arshland .

Movements. — R. W. Fredrickson recorded a small flock over
the Reservation on April 14, 1952.

Chen hyperborea (Pallas)

Snow Goose

Status. — Abundant migrant in both spring and autumn.

Habitat. — Marshland; arctic tundra in breeding season.

Movements. — The remarks in the species account of the blue
goose apply almost equally well to the snow goose. Relatively few
of the flocks seen were composed entirely of either kind, but both
snow geese and blue geese were usually in the same flock in varying
ratios. Also, a high proportion of individuals in a flock often were
intermediates, that were partly white and partly "blue" in their
plumage. Complete freedom of interbreeding is indicated.

Chen caerulescens (Linnaeus)

Blue Goose

Status. — ^Abundant migrant in both spring and autumn.

Habitat. — Marshland; arctic tundra in breeding season.

Movements. — In 1949 flocks were seen frequently between Oc-
tober 22 and November 13. In 1950 the northward migration oc-
curred mainly during the first week of March. In 1951 the fall
migration reached its peak on October 14. In 1952 the main spring
flight lasted for approximately two weeks in mid-March. In 1953
the main spring migration occurred in the period March 8 to 15,
but flocks were seen also on March 18, 19, 23, 25, and 30, and on
April 1, 2, 3, 4, 8, and 17. A large flock seen on the latter date was
flying south. Southward migration in autumn was first recorded on
October 13 and reached a peak about October 24, but flocks were
recorded also on October 28 and December 20. In 1954 spring
migration was noted on March 7, 8, 10, and 12, reaching a peak be-
tween the two latter dates. The fall migration was recorded only
on October 7 and IS. In 1955, spring migration was noted chiefly

174 University of Kansas Publs., Mus. Nat. Hist,

between March 10 and 31, and fall migration was first noted on
October 7. In 1956 spring migration began on February 22, reached
a peak on March 19, and was last noted on March 26. In 1957
spring migrants were noted on March 1, 2, 3, 4, 12, 15, and 27, and
on April 6, fall migration began the second week of October.

Anas platyrhynchos Linnaeus

Mallard

Status. — Irregular transient over area, occasionally stopping at
pond when water supply is adequate.

Habitat. — Marshes, ponds, lakes and slow-moving streams.

Movements. — Mallards have been recorded at the pond on the
Reservation on only a few occasions: March 19, 1950 (flock of
eight); June 11, 1951 (lone female); October 31, 1951 (pair); No-
vember 1, 1951 (group of three); February 26, 1952 (pair); Febru-
ary 28, 1952 (pair); March 30, 1952 (about 35); April 3, 1952
(about 35); November 11, 1955 (lone female). Low water, or
complete drying of the pond, in late 1952, 1953, and 1954, dis-
couraged use of the pond in those years by ducks; also probably
the population was low as a result of widespread drought. On
many occasions, including December 9 and 24, 1951, November -5,
1954, March 3, 1955, February 21, 1956, and January 20 and 24,
1957, flocks of mallards were seen or heard flying over the area,
although they did not stop there. Most often such flights were
noted after dark.

Anas acuta Linnaeus
Pintail
Status. — Occasional transient.
Habitat. — Ponds, lakes and streams.

Movements. — The first evidence of the occurrence of this species
on the Reservation was the strewn feathers of an adult male where
it had been eaten by a predator (probably a horned owl) in De-
cember, 1948. The remains were found approximately 100 yards
from the pond, where the duck probably had stopped. It may have
been wounded in the hunting season. On many occasions ducks
seen flying over the Reservation were too high to be identified posi-
tively. Some may have been pintails. R. W. Fredrickson recorded
a small flock of pintails flying over the Reservation on March 5, 1949,
and a lone individual on April 6, 1952.

Home Ranges of Vertebrates 175

Anas carolinensis Gmelin
Green-winged Teal
Status. — Uncommon visitant in late autumn and early spring.
Habitat. — Marshes, ponds, lakes and slow-moving streams.
Movements. — Flocks were recorded at the pond almost daily from
November 7 to 14, 1949, March 3 to 11, 1950, and were also recorded
on March 4 and 31, 1955.

Anas discors Linnaeus
Blue-winged Teal
Status. — Common visitant, chiefly in spring and autumn.
Habitat. — Marshes, ponds, lakes, and slow-moving streams.
Movements. — From one to 40 individuals were noted at the pond
on the following dates: June 27, 1949; April 12, 13, 15, 20 and 23,
1950; April 13, 15, 16, and 19, and October 29 and 30, and No-
vember 22, 1951; April 1, 2, 7, 21, and 23, 1952; April 6, 1953;
April 17, 20, and 29, 1955; and April 9 and 30, 1956.

Spatula clypeata (Linnaeus)
Shoveller

Status. — Uncommon visitant.

Habitat. — Marshes, ponds, lakes, and slow-moving streams.

Movements. — Throughout April, 1952, one or more groups of
these ducks were seen frequently at the pond, which was then full
of water. Often the shovellers were associated with blue-winged
teal. The number seen at one time ranged from six to fifteen. In
other years none was seen at the Reservation.

Aix sponsa (Linnaeus)

Wood Duck

Status. — Uncommon visitant.

Habitat. — Ponds and slow-moving streams in wooded areas.

Movements. — This duck has been noted on the Reservation on
only a few occasions: May 29, 1950 (adult male); April 17, 1951
( adult male ) ; November 3, 1951 ( adult male ) ; November 4, 1951
(adult male); and March 20, 1952 (pair). Although the records
in 1951 extended over more than six months, it is almost certain
that the same individual was involved in all three instances. This

176 University of Kansas Publs., Mus. Nat. Hist.

was a crippled and flightless male. On the night of November 3,
the pond froze over and when the duck was found on November
4, he was 200 yards farther down the gully, where there was still
a trickle of water.

Aythya coUaris (Donovan)
Ring-necked Duck
Status. — Uncommon visitant.
Habitat. — Ponds, lakes and streams.

Movements. — Recorded at the pond on the Reservation in 1950
only; on March 11 there were approximately 15, along with a larger
number of green-winged teal; on March 20 there were approxi-
mately 20; on March 21, seven.

Aythya affinis (Eyton)

Lesser Scaup Duck

Status. — Rare visitant.

Habitat. — Ponds, lakes and streams.

Movements. — On March 15, 1952, a lone female was flushed
from the pond, and a flock of approximately ten were seen at a
farm pond half a mile south of the Reservation. On the following
day they were seen again at the same farm pond, along with a
group of shovellers. On March 24 several were seen flying to the
pond on the Reservation late in the afternoon.

Cathartes aura (Linnaeus)
Turkey Vulture
Status. — Regular summer visitant.

Habitat. — Regularly patrols all parts of the Reservation, and
most other areas in northeastern Kansas that are not subject to
too much disturbance by humans.

Movements. — Vultures on their northward migration usually
reach the Reservation in the last week of March. For the years
1951 through 1957, arrival dates were the 25th, 19th, 20th, 25th,
28th, 27th and 31st of March. In all but one instance the early
records were of lone birds, but in succeeding weeks, vultures were
most often seen in small groups. A close-knit social group of four
pairs used the Reservation regularly. In their flight maneuvers
and search for food the members of this group tended to stay near
each other, and usually several or all of them were seen simul-
taneously. Within the group, pair-associations were nearly always

Home Ranges of Vertebrates 177

evident, as the members of each pair stayed closer to each other
than to the other vultures belonging to the group. Activity of the
entire group centered over "East Woods" and "Wall Woods," an
area of rocky hilltops and steep slopes near the eastern boundary
of the Reservation. When this part of the area was visited, the
vultures were almost invariably seen maneuvering over it, whereas
they were seen comparatively seldom over the part of the Reserva-
tion near the headquarters, two-thirds of a mile farther west. A
large proportion of their time seemed to be spent in an area esti-
mated to be less than 200 acres in extent, but whether this area
provided their main food supply, or merely served as a rallying
place where members of the group maintained contact and per-
formed maneuvers primarily of social significance, is uncertain.
Vultures that probably were members of this same group were often
noted approximately I/2 miles farther south at the north edge of
the Kansas River Valley, about the same distance north, and,
occasionally, over the west boundary of the Reservation, a mile west
from the place where they were most frequently seen. Nothing was
recorded of their movements east of the Reservation, but a mini-
mum area of perhaps six square miles is estimated as the range
of this group.

In the area where the group's activity was concentrated, a nest
was discovered in 1952, and the same site was used each year sub-
sequently, through 1956. This nest site was in an elm more than
two feet in trunk diameter, and having a large hollow in the trunk
ten feet above the ground. When the nest site was discovered,
diere was evidence that it had been robbed recently. The rough
bark of the tree was coated with the vulture's down feathers, where
the bird might have struggled with a chmbing predator that was
either approaching or leaving the nest. In 1954, also, the nest
was robbed by a predator before the eggs had hatched. The rac-
coon was considered the most probable predator.

Accipiter gentilis (Linnaeus)
Goshawk
Status. — Rare and irregular winter visitant.
Habitat. — Typically northern coniferous forests, but because of
its migratory habits may appear temporarily in much different habi-
tats, such as deciduous woodlands in eastern Kansas.

Movements. — Recorded on the Reservation on only two occasions.
One was on December 29, 1950. At 10:00 A. M. alarm calls of a
flicker and squaUing of a rabbit were heaid on a heavily wooded

178 University of Kansas Publs., Mus. Nat, Hist.

north slope near the Reservation headquarters. Two cottontails,
alarmed by the sound scurried from the edge of the woods and hid
in nearby blackberry thickets. Hurrying toward the sounds, I
flushed the goshawk, from beneath an osage orange tree. The hawk
was carrying an adult cottontail but soon dropped it. The rabbit
struggled to its feet and ran heavily a few yards to an old woodrat
house where it found shelter. The hawk boldly perched within 100
feet of me, watching intently. I easily caught the rabbit, which was
near collapse, and it died a few minutes later. Dissection disclosed
that the jugular vein had been pierced, and there were masses of
clotted blood in subcutaneous wounds. The back of the head, right
side of the neck, and throat had been badly lacerated. A second
goshawk was recorded on the area by Donald W. Janes in the winter
of 1955.

Accipiter striatus Vieillot

Sharp-shinned Hawk

Status. — Occasional transient, chiefly in winter.

Habitat. — Chiefly woodland.

Movements. — The few records seem to pertain to wandering in-
dividuals, since most are well separated in time and space. None
has been noted in summer. Specific dates of the records are: Sep-
tember 19, 1956; September 23, 1952; October 13, 1948; November
3, 1956; November 16, 1951; November 17, 1954; December 1, 1956;
December 2, 1956; December 15, 1953; December 16, 1954; Decem-
ber 31, 1955; January 9, 1954; January 18, 1952; January 24, 1953;
February 2, 1954; February 4, 1952; February 14, 1954; March 8,
1953; April 15, 1950; April 26, 1956, and May 15, 1952.

Accipiter cooperii ( Bonaparte )
Cooper Hawk
Status. — Uncommon summer resident; occasional winter visitant.

Habitat. — Chiefly woodland, and mixed habitat of woodland al-
ternating with open fields.

Movements. — Cooper hawks have been recorded on the Reserva-
tion most frequently in spring and early summer, the nesting season,
when usually at least one pair resides on the area. Records in winter
are sporadic, and seem to pertain to wandering individuals that are
passing through.

A nest, in dense oak-hickory woods, on a north slope 100 yards
south of the Reservation headquarters was used in 1950, 1951, 1952,

Home Ranges of Vertebrates 179

and 1953. In 1954 a new site approximately 100 yards east, on the
same slope was occupied. In 1955, 1956, and 1957 the territory was
not occupied, but Cooper hawks were seen from time to time in the
area.

Each year that the territory was occupied, the hawks appeared
abruptly in spring and passed through a period of maneuvering and
display before nesting actively began. In 1952 the hawks appeared
at about the beginning of March, and were heard cackling fre-
quently in the vicinity of the nest site. In 1953 first appearance was
nearly a month earlier. Although the bird that arrived first was
seen frequently in the territory in February and March, it did not
obtain a mate for several weeks. On April 1, 1953, the pair of hawks
were seen in a courtship flight. The wings of the male were raised
high over his back at each stroke, and moved in a wide arc, giving
the flight a peculiar mechanical quality faintly resembling the flight
of a nighthawk. Similar display flights were observed frequently
throughout most of April. On March 8, 1954, the first Cooper hawk
of the season was seen circling with the same slow rhythmic flap-
ping, over a hilltop field area of 32 acres. Apparently this individual
was a female.

Because Cooper hawks were ordinarily shy and secretive, the
extent of their territories was difficult to ascertain. In late July
and early August, 1951, a hawk was seen to bring young chickens
regularly from adjoining farms. The farmyard which probably con-
tributed most of the chickens was just under % mile from the nest,
while the other farm was approximately .6 mile from the nest. The
chickens appeared to be about one-third grown, and were heavy
loads for the hawk. Both farmyards were on a flat hilltop approxi-
mately 115 feet above the elevation of the nest. The hawk, prob-
ably the female in each instance, was favored by the lower elevation
of the nest, when burdened with heavy prey on her homeward
ffight. The area over which the pair of hawks were seen to fly, and
which they were known to exploit for prey, consisted of at least half
a square mile, but the actual territory was probably larger. In 1950,
1951, and 1952 other Cooper hawks were seen frequently on a
wooded slope a little less than half a mile south southeast of the
nest near the headquarters. Although a nest was never found in
this second area, nesting almost certainly occurred there. In 1952
a nest was found in a ravine .6 mile east of the nest under observa-
tion near the headquarters, and approximately .8 mile north north-
east of the second supposed nest site.

180 UNnTRSiTY OF Kansas Publs., Mus. Nat. Hist.

In 1951 the three young hawks left the nest near the Reservation
headquarters in the last week of July. For three weeks subsequently
they were heard or seen daily on the slope where the nest was lo-
cated and within 100 yards of it.

Buteo jamaicensis (Gmelin)

Red-tailed Hawk

Status. — Year-round residents, winter residents, and transients
occur on the area.

Habitat. — Red-tailed hawks use the entire area. While most
perches and nest sites are in woodland, the hawks hunt their prey
chiefly in open land. Parkland and woodland edge situations are
especially favorable.

Movements. — Individual hawks could not be recognized with
certainty except for relatively short periods when missing remiges
or retrices occasionally permitted recognition at close range. Be-
havioral traits provided more useful but not entirely reliable means
of recognizing individuals. In most instances, however, these means
of recognition were inadequate to trace the turnover in the resident
population. Of the red-tailed hawks seen on the area, most were
wanderers that did not stay long.

Most of the section of land within which the Reservation is located
was divided between the territories of two pairs of red-tailed hawks
(Fig. 11). These two territories were maintained in essentially the
same form throughout the period of my observations, from 1948
through 1957, although there was partial replacement of the hawks
themselves. The configurations of the territories were largely de-
termined by local physiography. The upland cuesta north of the
Kansas River Valley has peninsular extensions. One of these exten-
sions, a little more than a mile long but less than 100 yards across
at its narrowest, crosses the Reservation diagonally from the north-
eastern to the southwestern corner, as a mesalike flat hilltop. There
is a second peninsular hilltop, more or less parallel to this one but
% mile farther southeast, and a third about 1/i miles northwest. These
parallel hilltops formed territorial boundaries, and the territories
consisted mainly of the intervening lowland areas, and slopes. Of
the two territories, the more western was by far the larger, com-
prising approximately 800 acres, but it included much cultivated
land that was less productive of prey. The more eastern territory
was approximately 535 acres of which less than a fifth was under
cultivation. The remainder was mostly woodland, with brush and

Home Ranges of Vertebrates 181

grassland of former pastures. The territory of this pair included
no human habitations nor roads, and was mainly inside the Reser-
vation on the part least visited by people. The pair of hawks occu-
pying this territory are believed to have been the same throughout
the eight-year period of observation. Their behavior diflFered
notably from that of other hawks in their boldness and defiance.
In May, 1949, 1 tried unsuccessfully to climb to their nest and there-
after they showed resentment whenever I entered their territory,
regardless of whether or not the intrusion was in the nesting season.
Upon seeing me as much as half a mile away both hawks might
leave their perches, fly toward me screeching, and circle low over-
head. Over the years their hostihty subsided somewhat, as their
nests were not further molested. In 1950 and 1957, the nests were
in trees considered unclimbable, and in other years nests were not
found while they were in use. The heart of this pair's territory con-
sisted of thick woods of approximately 34 acres on a west-facing
slope. On dozens of occasions, the hawks were seen perched on
snags on this hillside.

Hawks of the second pair were much shier and more wary. Their
territory included the Reservation headquarters, and the residences
of four different farms and several roads. As a result, the hawks
often encountered humans and probably had been shot at on many
occasions. It is certain that in this pair there was occasional replace-
ment when one was shot and the survivor obtained a new mate to
share the territory. Whether there was continual tenure through
such a line of survivors over the eight-year period is uncertain, but
probable in view of the fact that the territory remained essentially
the same. This territory centered on a brushy hillside, which had
been wooded until it was cut and bulldozed in the winter of early
1949. Cottontails, rodents and small birds were abundant in the
brushland habitat. Hunters, with shotguns or .22 rifles, often visited
this hillside. When disturbed by humans, the hawks habitually
flushed at distances of hundreds of yards and flew to distant parts
of their territory. Sometimes they screamed at the moment of
flushing, and sometimes flew silently. Their survival obviously
was dependent on wariness and prompt response to the presence
of humans. A melanistic individual was co-occupant of the territory
from the autumn of 1948 until November 3, 1951, when it was found
dead beside the county road at the west edge of the Reservation.
On three other occasions buteos recently shot were found along this
road. Probably wandering individuals suffered the most casualties.
Despite their usual wariness, these hawks when hunting were

182 Uni\trsity of Kansas Publs., Mus. Nat. Hist.

several times seen to fly through or past a barnyard, in a surprising
display of boldness. Although they were not seen to catch any-
thing in or near the barnyards, their behavior suggested that young
poultry might be the object of their forays.

In May and June, 1955, the hawks were seen on various occa-
sions carrying prey (usually a snake) to the nest. In each instance,
the hawk was within a quarter mile of the nest when first seen.
On June 19, a fledghng left the nest when attempt was made to
climb to it. For the next six weeks, the fledglings were heard and
seen frequently in the parents' territory. On several occasions they
were seen soaring with the adults, but tended to be much more
sedentary, and often perched for periods of hours in the same small
area, calling to be fed.

Territorial encounters between pairs were relatively few. Even
when such encounters occurred, they were usually brief and rela-
tively mild, and might consist of circling and screaming with no
actual fighting. This was in striking contrast with my observations
on the species at the San Joaquin Experimental Range in the Sierran
foothills of central California (Fitch, Swenson, and Tillotson, 1946),
where habitat conditions were somewhere near the optimum, with
high populations of diurnal rodents, and rabbits, sparse cover, and
well distributed perch trees. Under these conditions territories
averaged only half a square mile, and territorial pressure was in-
tense. Each pair of hawks spent a substantial part of their time
in fights and maneuvers wdth their neighbors as a result of terri-
torial pressure.

Trautman (1940:212) found that red-tailed hawks, wintering in
the vicinity of Buckeye Lake, Ohio (where the species is absent in
summer), settled in individual territories of t\vo square miles or
less in extent. Having settled in such territories, the hawks ordi-
narily remained for the winter, unless forced to move by unfavor-
able weather conditions such as a heavy snow cover. After being
forced out of its regular territory by a winter storm, Trautman
found, an individual hawk would return as soon as conditions again
became favorable.

Before there were buildings on the Reservation, in 1948 and 1949,
the small valley that was later the site of the headquarters installa-
tion was much used by the hawks. The residence and other build-
ings were erected in the winter of 1949 and 1950. Thenceforth
human activity greatly increased, and the hawks habitually avoided
this part of their territory, and used the valley much less than they
had before. They rarely came within 100 yards of the buildings,

Home Ranges of Vertebrates

183

and perched in the valley within sight of the headquarters only at
times when there had been no human activity outside for periods of
hours. They were most often seen perched 200 yards or more from
the house in early morning. As soon as a person came out of the
house, they would flush and leave the vicinity and would not reap-
pear for the remainder of the day.

4^

Fig. 11. Approximate extent of two territories of red-tailed hawks as they
existed over a period of years. The long, narrow hilltop extending from
northeast to southwest across the Reservation was the common boundary.
The territory on the northwest was larger, as it contained a high proportion
of cultivated land that, for the hawks, was unproductive of food. Dots
show locations of nests for the years in which they were found.

Buteo lineatus (Gmelin)
Red-shouldered Hawk

Status. — Uncommon resident of this general area near western
limit of breeding range.

Habitat. — Deciduous forest is preferred by this hawk. On the
Reservation it was rarely seen in situations where there were mature
trees of climax species. Rather it showed preference for such places
as the pond and the adjacent willow groves, hilltops with low trees
and thorny tangles, and areas of mixed grassland and second growth
woodland.

184 UNTVTRsiTi' OF Kansas Publs., Mus. Nat. liisT.

Movements. — Compared with other buteonine hawks on the area,
red-shouldered hawks are more secretive. Although I had heard
the characteristic call from time to time, I was unfamiliar with it,
and failed to recognize the species' presence until June 22, 1953,
when a juvenile was flushed at close range at the edge of the pond.
Judging from the appearance of this individual, it had not been
fledged long, and could not have travelled far from the nest. No
m.ore definite evidence of nesting on the area has been obtained.
Throughout the latter half of 1953, 1954 (except March and De-
cember), 1955 (except January and February), and early 1956, a
red-shouldered hawk was seen and heard frequently on the Reser-
vation. Twice in May, 1955, two were seen or heard simultaneously,
near together and perhaps paired, but the records pertain mainly to
a lone individual that spent most of its time on the Reservation,
but occasionally sojourned elsewhere for periods of weeks.

Reason for considering the records to pertain to a single individual
is that the hawk was nearly always seen or heard in the same gen-
eral area, an elliptical territory of 180 acres, approximately twice
as long as broad, mostly in the northwestern quarter-section of the
Reservation (Fig. 13). Habits of this individual were noted to
change over periods of weeks or months. It tended to spend much
of its time in a small perching area, but from time to time neglected
or altogether deserted one perching area in favor of another that
was as much as half a mile away. In March and April, 1955, a red-
shouldered hawk was seen at several places along the eastern edge
of the Reservation, more than half a mile from the supposed terri-
tory. This may have been a second individual, but more probably
it was the same. Stewart (1949:26) found red-shouldered hawks
in a density of one pair per .8 square mile in the coastal plain of
Maryland, in the habitat to which the hawks were confined —
wooded stream courses.

The territory of the lone individual on the Reservation overlapped
to a large extent the territory of a pair of broad-winged hawks, and
also broadly overlapped a red-tailed hawk territory. Although not
limited to mutually exclusive territories, these three species of large
hawks were somewhat intolerant of each other. On the Reserva-
tion interspecific maneuvers, chases or fights involving two species
were observed much more frequently than intraspecific encounters.
Domination of the red-shouldered hawk by a red-tailed hawk was
observed on various occasions. The red-shouldered hawk prob-
ably was able to maintain its territory only because it was limited

Home Ranges of Vertebrates

185

chiefly to areas little used by red-tailed hawks. In encounters be-
tween the red-shouldered hawk and broad-winged hawks, domi-
nance was not clear-cut, nor was any actual fighting observed. The
encounters consisted of soaring in overlapping circles with an oc-
casional short swoop toward, but not near to, the opponent.

Fig. 12. Approximate extent of the territory of a pair of
broad-winged hiiwks, as it was maintained each summer
over a period of years. It was an area of hilltop pasture
and rocky, brushy south slope. Dots show approximate lo-
cations of nest sites in four different years, but nests were
not actually found in 1951 and 1955.

Fig. 13. Approximate extent of territory of a red-shoul-
dered hawk, in the northwestern part of the Reservation
and adjacent areas. This territory was shared with red-tailed
hawks, and (in summer) broad-winged hawks, but there
was constant friction between all three species.

186 Univtrsity of Kansas Publs., Mus. Nat. Hist.

Buteo platypterus (Vieillot)

Broad-winged Hawk

Status. — Moderately common resident in summer, absent from
October to late April.

Habitat. — Mixed woodland, and adjacent open areas, especially
of rock ledge.

On the Reservation, the abandoned rock quarry was a favorite
haunt of these hawks, and on many occasions was the scene of
territorial fights. The abundance of reptilian prey of several kinds
at this open, rocky place made it attractive to the hawks. The
hawks were seen chiefly along the south slopes of the northwest
quarter of the Reservation section, where the aspect of the vege-
tation was xeric.

Movements. — Upon their arrival in spring, broad-winged hawks
attract attention by their soaring flight and whistling calls. In
autumn before migrating south they spend much less time soaring
and are relatively silent; consequently the time of departure is
diflacult to determine. Arrival dates over an eight-year period,
1950 through 1957, were all in April: the 15th, 18th, 16th, 28th,
20th, 15th, 20th, and 23rd. For these eight years, April 19 was
the average date of appearance. After the first appearance the
hawks were usually heard and seen many times each day for the
remainder of spring. Within the first two weeks they were espe-
cially noisy and conspicuous, while they were establishing and
maintaining territories against competitors. At this season as many
as five have been seen soaring together, with occasional threatening
swoops and frequent squealing calls. Activity of broad-winged
hawks was mainly hmited to the areas of south-facing slopes with
xeric second growth woodland, rocky hilltop edges, and upland
pasture. Nests were found only in 1954 and in 1957, but probably
the hawks nested on the Reservation every year. In 1951 and 1955
they were seen repeatedly, carrying prey items toward the same
place, a hilltop edge, with low thorny trees and dense tangled
undergrowth, but my many attempts to find the nest were unsuc-
cessful.

For these hawks territorial boundaries were especially difiicult
to define, since the birds were usually seen circling high over the
hillsides. The area over which they were known to forage regu-
larly comprised approximately 250 acres (Fig. 12), but the western

Home Ranges OF Vertebrates 187

and northern limits, being outside the Reservation, were not well
determined. Most satisfactory information was obtained concern-
ing the southern limit of the territory, since it followed the contour
of the slope just north of the residence. In early summer the
hawks were seen many times each day on this slope, but they only
rarely ventured as far south as the residence. The opposite slope
to the south supported a more mesic type of forest and obviously
was not such favorable habitat. Occasionally, however, a broad-
winged hawk was seen to fly out of sight in this direction, in a
foray well beyond the limits of the usual territory. Within the
territory, a much smaller, elongate area of hilltop edge and upper
slope only a few acres in extent included the favorite perches and
the places where the hawks most frequently flew.

In 1954, when the nest of these hawks was studied, prey included
collared lizards of a marked population at an old quarry nearly
a quarter mile east, and several Great Plains skinks, believed to
have been caught along a pastureland outcrop % to % mile north-
west, where these lizards were especially numerous.

Buteo lagopus (Pontopiddian)

Rough-legged Hawk

Status. — Rare transient or migrant in winter.

Habitat. — Typically far northern forest, and tundra.

Movements. — Occasionally rough-legged hawks are seen in north-
eastern Kansas in winter. One was seen on the Reservation on
November 2, 1948; one of the dark phase was seen as it was
heckled by a pair of red-tailed hawks, over the southeastern part
of the Reservation on November 14, 1951.

Haliaeetus leucocephalus ( Linneaus )
Bald Eagle

Status. — Rare transient.

Habitat. — Most typically, forested areas bordering sizable bodies
of water.

Movements. — Only one record of a bald eagle on the Reservation
has been obtained. On the afternoon of December 9, 1951, my
attention was attracted by the screaming of red-tailed hawks over-
head; three of these hawks were chasing an adult bald eagle, which
was flying northwest. It kept to this course until it was out of sight.

188 UNa\'ERsrri' of Kansas Publs., Mus. Nat. Hist.

Circus cyaneus (Linnaeus)

Marsh Hawk

Status. — Appears from time to time at all seasons but mainly in
winter, when nomadic individuals often wander over Reservation.

Habitat. — The marsh hawk usually frequents open fields of
various types. In apparent order of preference, marsh hawks prefer
marshland, prairie, bottomland pasture, upland pasture, parkland,
and woodland. Although marsh hawks have been seen on many
occasions flying over woodland, they were always flying well above
the treetops and presumably were crossing from one open field to
another. On the Reservation they were seen frequently on all open
field areas and especially in the upland field at the north edge, that
had a mixture of weeds and native grasses.

Movements. — Marsh hawks were seen on the Reservation each
month from September through April, inclusive, but ordinarily
they were absent during May, June, July and August. The in-
dividuals that used the area in winter were all wanderers that did
not maintain regular beats; none was recognized on consecutive
days, and there was no regularity in the times or places of their
appearance. In 1951 a pair nested on the prairie tract at the north
edge of the Reservation. The unusually heavy precipitation
probably created conditions favorable for them. On May 8 the
male was heard cackling and squealing, and was seen in a display
flight over a grassy hilltop approximately 3000 feet south southwest
of the spot where the nest was found later. On June 20, when I
was walking across the same hilltop, 400 yards from the nest, the
male flew toward me, cackling, and swooped with a display of
aggressiveness, then withdrew in the direction from which he had
come. As I moved farther in that direction, he again flew to meet
me, this time accompanied by the female, and both swooped at
me. The male was the more aggressive and the female soon re-
tired. Moving back and forth across the field, I was able to detect
corresponding increase or decrease in the male's excitement and
aggressiveness, and was led to the nest which contained a single
downy young. Five days later this young was destroyed by an
unknown predator and the adults disappeared from the area. On
April 7, 1952, a pair was observed in courtship flight in this same
vicinity, but, so far as known, nesting did not occur.

Home Ranges of Vertebrates 189

Falco mexicanus Schlegel
Prairie Falcon

Status. — Rare transient.

Habitat. — Typically grassland, in rugged terrain such as that with
cliffs and buttes.

Movements. — Only one definite record of a prairie falcon on the
Resei-vation has been obtained. On April 16, 1951, one was seen
flying south across the hilltop on the northern part of the Reserva-
tion. On October 18, 1951, a medium-sized falcon, not definitely
identified, but perhaps of this species, was seen flying rapidly over
the southern part of the Reservation, on a southward course.

Falco sparverius Linnaeus
Sparrow Hawk

Status. — Occasional on migration; individuals that live on nearby
land wander farther than usual from time to time, encroaching onto
Reservation.

Habitat. — Requirements include open ground, with sparse or
short vegetation, and high perches from which the hawk may exam-
ine surrounding terrain. In 1948, a pair of sparrow hawks having a
territory centering in grain fields south of the Reservation were seen
frequently in the bottomland pasture of its eastern part, at a time
when many cattle were pastured on the area and grass was kept
cropped close. In subsequent years, after removal of the livestock
and growth of high grass, sparrow hawks no longer included this
area in their range.

Movements. — Sparrow hawks have been seen on the area mainly
in winter and early spring, when the unfavorable effects of tall
grass and other rank herbaceous vegetation are minimized. On
February 23, 1955, at a time when the ground was snow covered, a
male that normally ranged over cultivated land adjoining the Reser-
vation to the west, was found 200 yards inside the Reservation at-
tempting to attack a song sparrow that was in a trap.

On the afternoon of March 23, 1955, this male or another was
found at a bird trap attempting to attack a chickadee. Half an hour
later at dusk, a person coming out of the laboratory building flushed
a hawk, that was probably the same individual, from an old phoebe
nest on the porch. Flying erratically in the semi-darkness the hawk

190 University of Kansas Publs., Mus. Nat. Hist.

crashed into the screen of a rear window, and a moment later,
having circled the building, it lit beside the door and walked in.
Its behavior indicated confusion and unfamiliarity with its sur-
roundings. It was banded and released, and lingered in the vi-
cinity, as in the next four days it was seen twice wdthin a few hun-
dred yards of the headquarters buildings. Cade (1955:11) mentions
one territory occupied by a pair in a vacant lot 200 feet by 300 feet.
He found that in southern California pairs usually remain together
on their territories throughout the winter, though some pairs break
up in fall and occupy separate winter territories.

On the Rockefeller Tract in 1957 sparrow hawks were frequently
present for periods of days and were recorded April 6 to 9, April
24, May 28, July 17, September 3 to 5, and 24 to 26. Probably those
that came from time to time were different individuals. Usually
only a single hawk was present, but on July 17 there was a pair,
and on September 24 a family group of five with recently fledged
young. Invariably activity was concentrated along a line of tele-
phone poles, extending west from the east boundary across the
pasture to the buildings near the center of the quarter-section, and
along a line of electric wdre poles extending north from the south
boundary to the buildings along a road with fallow fields on either
side. There were 12 poles in all. The hawks usually perched on
them ( sometimes on the wires stretched between them ) and foraged
by making flights of usually less than 100 feet from the line to the
ground to secure prey. Food was obtained chiefly in this elongated
strip .43 mile in length and encompassing perhaps ten acres, but at
the maximum a hawk followed over periods of hours and flushed
frequently ranged over an area of somewhat more than 80 acres.
This was a female present on the area on April 6, 7, 8, and 9.

Colinus virginianus (Linnaeus)
Bob-white

Status. — Common resident.

Habitat. — Although quail range over almost every part of the
Reservation, areas in an early stage of secondary succession are
preferred. My records of coveys are concentrated at the pond, the
old quarry, and the formerly cultivated fields, which had grown
up with weed patches adjacent to brush and woodland. Coveys
were noted in woodland of various types but were rarely seen
where stands of awnless brome grass were dominant in former pas-
tures. In the nesting season when coveys had dispersed, pairs were

Home Ranges of Vertebrates

191

often seen in habitats different from those used by the coveys at
other times of year. Pairs were often seen along the edges of pas-
tures.

Movements. — In autumn, winter and early spring, there were as
many as eight coveys using the area, but the number was variable.
Fallow fields of 16.7 acres at the middle of the Reservation on its
western edge provided headquarters for one covey, which likewise
ranged over adjacent fields, to the west, cultivated for sorghum,
corn, and other crops. A second covey ranged over the old quarry
on a hilltop edge (where an abundant crop of lespedeza, sweet
clover and ragweed furnished an ample food supply), the pond
area at the foot of the same hill, the intervening brushy wooded
south slope, and field cut by a deep gully. This area was roughly
rectangular, approximately 1800 feet long by 1000 feet wide, and its

Fig. 14. Approximate extent of territories of five different pairs of bob-
white arranged along woodland and field border, in the late spring and
' ' early summer of 1955.

computed area was 40 acres. Two or three other coveys ranged
over an area of fallow fields, of 42.8 acres in the northeastern part
of the Reservation, and at least two others in another fallow area
of 48.5 acres in the southeastern and south-central part. A hilltop
field, adjoining the southwestern corner of the Reservation was
headquarters of still another covey, which used the area from time
to time.

In spring, territorial calhng usually has been heard in or near the
field where the Reservation headquarters are situated, but in the
■years when Cooper hawks nested nearby, relatively few quail were
heard or seen there. In 1956, when the hawks were not present.

5—3117

192 University of Kansas Publs., Mus. Nat, Hist.

quail were especially in evidence. In May and early June some
13 calling males were recorded along the edges of this field, each
in a distinct territory, and presumably accompanied by a mate.
The males' habit of answering the calls of neighbors made it possi-
ble to establish the approximate position of territorial boundaries.
Best data were obtained on the male whose territory included the
area around the residence building, as he was seen and heard many
times each day, and he was less wary than other individuals, having
become conditioned to the presence of humans. On various occa-
sions this male was seen, in early morning or after sunset, foraging
with a female. When flushed on such occasions the birds would fly
less than 100 feet, and, staying within the territory, would veer back
to thickets along a ravine, always alighting close together. On most
occasions the male was seen or heard alone, while presumably tlie
female was occupied with laying or incubation. The male spent
much time each day in territorial calling, especially in early morn-
ing, late afternoon, and in cloudy periods. The oval area within
which he stayed was approximately 1.4 acres, but territorial calling
and most foraging was mainly confined to the central part of it.
an area only about 140 feet in diameter. Four intervals between
territorial centers of adjacent calling males averaged 245 feet,
indicating territories of approximately 1.1 acres. All the territories
tended toward elliptical shape. Two, situated where the transition
from woodland to grassland was abrupt, had their long axes along
the edge. Three others that were partly in groves in a field adja-
cent to woodland, had their long axes at right angles to the edge.
Some of these bob-white remained in their territories until late
August, as they were heard calling from time to time. Nevertheless,
territorial calling steadily declined after the end of May, and was
not often heard after midsummer.

Grus canadensis (Linnaeus)

Sandhill Crane

Status. — Uncommon migrant.

Habitat. — Northern prairie and tundra in breeding season.

Movements. — On several occasions migrating groups of cranes
have been seen near the Reservation, but ordinarily they fly high
and do not stop in the vicinity. On the early afternoon of November
13, 1957, a lone crane, which had one leg danghng as if it was

Home Ranges of Vertebrates 193

injured, was seen flying west across the northwestern part of tlie
Reservation. A short time later it was seen flying back in the
opposite direction. In the following week it was seen daily, mak-
ing trips back and forth over approximately the same route, which
crossed the northern part of the Reservation or the southern part
of the Rockefeller Tract. On two occasions it was flushed from the
small pond on the Rockefeller Tract. A few days later feathers,
bone fragments, and the legs of the crane were found strevra at
the pond edge, and along a trail leading off to the south. Probably
the crane had been caught and eaten by a coyote.

Rallus limicola Vieillot

Virginia Rail
Status. — Rare transient.
Habitat. — M arshland.
Movements. — One was seen at the pond on May 8, 1955.

Porzana Carolina (Linnaeus)

Sora Rail

Status. — Occasional transient.

Habitat. — M arsh.

Movements. — Only one record has been obtained to date; on
May 14, 1950, O. M. King and R. W. Fredrickson saw one at the
pond on the Reservation. This individual was almost certainly a
migrant. At the time, marsh vegetation was abundant at the pond.

Fulica americana Gmelin

American Coot

Status. — Occasional transient.

Habitat. — Lakes, marshes and ponds.

Movements. — In April 1950, remains of a coot were found in a
homed owl nest in the southwestern part of the Reservation. This
coot had probably been caught at a small hilltop pond that was
a little less than 200 yards from the nest. At the larger pond, on
the Reservation, coots were recorded on April 22, 1955; May 7,
1951; May 21 to 24, 1953; and October 7, 1955. The pond seemed
to provide favorable habitat with ample food and cover at times
when coots stopped there, but probably was too small to hold
them permanently. Although records are few, their concentration

194 UNRrERSiTY OF Kansas Publs., Mus. Nat. Hist.

in late spring suggests that they pertain to individuals dispersed
from flocks in search of habitat favorable for nesting.

Charadrius vociferus Linnaeus

Killdeer

Status. — Regular transient, chiefly in early spring and autumn.

Habitat. — Typically in or near wet places with open or partly
open expanses of mud, sand or gravel.

In the absence of such habitat on the Reservation, the killdeer
has never been known to stop and all 37 recorded dates pertain to
birds flying over, singly or in pairs or groups.

Movements. — The records are distributed throughout the year
except for the months of December and January, but November 7,
1951, is the latest date in autumn. More than half the records (21)
are for March. Earliest records in each of seven years, 1950 through
1956, were the 4th, 26th, 16th, 5th, 1st, 3rd, and 2nd of that month,
but in 1957 one was heard on February 9 and in 1958 one was heard
on February 26. Probably these numerous early spring records
pertain to birds still on their northward migration, or at least still
wandering in search of nesting territory. In 1958 precipitation was
unusually heavy, and nesting occurred in a partly barren field on
the Rockefeller Tract. Young were fledged. In 1951 also, several
summer records were obtained (June 22, July 28, August 7, Septem-
ber 13). Most records of killdeers on the Reservation were ob-
tained in rainy weather or soon afterward. Killdeers noted in
spring and autumn were usually moving about erratically rather
than holding to a direct course on migration. On October 10, 1951,
at 10 A.M., five were seen in a group, flying south. On October 18,
1951, a group of 12 circled over a hilltop field.

Philohela minor (Gmelin)
American Woodcock

Status. — Rare migrant.

Habitat. — Wet places in or near deciduous forest.

Movements. — Several were recorded in the late summer of 1948,
one on November 4, 1949, and one on August 30, 1951. In the un-
usually wet summer of 1958, woodcocks, probably a pair, were
seen regularly along a 500-foot stretch of woodland edge, in a
brushy, damp, upland area. Probably nesting occurred.

Home Ranges of Vertebrates 195

Capella gallinago (Linnaeus)

Wilson Snipe

Status. — Occasional migrant.
Habitat. — Wet meadows.

Movements. — On several occasions snipe have been seen on the
Reservation, but my only definite record is for October 24, 1951.

Bartramia longicauda (Bechstein)

Upland Plover

Status. — Rare transient.

Habitat. — Prairie, especially of short-grass type, or where heavily
grazed.

Movements. — On May 12, 1957, one was flushed from the blue-
grass pasture on the Rockefeller Tract, It circled and lit in a newly
cultivated field where it remained for more than an hour. When
flushed again it flew back into the pasture. On the following day
it was gone.

Actitis macularia (Linnaeus) ,

Spotted Sandpiper

Status. — Occasional transient.

Habitat. — Shorelines of streams.

Movements. — On May 2, 1949, one was seen at the edge of the

pond.

Tringa solitaria Wilson

Solitary Sandpiper

Status. — Occasional transient.

Habitat. — Edge of water, in a variety of situations.

Movements. — On April 8, 1953, two were foraging along the edge
of the pond, which then contained but little water. On July 14,
1952, one was foraging on mud at the pond edge, late in the after-
noon.

Tetanus flavipes (Gmelin)

Lesser Yellow-legs

Status. — Occasional transient.
Habitat. — Shorelines.

Movements. — The only definite records are of individuals stop-
ping on northward migration in spring. On May 7, 1953, a group

196 Unis'ersity of Kansas Publs., Mus. Nat. Hist.

of six were foraging, wading in shallow water at the edge of the
pond in late afternoon. When flushed, they flew northeast. On
April 2.5, 1956, one feeding at the edge of the pond was seen to
catch a small frog (probably a cricket frog). When flushed, this
bird returned within a few minutes, but after being disturbed a
second time, it circled high over the area and left. On April 27,
1956. one that may have been the same was reported at the pond.

Larus pipixcan Wagler

Franklin Gull

Status. — Occasional transient.
Habitat. — Inland marshes and lakes.

Movements. — On May 2, 1951, late in the afternoon, four were
seen flying high over the pond headed northeast.

Zenaidura macroura (Linnaeus)

Mourning Dove

Status. — Common summer resident; rarely seen in winter but
flocks or lone birds, perhaps from more northern breeding popu-
lations, occasionally cross the area or stop on it briefly.

Habitat. — Most of the area is used by mourning doves for nest-
ing, and nests have been found in a variety of situations. Two
were on the mat of old dead grass in the bluestem prairie of the
northwestern comer. Most others were in trees and shrubs. Park-
land situations at the edge of the woods, and open glades with sumac
patches on south slopes are favored situations for nesting and
foraging. Doves were rarely seen in the denser oak-hickory woods
of north slopes. The fallow fields with their abundant crops of weed
seeds, provide a potential food source, which was much used in
1948 and 1949. Since 1949 these fields have been Httle used, be-
cause the accumulated dead vegetation makes foraging difficult
for the doves.

Movements. — The flocks of mourning doves that are occasionally
seen in Douglas County in winter tend to keep to cultivated land,
and have rarely been seen on the Reservation. On mild days of
early spring, doves appear abruptly and begin territorial calling.
These probably are new arrivals that have spent the winter farther
south. First recorded dates of territorial calling each year from
1950 through 1957 were: March 17, March 26, March 28, March

Home Ranges of Vertebrates 197

12, March 7, April 15, March 6 and March 2. Soon after arrival
they begin to nest. Each year doves nested in the vicinity of the
headquarters buildings, and on many other parts of the Reservation.

Males moving about and calling from territorial perches have
been seen to use stations separated by as much as 700 feet. Others
kept under observation for periods of hours, have stayed within an
area 100 feet in greatest diameter. In the vicinity of the Reserva-
tion headquarters, distances between adjacent calling males in
seven instances averaged 689 feet (1300 to 100). If this figure is
representative of a territory's diameter, nesting territories averaging
approximately eight acres are indicated.

Only rarely were doves seen to come to the ground to forage on
the Reservation, On the contrary, those that bred on the area
seemed to feed on other areas remote from their territories. Doves
that nested near the Reservation headquarters were often seen
travelling in strong and direct flight to the west for half a mile or
more, to cultivated fields, or were seen returning from such flights,
to nests, territorial perches or feeding of young.

A typical instance of territorial defense was recorded on April
20, 1954. A pair engaged in courtship in an elm tree beside the
house. After a period of preening, three feet apart on a horizontal
hmb, they showed increasing awareness of each others' presence.
One, presumably the female, postured with tail elevated and wings
vibrating. The mate approached, and standing behind, pecked
lightly at the other several times, then flew to another tree 50 feet
south. The female followed. At this stage proceedings were inter-
rupted when a second male began calling in another elm 100 feet
farther south. He soon flew to telephone wires 150 feet west of
the pair, and resumed calling. The male of the pair flew and
perched on the wire beside him, and several times in succession
lunged at him, causing him to fly a short distance. The intruding
male moved to a nearby tree, and the female joined her mate on
the wire. The pair followed the intruder, perching on the opposite
side of the tree. After about ten minutes the unmated male, then
the mated male, and finally the female flew to still another tree.
Soon all three flushed again, travelling in rapid and erratic flight,
the unmated bird closely pursued and harrassed by the mated male,
while the female followed farther behind. The pursuit led for sev-
eral hundred yards over a wooded north slope, and the outcome
was not seen.

198 Uxix-ERSiTi' OF Kansas Publs., Mus. Nat. Hist.

Nests were found each year from April through September. Few
were successful, and most that I saw were destroyed by predators
within a few days of their discov^ery. Often re-nesting, presumably
by the same pair of doves, occurred promptly within a few yards
of the site where a nest was destroyed. Individuals were not
recognized.

Coccyzus americanus (Linnaeus)
Yellow-billed Cuckoo
Status. — Common summer resident.

Habitat. — Cuckoos utilize all types of woodland present on the
area, but evidently prefer edge situations, where there are thickets.
Nests often are situated in thorny trees or shrubs such as osage
orange or honey locust. Territories may include extensive open
areas, but these are crossed in rapid direct flight and most of the
time cuckoos keep out of sight in dense foliage.

Movements. — Each year cuckoos have been first noted on the
Reservation in May; the 15th (1950), 24th (1951), 17th (1952),
21st (1953), 3rd (1955), 12th (1956), and 9th (1957). Within a
few days after recording of the earliest arrival, cuckoos are present
in abundance and are establishing breeding territories. Nests have
been found from May through September, but with a high propor-
tion in the latter half of the summer, suggesting that more than one
brood may be the rule. Most of the nests found were destroyed
within a few days by predators, and some were blown dowTi in
storms. Because cuckoos tend to be silent after late August, and
are somewhat secretive at all times, their departure in autumn
usually passes unnoticed. On several occasions they have been re-
corded in the first week of October, but none has been seen later
in the season.

Each year throughout the summer cuckoos were present in the
headquarters area and were seen and heard daily. Territorial call-
ing of each male tended to be concentrated in one or a few favorite
perch trees. In July, 1955, a territory encompassing the grove of
elms and other large trees in the headquarters area was approxi-
mately 250 feet in diameter and roughly circular.

As cuckoos tended to be secretive, keeping out of sight in dense
foliage, their movements could not be readily traced. The spacing
of calling males that answered each other provided some indication

Home Ranges of Vertebrates 199

of the size of areas occupied. In 17 such instances the interval
averaged 615 feet ( 160 to 1000 ) . On 13 other occasions linear dis-
tance covered was measured for individuals that were kept under
observation for considerable periods, or that were seen to make
relatively long direct flights. These distances averaged 323 feet
(140 to 650), with more than half concentrated between 200 and
310 feet. A typical territory of a yellow-billed cuckoo might be
approximately six acres.

On August 13, 1956, a cuckoo was watched while it was occupied
with nest-building. It flew into a large elm in which one main limb
was dead and devoid of foliage. It broke off a twig approximately
five inches long, and carrying the twig crosswise in its bill, flew
back to the nest tree. It paused, then made its way through thick
foliage to the partly built nest situated in a horizontal crotch. As
it passed through the thicker part of the tree, its movements were
impeded by the transverse stick catching on twigs and leaves. In
the next half hour it repeated the procedure time after time with
but little variation. It always went to about the same spot on the
dead branch, about 60 feet from the nest.

Brumwell (1951:220) at Fort Leavenworth, found the cotton-
wood-elm association along the Missouri River to be a favored
nesting habitat. He observed several adults fly across the river and
return with food for the young, making a round trip of approxi-
mately a mile.

Coccyzus erythropthalmus (Wilson)
Black-billed Cuckoo
Status. — Occasional transient and summer resident.
Habitat. — Deciduous forest and edge. The relative rarity of this
species on the area suggests that its ecological tolerances are nar-
rower than those of the yellow-billed cuckoo, and perhaps it is
limited to habitats providing better cover.

Movements. — On September 6, 1951, one that was flushed in thick
woods of elm, locust and osage orange, on a south slope, was carry-
ing a caterpillar in its bill. It uttered rabbit-like squalling sounds
and was joined by a mate. After their excitement had subsided
somewhat, they gave frog-like rolling notes at intervals of a few
seconds. A nest was nearby, about 13 feet above the ground.
Others were recorded on the Reservation on May 17 and 22, 1952,
and on September 24, 1957.

200 Uni\'ersity of Kansas Publs., Mus. Nat. Hist.

Tyto alba (Scopoli)
Bam Owl

Stottis. — Occasional transient.

Habitat. — Typically, grassland with nesting sites such as are
provided by caves, hollow trees, old buildings, mine shafts, or
niches in gully cutbanks.

Movements. — On December 7, 1949, a pair was flushed from a
dense thicket at the edge of an eroded, fallow upland field. After
being flushed both owls flew down a ravine and were heckled by
small birds. A few minutes later one of the owls flew over, about
100 feet high and it veered, suddenly spiralling downward to the
same place from which it originally flushed. On December 1, 1951,
one was flushed on a wooded slope 200 yards east of the Reserva-
tion headquarters. No other bain owls have been seen on the Res-
ervation but occasionally at night their calls have been heard. On
February 7, 1952, at about 9 P. M., one was heard calling and from
these calls its flight could be traced along the narrow, flat, grassy hill-
top that extends across the Reservation. Those living on nearby
farmlands may occasionally wander on the Reservation in the course
of their hunting. For the barn owl, habitat conditions on the Res-
ervation have worsened since areas that were formerly grazed or
were cultivated fields, have become covered with a dense stand of
rank vegetation.

Otus asio (Linnaeus)

Screech Owl

Status. — Probably regular resident, but records were not obtained
with sufficient regularity to indicate that area was occupied by
permanent population.

Habitat. — Woodland and parkland.

Movements. — Twelve records in eight different months were
obtained in 1951, all within a radius of 100 yards, and probably
all pertaining to the same pair. Activity seemed to center on a
wooded north slope and the ravine along its bottom, south of the
Reservation headquarters. On March 17 one of the owls was
found sleeping in the hollow trunk of an oak near the center of
the area where calling had been heard. On several occasions two
owls were heard calling and answering each other. Screech owls
were heard also in 1950 (once), 1952 (once), 1954 (twice), 1955

Home Ranges of Vektebrates 201

(three times), 1956 (five times), and 1957 (once). All but two
of these records were within the area encompassed by the 1951
records.

Bubo virginianus (Gmelin)

Horned Owl

Status. — Abundant (for a large raptor) resident, using nearly all
parts of the Reservation.

Habitat. — By day usually seen in dense woods where roosts are
located, but at night it is less restricted, ranging far and wide,
through woodland, woodland edge, and open fields.

Roosts are usually in large trees such as elms, oaks or honey
locusts, especially where thick branches or vine tangles provide
concealment.

Movements. — Ordinarily homed owls are territorial. Each ter-
ritory is occupied by a pair of adults. Territories are occupied
throughout the year, and perhaps continuously for long periods of
years. Most information concerning territories was obtained in
the months of November, December, and January because the owls
were especially vocal then. Their hooting would begin a few
minutes before sunset and might continue sporadically throughout
the night. However, the most concentrated vocalization occurred
in a period of perhaps 15 to 20 minutes after sunset. The first owl
to hoot would usually be answered, after it had called a few times,
by its mate or by a rival on an adjacent territory. Soon still others
would begin calling until as many as six pairs could be heard from
one vantage point.

In late 1955, approximate centers were located for eight terri-
tories, six of which were on the Reservation, while the other two
were near the boundary. For 13 instances of territories adjacent
to each other, with continuously favorable habitat, the distances
intervening between territorial centers ranged from 3070 to 1700
feet ( average 2400 ) . In 23 instances of owls answering each other
from adjacent rival territories, the intervening distances ranged
from 2360 to 686 feet (average 1360).

These figures represent conditions obtaining when the popula-
tion was high. The number of territories on the Reservation varied
somewhat from year to year. No figures are available for 1948 or
1949, but certainly the owls were moderately abundant in those
years. In the winter of 1950-51, eight territories were represented

202 Unin'ersity of Kansas Publs., Mus. Nat. Hist.

on the Reservation. Three of these were almost or entirely confined
to the area, and each of the other five overlapped it extensively.
In the winter of 1951-52, there were nine territories represented.
In general, they corresponded with the territories of the previous
year, but with some shifting of boundaries and centers of activity.
The increase in numbers of owls on the area did not represent a
higher population density. In 1952-53 only six territories were
known to be represented on the area. In correlation with this
decrease, the territories were exceptionally large. The best known
territory, which was known over the six-year period, reached its
greatest extent that winter; it was .6 mile long and .42 mile wide.
Baumgartner (1939:280) recorded three or four male horned owls
per square mile in favorable habitat south of Lawrence. On a five-
square-mile area he recorded an average of two males per square
mile.

Low population density of the owls was probably correlated with
deficient rainfall and, more directly, with the low population of
cottontails. In the summer of 1952 vegetation was relatively sparse
because of drought. Having less food and cover than usual, cotton-
tails did not increase normally in the breeding season. Only
meager data concerning the numbers and distribution of owls were
obtained in the winters of 1953-54 and 1954-55, but there were
probably seven or eight territories in those years. In the winter
of 1955-56 there were eight territories. Only one of these was
entirely within the Reservation, but each of four others had more
than half of its extent within the area.

On many occasions the presence of horned owls was revealed
to the observer by the clamoring of crows, which heckled and
pursued them on sight. Especially in winter, when the population
of crows was high, and trees were bare, so that concealment was
more difficult, crows spent many hours each day heckling the owls
on the Reservation. An owl might be followed and harrassed for
an hour or more at a time, and hundreds of crows might congregate.
The owl might flush a dozen times or more before ridding itself of
the crows. When it flew, excitement increased, as the crows pursued
it. Harrassment by crows must be an important factor in the ecology
of the horned owl in this region. On February 24, 1953, the course
of an owl was recorded for about an hour that it was followed by
crows. It flushed six times, with flights of 100 yards to a quarter
mile, travelled a linear distance of .9 mile, and was finally left by
the crows when it was a little more than .6 mile south of the starting
place.

Home Ranges of Vertebrates 203

From time to time small accumulations of pellets were found
where horned owls had roosted during the day. Such accumula-
tions rarely consisted of more than a dozen pellets, and it was ob-
vious that the owls of each pair changed roosts frequently. Never-
theless certain favored trees were returned to repeatedly and had
new accumulations of pellets on many occasions over periods of
years. The roosting area is only a small part of the territory, gener-
ally 100 yards or less in diameter. The members of a pair have
never, in my experience, been flushed from the same tree, but are
usually in nearby trees where each may be seen by the other.
Upon leaving the roost at dusk an owl may fly to a distant part of
its territory, followed by its mate. After dark the owls may move
away from the woodland into relatively open situations, perching on
fence posts, snags or isolated trees that command a view of the
surroundings. Many of the owl kills found have been in fields or
pastures, sometimes hundreds of feet from the edge of the woods.

In 1950 a nest was observed from time to time throughout incuba-
tion, and subsequent development of the young owls. Both young
were banded. The nest was in a honey locust in a hilltop field.
After the young had left the nest, they were found perched close
together in the edge of the woods about 100 yards from the nest and
about 15 feet above the ground. In subsequent weeks these young
were seen frequently. At first they stayed within a small area and
tended to stay close together. Gradually they shifted in roosting
from the southern to the northern part of the territory and even
beyond it, and they were no longer found together. Through tlie
spring and early summer these young presumably were still being
fed by the adults as their hunger calls were heard often at night,
but the adults were never with them when they were flushed in the
daytime. Five years later one of these banded young was shot by
a farmer at Tonganoxie, some five miles from its parents' territory.

Strix varia Barton

Barred Owl

Status. — Resident; one pair believed to have been in continuous
tenure of territory from 1949 through 1957, and others have been
present on various parts of Reservation from time to time.

Habitat. — Deciduous forest.

The relatively small portion of the Reservation where a pair of
these owls stayed, differed from other woodland areas in more nearly
approximating the Oak-Hickory climax, as it had many large, ma-
ture trees, with a high proportion of red oak, chestnut oak, black

204 University of Kansas Publs., Mus. Nat. Hist.

oak and shagbark hickory. The numerous hollow trees provided
nesting sites and perhaps daytime roosts for the owls. This was
at the head of a small valley, where two deep ravines joined, near
wooded slopes, and a brushy bottomland. In other woodland and
field-edge situations where barred owls were seen from time to
time, they did not remain for long.

Movements. — The pair of owls that lived near the eastern edge
of the Reservation were rarely seen but were heard hooting on
numerous occasions, on overcast afternoons or at dusk. My imita-
tion of their calls often caused them to begin hooting. They were
almost always heard when the territory was visited after dark.

Most of their hooting was heard witliin an area estimated to be
only 200 yards in diameter. However, these records mostly pertain
to daytime or dusk, while the owls were on or near their diurnal
roosts, and therefore are not adequately representative of the entire
territory. The more dispersed records indicate a territory of ap-
proximately a quarter-mile in diameter.

When these owls were caused to hoot, by an imitation of their
calls, the response of the first individual was usually followed within
a few seconds by that of its mate. The calls of the two individuals
came from different points, but within the same general area, evi-
dence that they normally roosted in separate trees. Sometimes the
calling was taken up by other barred owls in woods east of the
Reservation boundary, at a distance in the neighborhood of half
a mile from the resident pair.

More detailed information concerning movements was obtained
in the vicinity of the Reservation headquarters, where a barred owl
— probably the same individual — occurred over periods of months,
alone at first, and later accompanied by a mate. Early records of
it were obtained on October 12 and 17, 1950, and on March 5, May
26, and November 13, 1951. This owl may have had a territory
farther north and east; if so it only occasionally ranged into the
area. However, it was seen more frequently in the autumn and
early winter of late 1952. Subsequently, in January, 1953, it
seemed to have shifted so that its territory centered near the Res-
ervation headquarters. Thenceforth, throughout 1953, it was re-
corded many times each month. From mid-May to mid-October
it was accompanied by a mate.

In winter, its activity was largely diurnal. On many occasions
it was seen hunting, and sometimes was kept under observation for
periods of hours. Voles seemed to be the chief food source, and

Home Ranges of Vertebrates

205

in winter their activity likewise tended to be limited to the daylight
hours. In hunting, the owl did not stay in the woods, but chose
trees in meadows or that bordered on them. Flushing distance
was usually 150 to 200 feet but sometimes the owl permitted ap-
proach to within 60 feet. When not disturbed, the hunting owl
usually remained on its perch from a few seconds up to fifteen
minutes. While on such a perch, it was alertly scanning its sur-
roundings with almost continual, short, quick movements of its
head. From time to time it would fly down into the grass in pursuit
of prey, and sometimes would remain as much as a minute. Often
it flew up with no evidence of a catch even after it had been on the
ground for as much as half a minute. It was seen to fly distances

Fig. 15. Northwestern quarter of the Reservation showing territory of a
pair of barred owls in 1953. Dots show locations where owls were heard
hooting. Broken lines show course of an owl that was followed as it hunted
in daylight on January 8 and 19. The circuit shown for January 8 was fol-
lowed three times in succession, with minor variations.

206 Universitti' of Kansas Publs., Mus. Nat. Hist.

as much as 100 feet to strike at prey, but usually the flight to the
ground was shorter. In moving from one perch to another it
sometimes flew only a few feet, rarely as much as 100 yards. The
flight seemed remarkably slow. On the difi^erent occasions when
it was followed, it covered more or less the same area, and returned
to many of the same perch trees, although not using them in any
definite sequence. The area that it covered was of irregular out-
line, approximately 1900 feet long and 1200 feet wide. Its diurnal
activity in winter was largely between dawn and 10:30 A. M. It
was occasionally seen in bright sunlight, but seemed to prefer
overcast days. On numerous occasions it was found later in the
day, inactive on a roost. It liad no special roosting place, but was
usually in some large tree with dense foliage, near the central part
of the territory.

After this owl obtained a mate, hooting was heard much more
frequently. Often the owls began hooting late in the afternoon,
answering each other while still on their roosts. They were usually
near together, but over periods of weeks their roosting areas
shifted until almost every part of the territory had been utilized
for this purpose. In fact the area encompassed by their perches
was considerably greater than that otherwise known for them; it
was bounded by seven marginal points, and was measured as 75
acres. The territory of these owls was separated by at least half
a mile from that of their nearest neighbors, the pair already dis-
cussed, that occupied an area near the eastern edge of the Reser-
vation. Obviously territorial pressure was not a factor limiting the
size of either territory.

Asio otus (Linnaeus)
Long-eared Owl
Status. — Frequent winter resident; occasional in summer.

Habitat. — For roosting this owl prefers dense thickets, and locally
most often uses clumps of red cedar which offer better protection
than bare deciduous trees. On the Reservation no large cedar
trees are present, and the owls have usually been found roosting
in thickets of osage orange, honey locust, or chinquapin oak. In
their nocturnal foraging, on the Reservation at least, these owls
most often seem to hunt over grassland, as analysis of pellets has
shown the prey to consist largely of prairie voles, cotton rats, and
harvest mice.

Home Ranges of Vertebrates 207

Movements.— On October 11, 1950, two were found in a thicket
of chinquapin oak at a hilltop edge. They flew about 100 feet and
lit in another thicket. On December 7, 1951, one was flushed from
a hilltop rock outcrop. On January 8, 1952, a group of three were
flushed from an osage orange tree on a wooded south slope. On
March 10, 1952, about 200 feet north of this spot, a group of at
least a dozen was flushed from within a 20-foot radius in a thicket
of chinquapin oak. Ground beneath the tliicket was littered witli
pellets and their fragments, some of which appeared to be many
weeks old. Later in the day the owls were located again, mostly to-
gether in an osage orange tree about 100 yards south of their regular
roost. At my approach they flushed one by one and headed back
in the direction of their former roost. White-footed mice and other
prey that might have been secured in the forest made up but a smaU
percentage of the items in the pellets that were examined; voles and
cotton rats that must have been obtained in open fields, made up
the bulk of the diet.

On January 22, 1954, one was flushed from a roost among project-
ing roots in the cut-bank of a gully. On February 2, 1954, strewn
feathers of one were found in the middle of a hilltop field, and it
seemed the owl had been attacked and eaten by another predator.
On March 23 and 26, 1954, a pair was seen near the same place in a
thicket on a brushy south slope.

On a December evening when it was nearly dark, a flock was
observed hunting over a hilltop field. They paid little attention to
my presence, but were wheeling and circling in a fairly compact
group within a few yards of the ground.

On April 19, 1951, a quarter mile southeast of the Reservation,
one was found incubating in a nest 20 feet up in a honey locust.
The only summer record from the Reservation is of one seen on
August 10, 1951.

Aegolius acadicus (Gmelin)

Saw-whet Owl

Status. — Rare transient in winter.

Habitat. — Breeds in northern coniferous forests but moves south
in winter and may appear in variety of habitats.

Movements. — Only two have been definitely recorded from the
Reservation. In December, 1948, one was seen on a wooded hill-
top. It was tame at first, but became more wary and flew farther

20S Untvebsity of Kansas Publs., Mus. Nat. Hist.

at each attempt to stalk it, until it was lost. One perched ten feet
above the ground in an osage orange tree on February 11, 1951,
at 1 P. M. was in the act of swallowing a white-footed mouse. It
permitted approach to within fifteen feet. Twice when it was
flushed, it moved only a few yards before settling on another low
perch, but it continued to watch my movements alertly.

Caprimulgus carolinensis GmeHn
Chuck-will's-widow

Status. — Irregular summer resident.

Habitat. — Woodland.

On the Reservation activity was concentrated on areas of south
slopes with medium to small trees and dense thorny undergrowth,
a type of woodland distinctly more xeric than that preferred by
the whip-poor-will.

Movements. — The first definite record was obtained on May 11,
1953, when one was heard calling on the slope north of the head-
quarters. All other records were obtained in 1956 and in 1957,
when the species was present throughout the late spring and early
summer and was heard almost nightly. One calling on April 27,
1956, was answered regularly by a whip-poor-will in the same
general area. In subsequent weeks whip-poor-wills were heard
less regularly than they had been in other summers and perhaps
were, in part, displaced by the chuck-will's-widows. On May 17
one of the latter began calling at 7:36 P. M., and soon a second
began answering nearby. I approached to within 200 feet and
heard a peculiar low growling sound, then saw one in close pursuit
of the other in an erratic flight through the trees. They settled and
resumed calling only about 50 feet apart and a third was calling
on a hillside 150 yards away. Another pursuit, accompanied by the
same peculiar growling sounds, was noted on June 6, as one chased
a rival from one woodland edge to another, across intervening fields,
a distance of some 320 yards. The pursuer quickly returned, and
both chuck-will's-widows called, answering each other.

A chuck-will's-widow engaged in territorial calling as it shifted
from perch to perch, moved, on the average, approximately 430
feet (640 to 240) at each shift. A supposed territory mapped on
the basis of 16 stations covered 23.7 acres, but there is some doubt
as to whether all the records pertained to the same individual.

An unusually late fall record of a goatsucker that appeared to be
of this species was obtained on December 6, 1953, when the bird

Home Ranges of Vertebrates 209

was flushed at close range, from the ground or a low branch. It
flew about erratically and lit in a large tree but could not be re-
located. Temperature was slightly above 50° F. at the time it was
seen, but light snow had fallen on the preceding night.

Caprimulgus vociferus Wilson

Whip-poor-will

Status.— Common summer resident, using most parts of Reser-
vation, but chiefly confined to woodland and vicinity of woodland
edge.

Habitat. — Woodland, especially that of more mesic types where
there are oaks, hickories, and elms.

Whip-poor-wills flushed from their daytime roosts were in dense
woods where the leaf litter was thick, and where there was under-
brush such as dogwood and gooseberry. Calling males at night
were frequently noted in edge or parkland situations.

Movements. — Whip-poor-wills arrived in April each year; from
1950 through 1956, earliest records were the 6th, 24th, 24th, 22nd,
10th, 17th and 20th. Late records include: October 11, 1949; Oc-
tober 1, 1950; September 23, 1951; September 16, 1952; September 8,
1953; September 20, 1954 and September 20, 1955. Throughout
May, June and July territorial calling is persistent, beginning at
dusk and continuing intermittently until daybreak. This "song,"
the familiar "whip-poor-will" call, serves to maintain spacing be-
tween pairs on adjoining territories. Open spots are chosen for
calling. The roofs of the house and laboratory buildings and top
of the chimney were especially favored, as were the top of the dike
at a dam, and the bare shaly bank of a ditch. In other instances the
birds perched on horizontal branches of large dead trees while they
were calling. On several occasions when a whip-poor-will was call-
ing, I attempted an imitation. If the whip-poor-will was nearby,
this imitation usually caused a strong reaction. The bird might
hover erratically, coming within a few yards of me and on one oc-
casion, one lit on the ground only 15 feet away, giving a peculiar
purring sound. This same sound has been heard while one in-
dividual was chasing another, and immediately afterward. Excep-
tionally favorable sites for territorial calling may be used by different
individuals at different times, neither of the two (or more?) being
able to establish absolute control.

In 24 instances the distance between whip-poor-wills that were
calling and answering each other, ranged from 1600 feet to 370 feet.

210 UNivERSiTi' OF Kansas Publs., NIus. Nat. Hist.

averaging 890. If this distance represents the diameter of a typical
territory, the actual size amounts to some 22.6 acres. Territories ac-
tually measured insofar as they were known, covered 14.9 acres ( 18
stations, 4 marginal), 27.5 acres (28 stations, 6 marginal), and 6.9
acres (13 stations, 6 marginal).

On June 10, 1955, a whip-poor-will flushed from leaf Litter in oak-
hickory woods on a northwest slope feigned injury. On the fol-
lowing day it was flushed again, and the two eggs were found at the
spot where it had been sitting. On June 17, the bird was flushed
again, and the eggs were approximately twenty feet from the spot
where diey had been before. On June 21, when the nest was re-
visited, the parent flushed from a spot ten feet away. The eggs
were at the same spot where they had been on June 17, and the
parent may have crept away from them before making herself con-
spicuous. Again she feigned injury, jumping and flopping, moving
a foot or two at a time until the sixth leap when she flew several
yards, and at the next start she flew out of sight, but soon reap-
peared, approaching the nest from another direction.

Bent (1940:149) mentions instances of the closely related chuck-
will's-widow moving its eggs after disturbance of the nest, and
quotes Audubon as stating that the eggs are carried in the mouth.
Bent (op. cit. :175) also cites instances of female whip-poor-wills
flushed from the nest carrying nesthngs for short distances grasped
between her legs.

Chordeiles minor (Forster)

Nighthawk

Status. — Regular summer visitant; no evidence of nesting on the
area has as yet been obtained.

Habitat. — The prey consists of flying insects, and where these are
numerous the nighthawk may be present, in any of a variety of
terrestrial habitats. The limiting factors are thus not so readily de-
finable in terms of plant associations or physiography as in most
other kinds of birds.

Movements. — Earliest records for the Reservation are May 18,
1952, May 22, 1954, and May 15, 1955. Latest records are Septem-
ber 21, 1948, September 20, 1953, September 20, 1954, and Septem-
ber 11, 1955. On September 18, 1954, soon after sunset, a group of
more than a dozen, flying erratically on separate courses, passed
over the field where the headquarters are located, and gradually
moved away to the southwest imtil they were out of sight. Ap-

Home Ranges of Vertebrates 211

proximately a quarter hour later, a similar flight occurred, perhaps
of the same individuals. In 1952 nighthawks were often seen for-
aging over the Reservation in late afternoon, and several times
were flushed from woodland in the daytime. In later years, while
drought conditions prevailed, they were rarely seen on the area
between May and September. Foraging nighthawks that were fly-
ing over the area covered ranges much larger than those of other
birds of comparable size. Their ranges seemed to be more than
half a mile in diameter, and, although some tendency was noticed
for an individual to make the same circuit several times in succes-
sion, they were not present consistently over periods of days or
weeks suggesting that they wandered extensively, or that the areas
they were seen to cover comprised only small parts of their total
ranges.

Chaetura pelagica (Linnaeus)

Chimney Swift

Status. — Usually present in summer but not known to breed;
those that appear from time to time evidently have large foraging
ranges that include parts of the Reservation, or possibly all of it.

Habitat. — Aerial maneuvers of the individuals that are seen on
the Reservation have little or no relation to the terrain or plant
associations beneath. The chimney of the residence was an
obvious focal point of activity, though the swifts were never seen
to enter or emerge.

Movements. — Over the eight-year period, 1950 through 1957,
chimney swifts were first recorded on the Reservation on the fol-
lowing dates: May 20, May 18, May 23, May 24, May 8, May 17,
April 28 and April 24. In several years they were noted somewhat
earlier within the Lawrence city limits than they were at the Res-
ervation: May 8, 1953, May 5, 1954, and May 1, 1955. Most rec-
ords for the Reservation are for late April, May, June, and July
(latest September 24, 1957). Swifts may have nested in the
chimney at the Reservation headquarters. At least, this chimney
was the focal point about which their maneuvers centered. Maneu-
vering was seen chiefly near dusk and on overcast days. On June
24, 1952, two pairs were flying in the headquarters area. Those
of each pair kept close together in their rapid flight, but seemed
to pay no attention to the other pair. They were flying to dis-
tances of hundreds of yards in each direction from the chimney,
about which their maneuvers centered. On May 25, 1953, two

212 University of Kansas Publs., Mus. Nat. Hist.

seemed to be chasing a third, while a fourth was flying alone. On
many other occasions, especially in early June, 1953, and in June,
1955, three were flying together in a group. On June 9, 1953,
there were two such groups. On May 8, 1954, the first time that
swifts were seen that spring, a pair was moving in rapid flight
about the chimney, in circles about 200 feet in diameter, from time
to time descending to the chimney in tight spirals, but not entering
it. On May 17, 1955, the first swifts noted that season were heard
chattering and then a moment later, were heard fluttering inside the
chimney.

Archilochus colubris (Linnaeus)

Ruby-throated Hummingbird

Status. — Uncommon summer visitant or transient.

Habitat. — Less limited to specific plant associations and habitat
situations than most other species of birds occurring locally. The
limiting factor in most instances is an abundance of suitable flowers
providing nectar. Hummingbirds shift their foraging areas fre-
quently according to the availability of the food supply at difiFerent
stages in the season. Germander (Teucrium canadense) is a
favorite source of nectar. This mint grew in dense patches in some
parts of the former pastures for several years after grazing was dis-
continued. It decreased in abundance and was replaced by grasses.

Movements. — Hummingbirds were generally seen only at times
and places where there were abundant supplies of flowers, and may
not have nested on the Reservation. Redbud and germander were
two of the favorite kinds of blossoms. A hummingbird seen on
May 18, 1952, was feeding from dandelion flowers. After 1953, as
succession progressed, with germander and other kinds of flowering
plants being replaced by grasses, hummingbirds became much
scarcer on the Reservation.

Megaceiyle alcyon (Linnaeus)

Belted Kingfisher
Status. — Occasional transient.

Habitat. — Vicinity of ponds and streams, requiring vertical cut-
banks for excavation of nest burrows.

Movements. — Kingfishers, either lone individuals or pairs, ap-
peared from time to time at the pond on the Reservation. In each
instance probably the birds were migrants, which happened to
notice the pond in flying over, and stopped there temporarily. Fish

Home Ranges of Vertebrates 213

were not present, and amphibians comprised the chief sources of
food. Specific dates that kingfishers were seen at the pond were:
July 8, 1950; April 7, 1951; May 19, 1951 (pair); February 17, 1952
(may have been one of a pair seen frequently along a slough two
miles south of the Reservation); April 16, 23, and 24, 1952; May
28, 1952; April 5, 13, and 29, 1953; March 31, 1955; April 22, 1955;
June 21, 1955; April 8, 1956, and May 3 and 4, September 18, and
October 4, 1957.

Upon leaving the pond kingfishers sometimes flew in a low
direct course as if headed for a predetermined goal. At other times
they circled over the area, gradually rising until almost out of sight.

Colaptes auratus (Linnaeus)
Yellow-shafted Flicker

Status. — Common winter resident and migrant.

Habitat. — ^Woodland, but preferably where alternating wdth open
situations such as grassy pastures.

Every part of the Reservation was utilized by flickers, but with
temporary concentration on certain habitats, depending on tlie
season, the weather, and availability of food sources.

Movements. — Flickers have not been recorded on the area from
about mid-April to mid-September; generally they are absent in
late spring and summer. The area is within the breeding range,
and failure of the species to remain through the summer may re-
sult from the lack of suitably open situations where flickers might
forage on the ground. Because the area is not grazed, or otherwise
disturbed, the parts that were formerly open have grown up to
thickets of young trees and shrubs, rank weeds, or tall grass.

First recorded appearances of flickers in fall of several different
years were: September 28, 1948; September 13, 1951; September 12,
1952; September 19, 1953; September 17, 1954; October 5, 1955;
September 21, 1956, and September 18, 1957. In late September
and early October numbers on the area were rapidly augmented
by new arrivals, and by late October flickers were among the most
conspicuous birds of the Reservation. However, unlike most other
wintering birds on the Reservation, individual flickers did not seem
to become attached to specific areas for more than a few days at a
time but were roving about v^dth constant fluctuations in the num-
bers present.

On October 5, 1953, after one of the first cold nights of the season,
many flickers were foraging on the ground near the house in com-

214 Univ-ersity of Kansas Publs., Mus. Nat. Hist.

pany with red-bellied woodpeckers and robins, finding grasshoppers
and other insects that were immobilized by the cold and were easily
obtained in the short vegetation. By 8:30 A. M., this group had
dispersed. In autumn, foraging is to a large extent arboreal, and
wild fruits such as those of grape and poison ivy are major food
sources. In winter relatively few remain, but from time to time
there is an influx of migrants. At times when the ground was cov-
ered with snow and ice, flickers often concentrated their foraging
on the cut-banks of gullies, digging in the loose soil for insects. On
warm days after heavy snowfall, weedy fields were favorite places
for foraging. There, the snow melted rapidly, being held oflF the
ground by the mat of dead vegetation, and the flickers were able
to find areas of exposed soil for digging.

Centurus carolinus (Linnaeus)

Red-bellied Woodpecker

Status. — Common resident; no seasonal shifts are apparent.

Habitat. — Woodland, with perhaps some preference for oak-
hickory type.

The main requirement seems to be the presence of large old trees
with numerous dead limbs and cavities. It matters little whether
the tree happens to be cottonwood, elm, oak, or honey locust. The
woodpeckers have been observed to utilize each of these kinds as
the headquarters of a family group or an individual. Such groups
or individuals are usually well scattered, with intervening wood-
land, often hundreds of yards in extent, that is little used or entirely
unoccupied.

Movements. — Red-bellied woodpeckers were observed to be no-
tably sedentary. Large old trees, including elms, oaks, cottonwoods
and honey locusts, were favorite foraging places, and an individual
might spend a disproportionately large part of its time in one or
several of these trees. At all times of year woodpeckers of this
species are somewhat social, and two or more may be observed
in the same tree. However, the bond between members of a pair
in winter is weak if it exists at all. At that time each individual
seems to be largely independent although it may at times asso-
ciate with another, or even with woodpeckers of different species.
In this respect the species differs from the hairy woodpecker, in
which a strong bond between members of a pair exists, even at
times of year other than the breeding season.

Home Ranges of Vertebrates 215

Each year, in warm clear days of early spring, males gave terri-
torial calls from high snags. First records of such territorial calling,
for several different years, were: March 24, 1951; March 28, 1952;
March 13, 1953; February 1, 1954 ( subsequently calling was heard
often through February); March 11, 1955; March 30, 1956; February
16, 1957; and February 22, 1958. Often two males on snags hun-
dreds of feet apart would answer each other regularly with the
"churring" territorial calls. Also, territorial quarrels were frequent,
and were notable for the persistence with which the intruder re-
mained within the territory of his opponent or returned to it. A
typical encounter between two males was observed on February
8, 1954, in a large honey locust in the edge of the woods, between
the headquarters and the pond. The two males, clinging upright
on opposite sides of a vertical branch, would engage in rapid spar-
ring, uttering loud rolling calls every few seconds and also low
grinding sounds. One would fly northeast 100 yards or more,
chased by the other. In a few seconds the pursuer would return
and perch on top of the snag, giving long rolling calls, but soon the
second would return and renew the contest. While quarreling, the
males drummed from time to time. A female, probably the mate
of the defender, was also present in the tree, but she remained aloof
and passive. A fourth woodpecker, possibly the mate of the in-
truder, was calling occasionally near the pond.

On many other occasions similar performances were observed.
Probably they involved former co-occupants of a small area of
favorable habitat, who became increasingly intolerant of each other
and finally excluded each other from their territories as the breed-
ing season approached.

Quarreling red-bellied woodpeckers strutted, bowed, postured
and sparred, often moving about from tree to tree; the one that
flew was immediately followed by the other. A quarrel might con-
tinue intermittently for hours, and might be resumed on successive
days.

In their foraging, red-bellied woodpeckers seem to have an aflBn-
ity for other kinds including hairy and downy, and often are ob-
served in the same tree with one or both of the two last named. The
abundant supply of insects probably attracts all three species, but
also there seems to be a tendency for such a group to maintain
itself in shifts from one tree to another. However, the nesting snags
are jealously guarded against other woodpeckers, and against other
kinds of birds that use cavities. In autumn the influx of flickers

216 U>avERSiTY OF Kansas Publs., Mus. Nat. Hist.

caused much uneasiness among the red-bellied woodpeckers that
lived near the Reservation headquarters. The larger and more
aggressive flickers moving in numbers through and near the fa-
vorite trees of the red-bellied woodpeckers, caused the latter to
scold and to make frequent short, restless movements. The red-
bellied woodpeckers seemed to be incompatible with the red-headed
woodpeckers, as the former were hardly ever seen or heard in an
area at the east edge of the Reservation occupied by a colony of tlie
latter, and from time to time when red-headed woodpeckers tem-
porarily settled in the area near the Reservation headquarters, there
was much quarreling. On April 2, 1956, a quarrel between a pair
of red-bellied woodpeckers and a male starling was observed in a
willow snag having a cavity offering a potential nest site for both
species. The starling was the most aggressive. It would dart first
at one woodpecker and then at the other, but it did not succeed in
driving them from the tree. After ten minutes of continuous quar-
reling it gave up and flew out of sight headed northwest.

One woodpecker that was color-banded and recorded from time
to time over a period of a year, used an area of 20.6 acres measured
within the five marginal points. Most of the recorded movements
and locations were near the headquarters and the pond, but a
single record at a place 900 feet north of the pond greatly increased
the total area involved. Certainly the entire 20.6 acre area was not
used regularly. In 1955 the nest was in a sycamore snag southwest
of the pond. In finding food for the nestlings the adult regularly
made trips to places 550 feet southwest, 375 feet north, and 200 feet
east of the nest.

In May, 1956, a nest was located in a willow snag beside a gully
in a relatively open situation. The adults were often observed fly-
ing to and from the nest on foraging expeditions. Because the nest
was away from woodland, the foraging expeditions probably were
to greater distances than are typical for the species. The adults
made expeditions in all directions from the nest, visiting not only the
woodland but isolated trees and groves. The main line of flight
was north northeast and south southwest, to an extreme distance
of 520 feet in the one direction and 820 feet in the other. The area
encompassed by all the observed foraging flights was measured
as 17.3 acres. This total area, however, included several major
portions that were not even crossed by the woodpeckers, and their
movements were almost entirely confined to an area of irregular
outline measured as nine acres.

Home Ranges of Vertebrates 217

A woodpecker that was actively foraging, with frequent shifts
from tree to tree, was followed for several hours on December 7,
1953. The hexagonal figure outlining its route was 9.6 acres, and
this probably represents the greater part of its home range at that
particular time. For one banded, and recorded several times in
November and December, 1953, a pentagonal range of twelve acres
was recorded. Much of this area consisted of an open field, which
was merely crossed by the woodpecker, staying mainly in the woods
along its borders on each side. A triangular area of wooded slope
within which another was noted repeatedly was 5.4 acres. This
probably was only a part of the actual home range.

Occasionally red-bellied woodpeckers were seen making long
direct flights, which were almost certainly beyond the limits of the
usual home range or territory. On September 15, 1953, one was
seen flying high, from a hilltop ("Sugarloaf") in the northwestern
part of the Reservation, across an intervening open valley to another
hilltop nearly half a mile farther south, a total distance estimated
as 2400 feet, although the exact spots where the flight began and
ended were not seen. On December 27, 1953, one that was moving
about from tree to tree frequently as it foraged was followed over
an area 1350 feet across.

Territories between ten and twenty acres in extent are probably
most typical of tlie species on the Reservation and in similar habi-
tat. The same general area is occupied by an individual through-
out the year, but with some shifting, occasioned chiefly by the
mutual exclusiveness of pairs in the nesting season.

Melanerpes erythrocephalus (Linnaeus)

Red-headed Woodpecker

Status. — Moderately common resident, but localized to small area
of woodland along eastern edge of Reservation.

Habitat. — Chiefly forest.

On the Reservation the one colony present from 1952 through
1956 was closely confined to an area of red oak woodland, the part
of the area most closely approximating a climax deciduous forest.

Movements. — Red-headed woodpeckers were occasionally seen
far from their usual habitat. These were wandering individuals.
Some were travelling, but others remained for hours before moving
on. It was not determined whether these wanderers had migrated
for long distances or were from nearby colonies. They were seen

218 UNivERsiri' OF Kansas Publs., Mus. Nat. Hist.

chiefly in late spring and early autumn, ha\dng dispersed just before
and just after the nesting season. By months, they were distributed
as follows: May — 5, June — 1, August — 2, September — 5, October
— 2. Some of those seen in September and October were in juvenal
plumage. From September 10 to 19, 1955, a small group settled in
the grove just northeast of the Reservation headquarters. During
the time that they stayed most of their activity was confined to
several large elms within a hundred-foot radius, entirely within the
territory of a pair of red-bellied woodpeckers. There was almost
constant quarreling between the two kinds. Scolding and pursuit
by the red-bellied woodpeckers may have been the motivation for
the red-headed woodpeckers to leave.

In 1956 there was a similar invasion at the same place in the last
week of August. On the morning of August 31 at least six were
present, and one or more had been heard in the vicinity for several
days previously. This group remained until September 6, then
disappeared abruptly, but in 1957 another group was present in the
same place over the period September 4, 5 and 6.

The colony often observed on the eastern edge of the Reservation
covered an area of more than 50 acres including almost level, creek-
bottom land with exceptionally large, old trees, and the adjacent
slope with oak woodland. More than a dozen of the woodpeckers
inhabited this area, but no territoriality was discernible; they seemed
to move about more or less at random, over the entire area, usually
wandering singly or in small groups, each individual tending to
keep within sight or sound of others.

Sphyrapicus varius (Linnaeus)

Yellow-bellied Sapsucker

Status. — Rare transient.

Habitat. — Woodland.

Movements. — On September 30, 1952, at 8 A. M., one in juvenal
plumage was working on an elm at the Reservation headquarters.
Another, also in immature plumage, was seen in oak woods on the
Rockefeller Tract on April 9, 1957.

Dendrocopos villosus (Linnaeus)

Hairy Woodpecker

Status. — Permanent resident, with population of several pairs.
Habitat. — Woodland, especially where there are large mature
trees, oaks, hickories, elms and locusts.

Home Ranges of Vertebrates 219

It is rarely seen in parts of the woodland where the trees are
sparse or scrubby, nor in isolated trees growing in the open. Oc-
casionally seen crossing broad, open fields, in high, direct flight.

Movements. — Hairy woodpeckers were especially noteworthy for
the year-round association of pairs in permanent territories. All
individuals on the area were shy (a flushing distance of approxi-
mately 150 feet was typical) and none was individually marked.
Therefore, individual recognition ordinarily was uncertain, but
pairs were followed from time to time and were seen in the same
places with such regularity that identity could be guessed with some
assurance.

On December 10, 1953, a territorial encounter between two males
was seen. Soon after one flew into a large elm, another came from
the opposite direction and perched nearby. They challenged each
other, bowing and scolding. One lunged at the other and when it
flushed he followed in close pursuit. The chase led in a circular
course, with two complete circuits approximately 200 yards in di-
ameter. Two days later a male which probably was one of those
involved in the chase, was followed as he moved about from tree
to tree, and the area covered corresponded roughly with that over
which the chasp extended.

In late 1953 and early 1954, two pairs were often observed sepa-
rately, and occasionally in contact, in the area about the head-
quarters. The woodland encompassed by the records of one pair
comprised approximately 37 acres. The second pair occupied an
adjoining territory consisting of two separate blocks of woodland
totalling approximately 22 acres, but with an intervening area of
open field that was almost as large. On several occasions these
woodpeckers were seen to make high, direct flights from one hill-
side to another one thousand feet or more away, crossing the open
field intervening. In these and shorter flights, the bird that flew
first (most often the female in observed instances) called as it
launched into flight, and the mate followed within a few seconds.
They were never seen to fly close together, but always made their
moves to new areas well spaced in tandem fashion. In their forag-
ing they tended to keep in separate trees, often as much as 100 feet
apart, but obviously aware of each other's whereabouts through the
sound of pecking and the occasional sharp squeaking calls.

Hairy woodpeckers were often noticed foraging in close associa-
tion vi^ith other kinds of woodpeckers, especially the downy and less
frequently the red-bellied. Such associations often were maintained
when the birds were shifting from one tree to another, suggesting

220 Uni\'ersity of Kansas Publs., Mus. Nat. Hist.

that actual sociability, rather than mere attraction to a common
food supply was involved. However, at other times hairy wood-
peckers have been seen to display varying degrees of intolerance
(scolding, threat, and actual pursuit) toward both these other spe-
cies. The reason for such intolerance is not understood. It does
not seem to be seasonal, and has been noted at times of year other
than the breeding season. Perhaps intolerance involves unfamiliar
individuals, while those that are met regularly are accepted as co-
occupants of the territory.

A roost in a hollow elm at the gully just south of the Reservation
headquarters was used regularly from 1952 to 1957, probably by the
same individual. Each evening near sunset this female hairy wood-
pecker came to trees near the roost, usually making a flight of hun-
dreds of yards, from a remote feeding area. She was remarkably
circumspect in approaching the roost, and sometimes spent as much
as an hour in the process, maneuvering and calling in the vicinity.

In May, 1955, a nest with young was discovered fifteen feet high
in a live honey locust, in thick woods on a north slope. The par-
ents were not observed to forage near this nest, but upon leaving it
they usually made flights of hundreds of yards to other hillsides.

Dendrocopos pubescens (Linnaeus)
Downy Woodpecker

Status. — Abundant resident.

Habitat. — Found in all situations where there are trees or large
shrubs.

It is numerous in all parts of the woodland, using every species of
tree on the area; perhaps even more abundant in parkland where
trees are scattered. This small woodpecker does part of its foraging
on annual weeds, notably the giant ragweed.

Movements. — In the breeding season each pair keeps to its own
small territory, excluding those of other pairs. As these wood-
peckers are sedentary in habits, they tend to remain year-round in
the same general area. Occasionally there is some displacement.
Especially after severe winter storms individuals have been seen
foraging in roadside weeds and in other situations removed from
their usual woodland habitat. Pairs may continue their association
throughout the winter. In February, 1952, a male was color-banded
near the Reservation headquarters and was seen frequently in a

Home Ranges of Vertebrates 221

period of weeks thereafter, usually associated with a female. All
records for this male were within a quadrangular area of 31.5
acres, with most remote locations separated by 2180 feet. Most
of the records were concentrated in an oval inner area of only 4.6
acres. Downy woodpeckers were perhaps especially in evidence
in 1952 and 1953, when elms were dying off as a result of the com-
bined effects of drought and attack by bark beedes ( Scolytus multi-
striatus). These minute but numerous beetles produced an ample
food supply, and the woodpeckers concentrated their attention on
heavily infested trees. About the bases of such trees, chips of bark
accumulated to depths of several inches. The same tree even
though small, might be returned to day after day.

Often in winter, a downy woodpecker or a pair were found in
association with mixed flocks of small passerines, usually including
black-capped chickadees and tufted titmice, and sometimes includ-
ing other kinds, such as brown creepers, and golden-crowned king-
lets. When such mixed flocks were kept under observation, they
were soon noted to undergo at least partial disintegration. The
woodpeckers, especially, were liable to be left behind because they
moved about less rapidly than the other kinds in foraging.

In a pair that is moving about, each keeps the other informed
of its whereabouts by an occasional sharp metallic call. In winter,
frequentiy, a pair has been observed to attack or chase a third in-
dividual or a second pair. This does not involve a defense of hard-
and-fast territorial boundaries, but is indicative of intolerance for
other individuals foraging in the neighborhood of a favored feeding
place or of the mate.

Although such hostility is noticeable at all times of year, it is
especially prominent in early spring. In mid-April, 1954, a dead
snag, having several hollows, at the edge of the woods near the
headquarters, was the scene of constant quarreling. Sometimes as
many as six downy woodpeckers (presumably those of three dif-
ferent pairs) were in the vicinity simultaneously. Though both
sexes participated in the quarrels, males were especially pugnacious.
Two would fly from tree trunk to tree trunk, uttering whinneying
calls, and perched on opposite sides of a trunk, would thrust at each
other with their bills. Territorial drumming was heard as early as
the first week of February but chiefly in late March and early
April.

222 University of Kansas Publs., Mus. Nat. Hist.

Tyrannus tyrannus (Linnaeus)
Eastern Kingbird

Status. — Often present in summer, but not knowai to breed on
the Reservation. Those that appear from time to time are wan-
derers that have not yet established territories or have left them
after the breeding season. Usually they do not remain for more
than a few hours in any one place.

Habitat. — Prefers relatively open ground with high perches, for
foraging.

On the Reservation, in the absence of grazing, the dense grass
or herbaceous ground cover evidently renders habitat conditions
unsuitable for this kingbird.

Movements. — In July, August and September, 1948, when field
work was begun on the Reservation, kingbirds were seen regularly.
Parts of the area were then heavily grazed pastures, and habitat con-
ditions were probably favorable for breeding of the kingbird. With
discontinuance of grazing in early 1949, habitat conditions de-
teriorated for this species because of the high rank growth of
herbaceous vegetation. Thenceforth, kingbirds were seen on the
area only on migration and, occasionally, as wanderers from adja-
cent farms where the habitat remained favorable.

Kingbirds have been seen on the Reservation chiefly in May, when
they have recently arrived from their northward migration, and
perhaps are still moving about in search of territories.

Kingbirds nesting at a farm a quarter mile north of the Reserva-
tion boundary were seen on the area near its northern edge on
several occasions. In late August of several different years, loose
groups of kingbirds were seen moving across the Reservation or
nearby areas, perhaps already embarked on their southward migra-
tion.

Muscivora forficata (Gmelin)

Scissor-tailed Flycatcher

Status. — Rare transient.

Habitat. — Short-grass prairie or closely grazed pasture with ele-
vated perches such as arc pro\'ided by scattered trees, fence posts,
or telephone wires, comprise suitable habitat for this flycatcher.
Presence of woodland and thickets alternating with rank ground
vegetation in more open situations renders the Reservation unfa-

Home Ranges of Vertebrates 223

vorable for occupancy. This area is on the margin of the species'
range, and breeding pairs have been recorded within a few miles.

Movements. — The only definite record for the Reservation is one
seen by Donald W. Janes on August 28, 1955, perched on a tele-
phone wire between the headquarters and the west boundary. It
was seen twice at the same place. On the second occasion, as a
vehicle approached on the road, it flushed and flew out of sight
to the west.

Frank W. Fitch, Jr. (1950:153) studying the species in east-
central Texas, found that "Territories . . . included the ground
within a radius of 35 yards from the base of the nest tree."
". . . an area of complete intolerance to all species immediately
surrounds the nest, with gradations of intolerance outside of that,
their extent depending on the species involved."

Myiarchus crinitus (Linnaeus)
Crested Flycatcher
Status. — Common summer resident.

Habitat. — Deciduous forest, and edge.

Ordinarily these flycatchers stayed in woodland, showing a dis-
tinct preference for large old trees, especially those that were dead
or dying and therefore had open crowns.

Movements. — Crested flycatchers first appeared on the Reserva-
tion in late April each year; dates were the 27th, 30th, 28th, 20th,
22nd, 26th and 22nd over the period 1951 through 1957. Within a
few days after the first one was heard they were present in abun-
dance, and were establishing territories. They were noted chiefly
in May and June. As the season advanced, their calling became
increasingly restricted to early morning and evening, and finally
ceased altogether. Latest seasonal records are August 27, 1952,
August 30, 1953, and August 31, 1955.

In early May crested flycatchers were already associated in pairs.
Quarrels between pairs, and pursuits have been noted chiefly in
May and June. Often two pairs perched in separate trees within
100 feet of each other where their territories adjoined and the males
called in seeming defiance, answering each other in turn. Pairs
have been seen nest-building in early June. In the nesting season
crested flycatchers have often been seen to chase other kinds of
birds, including phoebes and cardinals.

In 1955 three adjoining territories were measured, one in the

6—3117

224 UNi\'ERSi'ri' OF Kansas Publs,, Mus. Nat. Hist.

headquarters area of the Reservation, and one on each side of it.
The appro.ximate areas computed were: 7.2 acres, 6.6 acres, and 5.6
acres. All three territories included areas of fields with groves and
scattered trees, and edges of woodland bordering the fields on
each side. Activity of the flycatchers was mostly confined to the
woodland, but in early morning and evening each ventured out
into the field and called from the tops of several of the larger trees,
in succession. In several instances such a favorite perch tree was
used at difi^erent times by the birds from each of t\vo territories.
Neither seemed able to establish exclusive control, but in each in-
stance the tree was on the borderline of territories that otherwise
did not overlap each other.

Engles (1949:287) describes two nests in dead trees where forest
had been killed by wind blown sand, on Shackleford Bank, a small
island on the North Carolina coast. These nests were, respectively,
274 and 167 "long paces" from the edge of the woodland where the
flycatchers did nearly all their foraging. As a result of their unusual
situations, the nests were relatively safe from predators, but it was
a hard struggle for the flycatchers to fly against the wind back to
their nests.

Sayomis phoebe (Latham)

Phoebe

Status. — Common summer resident, several breeding pairs present
each year.

Habitat. — The distribution of bridges and buildings providing
sheltered nest sites seems to be the chief factor in determining the
number of phoebes on the area.

There were perennial nest sites at the Reservation headquarters
buildings, at a wooden bridge over a deep gully of the creek in
the southeast part of the area, and inside a seven-foot concrete cul-
vert under the county road at the west edge of the Reservation.
In their foraging, phoebes seemed to prefer parkland situations,
perching in large elms, but also spent much time in nearby fields
of brome grass, where they perched in isolated trees or on stakes
and weed stalks.

Movements. — Phoebes arrived on the area each year in early
spring, and their annual sojourn was of longer duration than that of
any other kind of bird that was not a permanent resident. Earliest
appearance each year was in March, and for 1950 through 1957
first recorded dates were the 21st, 22nd, 12th, 14th, 12th, 10th, 6th,

Home Ranges of Vertebrates 225

and the 13th. In most years the first phoebe noted was in the head-
quarters area, where nesting occurred annually. Several times such
recent arrivals were noted at daybreak or soon afterward, presum-
ably having arrived in the night. The fact that they preceded the
main wave of migration, and that they came directly to a nesting
territory, suggested that these individuals were survivors tliat had
nested there the year before. However, in 1954, the first phoebe,
recorded on March 12, was more than 100 yards from the head-
quarters area, and it did not come to the buildings. After a few
minutes it left, and no others were recorded until March 21.

Buildings were erected at the Reservation headquarters in early
1950, and these provided three perennial nesting sites. One was
on a bend of the drain pipe beneath the eaves of the residence at
the southwest comer, one was in the garage (or under its eaves)
150 feet east northeast, and the third was on the porch of the labora-
tory between the other two buildings and approximately 50 feet
from the garage. Two pairs always occupied this vicinity; the house
and the garage were in diflFerent territories. The laboratory nest
site was usually used as an alternate by the pair whose territory
included the garage, but occasionally it was used by the pair from
the house.

The territories occupied were contiguous, and they were similar
from year to year. In each, foraging was concentrated in a parklike
grove, mainly of large elms, in an area that was roughly circular,
and approximately 170 feet in diameter, a little more than half an
acre (Fig. 16). Several favorite perches, five to fifteen feet high,
were used with such regularity that conspicuous accumulations of
excreta and pellets formed beneath them. However, the phoebes
often foraged far beyond the limits of the groves where their activi-
ties were concentrated. The pair living at the garage often used the
open field to the west, perching in the several isolated trees or on
weed stalks, as far as a gully that was 600 feet from the nest site.
At times, they spent hours daily along the cut-banks of the gully
and in adjacent trees and shrubs, and seemed to be preparing to
nest there, but no nest was ever found at the gully. In the op-
posite direction this pair foraged to the pond, approximately 580
feet east from the nest and their territory was about 100 yards wide.
Their territory was measured as 7.1 acres, while the pair that nested
at the house had a less elongate territory of 3.3 acres.

Throughout the breeding season territorial pursuits and attacks
were frequent, owing to the proximity of the two nests. Conjlict

226

Untversity of Kansas Publs., Mus. Nat. Hist.

Fig. 16. Territories of two pairs of phoebes; activities of both pairs were
concentrated about the Reservation headquarters, as shown by the lines
of crosses. Little used outlying portions of the territories are enclosed by the

dotted lines.

was especially severe early in the season before territorial bound-
aries were fully established. The sequence of events in 1953 well
illustrates the establishment of territories, and the seasonal cycle.
The first phoebe noted was present on the morning of March 14,
but no other was seen that day, or on March 15 and 16. At day-
break on March 17, two were calling near the old nest at the south-
west corner of the house. Later in the morning they were seen
near the garage and at other places nearby, perched near together
and seemed to be paired. Once there was a spirited pursuit. On
March 18 and 19, two phoebes, probably the same, were present in
the vicinity. On March 20 at 8 A. M., a third phoebe was present,
and the three were involved in a violent quarrel, centering at the
old nest on the drain pipe at the corner of the house. The two
already paired were trying to drive away the one newly arrived.
Most maneuvering was within thirty feet of the nest, and one mem-
ber of the pair was much more aggressive than the other. Attacks
were most severe when the intruder lit on or beside the nest. A de-
fender would fly at it, and would follow close behind as it darted
about erratically. Sometimes the pursuer would catch the other,
and cling to it persistently, forcing it to the ground where the two
might continue to struggle for a few seconds before separating.
After escaping, the pursued bird would soon return to approach the
nest again. Its persistence suggested that it may have used this

Home Ranges of Vertebrates 227

nest site previously. However, by 9:30 A. M. it had w^ithdrawn
from the contest for the nest site and was keeping to the vicinity
of the garage and laboratory apart from the pair. The latter were
staying mostly within thirty feet of the nest site, alternately forag-
ing and courting. The female tended to fly away a short distance
each time the male approached. A little later the male flew to the
single bird and perched beside it momentarily, twittering as in
courtship. Later he repeated this procedure with increasing fre-
quency. In the latter part of the morning he was distributing his
time between the two females, spending about two-thirds of it with
his original mate and the remainder with the second female. On
March 21 wintry weather returned, with high wind and freezing
temperatures. On March 22 weather moderated, but the phoebes
were preoccupied with foraging, and no courtship or quarreHng
was noted. Once, all three were seen perched in the same tree.
After dark two were flushed from the garage, suggesting that the
lone female had acquired a mate, and on the morning of March 23
four phoebes were present, behaving as two pairs on separate ter-
ritories. There was persistent quarreling and maneuvering in the
area between the buildings on this morning, and probably the ter-
ritorial limits of each pair where their ranges met were established
as a result. Later in the day, one (female?) was seen carrying
nesting material to the garage, from the ravine 250 feet south of it.
On March 25 the nest in the garage was well underway and one of
the phoebes made frequent trips to a ditch 190 feet north, carrying
back mud for building material. On March 26 a territorial quarrel
involving all four phoebes lasted intermittently for more than an
hour in the area south of the house. On March 28, a phoebe was
observed nest-building at the comer of the house for more than an
hour. It was making frequent trips to the ravine 225 feet south-
east, and carrying back moss and perhaps mud. On April 3 this
nest seemed to be completed. On April 4 the female was acci-
dentally entangled in a net and was found dead. She contained
a fully formed egg ready to be laid. The nest in the garage probably
received its first egg on this day or the next. The clutch of four
was completed on April 8. At dawn this same day a new female
had replaced the one accidentally killed and courtship was in prog-
ress beside the nest at the corner of the house. Spirited pursuits
and fighting were noted later in the day, doubtless resulting from
the new female's unfamiliarity with the limits of the territory she
had inherited. On April 11 an egg was found broken beneath the

228 University of Kansas Publs., Mus. Nat. Hist.

nest in the garage. On April 13 the garage nest had again been dis-
turbed by an unknown marauder; only two eggs remained and the
phoebes had deserted. The nest on the house had been raided, torn
loose from its foundations, and its lining ripped out. The pair of
phoebes that had used this nest were making frequent trips to a
puddle 250 feet west and carr)'ing back mud to repair the old nest
on the porch of the laboratory.

On April 20 the nest in the garage was empty. On April 21 it was
being repaired, partly with spider webs being gathered by the
phoebes in a thicket 275 feet south. On April 26 at dusk, a phoebe
was flushed from the renovated laboratory nest, which contained
a newly laid egg. On April 29 there were four eggs in the garage
nest. On April 30 and May 1, the second and third eggs were
added to the laboratory nest. The eggs of the garage nest all
hatched on May 11 and 12; those of the laboratory nest hatched
May 19 and 20. Both broods were fledged. Those of the labo-
ratory nest left June 3 to 6. On June 10 the adults were seen in
courtship beside the nest and on the following day one was carrying
material to repair the nest. A new clutch of eggs was completed on
June 19 and the young that developed from them were fledged on
July 18. Thenceforth, phoebes were little in evidence about the
headquarters buildings where they had spent the spring and early
summer, but throughout August and September the buzzing court-
ship calls were heard almost every day at dawn at one or more of
the old nests. This behavior was last noted on October 18.

If not prematurely disturbed, the young phoebes stay in the nest
until well feathered and upon leaving fly strongly to an elevated
perch. When one flies from the nest, others may follow close
behind it and alight near its perch. Or, if the brood is scattered
at first, the hunger calls given by all the young guide them in as-
sembling. Characteristically they perch in a tight cluster in the
same bush or tree, and at least one parent stays nearby foraging
in the vicinity and feeding them in turn. Whenever one young
flies, others are likely to follow. Presumably such an aggregation
of fledged young is tended largely by the male parent, and in some
instances the female was already incubating a new clutch while
the previous brood of fledglings was still dependent. So far as
observed, the dependent fledglings remained in the adults' nesting
territory but shifted to a part of it remote from the nest itself.
For aggregations of young of seven different broods recorded over
a period of several years, the distance from the nest tended to
lengthen somewhat with elapsed time, as follows: 1 day, 70 feet;

Home Ranges of Vertebrates 229

2 days, 200 feet and 260 feet; 3 days, 90 feet and 100 feet; 5 days,
460 feet; 8 days, 120 feet; 11 days, 240 feet; 12 days, 300 feet.
Smith (1942:410) recorded similar tendencies in fledglings in
Vermont. He observed the young all within a 75-foot radius, 150 to
200 feet from the nest the first day after leaving. At dusk on the
third day the young were observed clustered on a branch 300 feet
from the nest. By the seventeenth day after leaving, the young were
seen with one parent 1300 feet from the nest, but on the eighteenth
day, all were back near the nest. Smith noted a gradual reduction
in parental attention for the fledglings, which had become partly
independent by the seventh or eighth day. After the eighteenth
day they were fully independent, and had dispersed.

On one occasion I observed a phoebe 850 feet from its nest, and
another was seen 750 feet from its home base, but these distances
are unusual. When feeding nestlings, phoebes may stay within 70
feet of the nest.

Latest recorded occurrences in six different years were: October
11, 1951; October 31, 1952; October 18, 1953; September 16, 1954;
October 26, 1955; and November 3, 1956. In each year phoebes
may have actually remained somewhat later in the season, as they
were relatively inconspicuous and silent, and might have been
overlooked.

Empidonax flaviventris ( Baird and Baird )
Yellow-bellied Flycatcher
Status. — Rare migrant.

Habitat. — Northern coniferous forests in wet places during the
breeding season.

Movements. — R. W. Fredrickson recorded one on May 12, 1950.

Empidonax traillii (Audubon)

Alder Flycatcher

Status. — Temporary resident in late spring.

Habitat. — Groves and woodland edges.

Movements. — These small flycatchers have been noticed on the
Reservation each year in May. In 1952 and 1956 they were es-
pecially conspicuous and seemed to be present in unusually large
numbers. The willow grove at the pond was their favorite habitat.
They were frequently seen also at various places along the wood-
land edges of the field in which the headquarters were situated.
In 1952 they were seen or heard almost daily from May 20 to June

230 University of Kansas Publs., Mus. Nat. Hist.

3. In 1956 they were noted from May 4 to approximately June 4,
with the main influx about May 9. In late May several could be
found at the willow grove at almost any time, and singing was
frequent. Each seemed to move about the grove freely rather than
being restricted to any definite part of it, but occasional squabbles
and pursuits were seen. Larvae of chrysomelid beetles were
abundant on the willow leaves and provided a ready food supply.
Several times the flycatchers were seen to depart from their usual
method of foraging to hover while picking off these larvae. In
1957 the only record was obtained on May 10.

Empidonax minimus (Baird and Baird)
Least Flycatcher
Status. — Occasional temporary resident in late spring.

Habitat. — On the Reservation seen chiefly at woodland edges
having thickets and dead trees.

Movements. — On May 17, 1952, one was seen chasing another,
at woodland edge beside the ravine about 100 yards south of the
Reservation headquarters. On the following day one foraging
near the same place moved from perch to perch in open tree
crowns, tending to keep over blackberry thickets. Several were
seen on May 17, 1955.

Breckenridge (1956) studied territoriality in this species in
Minnesota. He found that territories were usually less than one
acre, but size depended partly on height of the trees. He regarded
territories as three-dimensional, extending vertically from the top
of the shrub stratum to the leaf canopy of the dominant trees.

MacQueen (1950:204) also studied territoriality in this species,
at Douglas Lake, Michigan. For 33 territories the size range was
from .5 to .03 acre and the average was .18 acre. The territory
was found to be defended chiefly by the male.

Contopus virens (Linnaeus)
Eastern Wood Pewee

Status. — Common summer resident in wooded parts of Reserva-
tion.

Habitat. — Deciduous forest especially in edge, parkland and
other situations where leaf canopy is somewhat discontinuous.

Movements. — Pewees were notably consistent in their arrival

Home Ranges of Vertebrates 231

dates; over a six year period earliest records were: May 16, 1952;
May 11, 1953; May 17, 1954; May 10, 1955; May 14, 1956; and May
10, 1957. They were seen and heard much less frequently in late
summer. The latest record is September 17, 1956. Dead trees or
branches seemed to be the favorite locations for territorial calling,
but in the course of its normal foraging each bird shifted frequently
from perch to perch through the crowns of the trees, in a circuit
of its territory.

In defense of territory pewees were especially pugnacious, not
only against members of their own species, but against other kinds
of birds. They have been seen to chase away mourning doves,
bluebirds, phoebes, and field sparrows. On May 22, 1953, one
pewee darted at another and caught it in midair. They fell into
the grass and after a brief struggle one escaped, closely pursued
by the other. A third followed these two.

A pewee that stayed in the headquarters area was observed and
followed frequently in August, 1954. On the morning of August 8
its activities were confined to an area measured as .34 acre. On
August 12 it covered .57 acre, overlapping only in part the area
covered August 8. The woodland edge grove within which it
usually stayed totalled 1.1 acres, but occasionally it visited outlying
isolated trees in the adjacent field. If these and the intervening
area of field were considered part of the territory, its size would be
more than two acres.

Notably larger territories were found by Odum and Kuenzler
(1955:131), perhaps in more open habitat. They used the wood
pewee to illustrate their standardized method of determining size
of territory in birds. The individual (male) most thoroughly
studied by them was determined to have a territory of 10.8 acres,
and fifty observations were necessary, as with each series of ten
observations, up to the fifth series, the size of the known range
increased by more than ten per cent. At this stage the known ter-
ritory was a nine-sided polygon. With subsequent observations,
the size of the known home range continued to increase, but at a
slower rate — below the one per cent level. With the total of 90
records finally accumulated, the territory was still a nine-sided
polygon, but with boundaries somewhat extended. Many of the
records were in sizable clusters ( 14, 10, 10, 8, 5 ) , and it was evident
that certain parts of the area were intensively used, whereas more
extensive intervening portions were used relatively little or not at all.

232 University of Kansas Publs., Mus. Nat, Hist.

Nuttallomis borealis (Swainson)
Olive-sided Flycatcher
Status. — Rare migrant.

Habitat. — In breeding season, northern coniferous forest; on Res-
ervation, on migration, woodland and edge.

Movements. — On May 22, 1951, at dusk, two were calhng at
different places at the edge of the woods near the Reservation head-
quarters. On October 13, 1951, one was seen by Maurice F. Baker
on a telephone wire between the headquarters and the west edge
of the Reservation.

Eremophila alpestris (Linnaeus)

Horned Lark

Status. — Common year-round resident in the general area, with
numbers augmented in winter by an influx of northern birds; on
Reservation, habitat has disappeared as revegetation of formerly
barren areas has progressed, and no horned larks have been re-
corded since 1949,

Habitat. — In eastern Kansas, chiefly cultivated fields; occasionally
other open areas that are nearly denuded of vegetation, such as
sand or dried silt of stream deposits, and heavily overgrazed pas-
tures.

Movements. — In June, 1949, a male was seen singing as it flew
over an eroded fallow field at the north edge of the Reservation,
and a cornfield adjacent on the north. Its territory seemed to in-
clude parts of both areas. In subsequent years none was seen at
this place or elsewhere on the Reservation, because the open type
of habitat that horned larks require had disappeared as plant suc-
cession progressed.

In the spring of 1957 horned larks were much in evidence on the
newly acquired Rockefeller Tract adjoining the Reservation on the
north. Fields which in 1956 had crops of milo and wheat, were
disked and sown with seeds of native grasses in March, 1957. In
early April, seedlings chiefly of annual weeds appeared on this
almost barren area, and by late May weeds on some parts of the
area had increased in height and density rendering the habitat
much less favorable for the larks. A northern block of field com-
prising slightly more than 36 acres was separated from a southern
block of a little less than 36 acres by an alfalfa patch, barnyards

Home Ranges of Vertebrates 233

and pasture. Six pairs of horned larks occupied the 72-acre area
of fields, with three pairs each on the northern and southern blocks.
The distance between the two fields varied from 150 feet to 360
feet and the field's edges formed natural territorial boundaries.
However, on many occasions a pair or a male living on one side
of the pasture or alfalfa patch flew across and invaded a neighbor-
ing territory. Fighting and pursuit were seen frequently. Some-
times a chase lasted for several minutes, one lark flying in the lead,
low and rapidly, with frequent sharp turns to avoid the one behind
it. Flight song was prominent throughout the nesting season, and
especially after territorial encounters. The lark would spiral and
tower over the territory in a flight often continuing for five minutes
or more, then would swoop abruptly to the ground. It might con-
tinue to sing at intervals as it wandered about the territory foraging.
Relatively little singing was done from perches. In the nesting sea-
son females were seldom seen in flight except when they were
flushed. Territories of approximately twelve acres were typical in
this habitat.

Riparia riparia (Linnaeus)

Bank Swallow
Status. — Occasional.

Habitat. — For nesting requires cut-banks; otherwise not closely
limited in habitat requirements.

Movements. — In May, 1949, a pair was seen patrolling up and
down a gully and the adjacent pasture on the Reservation, west of
the headquarters area. These swallows may have been nesting or
preparing to nest in the bank of the gully, but a few days after they
were first seen they disappeared, and none was recorded in sub-
sequent years.

Stelgidopteryx ruficoUis (Vieillot)

Rough-winged Swallow

Status. — Occasional in summer.

Habitat. — Usually near bodies of water.

Movements. — Rough-winged swallows have been identified fly-
ing over the Reservation on a few occasions in summer. Locations
of breeding colonies in surrounding areas are not known. The swal-
lows seen over the area were probably either migrants or individuals
that had wandered farther than usual from their normal ranges.

234 University of Kansas Publs., Mus. Nat. Hist.

Hirimdo rustica Linnaeus
Barn Swallow

Status. — Present in summer; many nest on nearby farms, but
none has nested on Reservation.

Habitat. — The chief habitat requirements are: nesting sites of
the preferred types, as provided by old farm buildings, and foraging
places, chiefly meadows or cultivated fields. Swallows were often
seen foraging over the Reservation but it is doubtful whether any
individuals had definite home ranges including parts of the Reserva-
tion. They were seen over every type of habitat represented.
The altitude of flight and the place chosen for foraging probably
varied according to the availability of insect prey, as determined
by season, temperature, and humidity.

Movements. — Barn swallows usually were not seen on the Reser-
vation until somewhat after their arrival in the general area, in late
spring. Earliest dates recorded over a period of years were: May
7, 1952, May 15, 1953, May 18, 1954, May 17, 1955, and April 25,
1956. There were nesting colonies in some years, at least, at farm-
sites located a quarter-mile north of the Reservation (this latter
acquired by the University in 1956 as the Rockefeller Tract), and
a half-mile west. These colonies were the sources of most that were
seen on the Reservation. On May 15, 1953, when conditions were
unusually dry, a pair flew to the Reservation pond and settled on
bare ground at the edge of the water, gathering mud for a nest,
then flew up a slope, headed for tlie farmsite some 2800 feet to
the north. Ordinarily material for building was collected much
nearer to the nest. In the nesting season most activity was within
a quarter mile of the colony and was concentrated about a small
pond 100 yards from the barn in the area between the pond and
bam, and in the adjacent pasture and wheat field. Only an occa-
sional individual crossed to the Reservation a quarter mile south.

On May 30, 1955, a pair came to the buildings on the Reservation
and spent several minutes circling the house, hovering under the
eaves, and clinging to the walls. These birds soon left, and did
not return. Probably they were from a nearby colony and having
failed in an attempt at nesting, were in search of a new nest site.

On July 9, 1952, there were many barn swallows on the wires
at the west edge of the Reservation and foraging over the fields to
the west. These were from the colony at the farmsite half a mile
west of the Reservation. The groups on the wires included many
fledglings, that were being fed from time to time by the adults.

Home Ranges of Vertebrates 235

In 1957 a pair nested in the barn on the Rockefeller Tract. Most
of their foraging was confined to an area of about 20 acres, the
farmyard, the 14-acre pasture to the east, and a five-acre alfalfa
field to the west, although they frequently cut across corners of
the cultivated fields adjoining. The pair of swallows perched
mainly on a 50-foot stretch of electric wire beside the barn. Latest
dates of records were September 12, 1957, and September 27, 1948.

Petrochelidon pyrrhonota (Vieillot)
Cliff Swallow

Status. — Occasional in summer.

Habitat. — Vicinity of water probably is preferred, especially
where cliffs or buildings provide nesting sites.

Movements. — No nearby colonies are known and appearances
over the Reservation are infrequent. My only specific record is
for July 23, 1948. In most instances swallows seen over the Reser-
vation were flying high and were not definitely identified as to
species.

Progne subis (Linnaeus)
Purple Martin

Status. — Transient in late spring and summer.

Habitat. — Breeds chiefly about farms and suburban situations.

Movements. — Martins are seen over the Reservation most often in
May. Those seen probably are still on migration or are wandering
in search of suitable nest sites.

Cyanocitta cristata Linnaeus
Blue Jay

Status. — Common resident.

Habitat. — Hardwood forest.

Locally jays keep mainly to the woodlands of oak, elm and hick-
ory. They are often seen crossing open areas, in high direct flight,
but have never been seen to stop on the ground in open fields. They
seldom stop in isolated trees in open fields, and spend little time in
such situations. In summer they often frequent .woodland edge
and parkland situations, but in winter, they seem more inclined to
stay in thick woods.

Movements. — Numbers of blue jays on the area varied widely
from month to month and from year to year. In part this variation

236 University of Kansas Publs., Mus. Nat. Hist.

was due to passage, through the area, of migrants from farther
north. Jays that breed on the area reside year-round in the vicinity,
but are incHned to shift their ranges seasonally according to avail-
ability of cover and food. The tendency of the jays to carry on most
of their activities as members of social groups, and the existence of
more or less permanent bonds between the members of a pair, are
factors that control, to a large extent, the movements of individuals.

In spring, the migratory northward movement through the area
occurs chiefly in late April, after the resident blue jays have begun
nesting. At 9 A. M. on April 29, 1952, 30 or more strung out in a
long loose group, were moving northward across the Reservation,
passing near the headquarters building. Many of the birds in this
group seemed to be associated in pairs. At 4 P. M. on the same day,
a smaller group was moving north along a hilltop edge, a quarter
mile farther east. At 8:20 A. M. on May 18, 1953, a long loose flock
of 22, mostly grouped in pairs, moved nortli across the Reservation
near the west edge. At 11 A. M. on April 24, 1954, a straggling
group of 27 was travelling north northeast, flying high over the
treetops. On April 26, 1956, between 9:00 and 9:07 A. M., a group
of 32 passed the Reservation headquarters headed nortli. Many
seemed to be in pairs. There was a distance of hundreds of yards
between the vanguard and those bringing up the rear. They were
moving steadily but unhurriedly, in flights mostly less tlian 100
feet between perches, except where there were open stretches with
no perches available. Like other migrating groups this one was
relatively silent, but with some low conversational notes between
nearby individuals. Each individual maintained contact with the
group mainly by sight. Most of the group passed through an elm
at the Reservation headquarters where a female was incubating.
Although some of the migrating jays passed within a few yards of
the nest, neither the female nor her mate, nor any other resident
jays in the vicinity paid any attention to the migrating flock.

Transients moving south in autumn are less easily identified,
because the residents are restless and noisy at that season, and their
numbers are augmented by the annual brood. Southward migration
probably occurs mainly in late September and early October —
beiore the advent of cold weather. On September 27, 1954, while
approximately a dozen blue jays were foraging in oaks near the
pond, 27 more came straggling down the slope from the north and
joined them, with much commotion. Obviously these new arrivals,
at least, were migrants, as there were not this many jays living in

Home Ranges of Vertebrates 237

the vicinity. In 1957 migrating groups were seen on September
27 and October 9.

In the winter of 1953-1954, when the population of jays was low,
there were three distinct groups on the Reservation, which kept
to separate ranges, one group in the northeastern part, another in
the southeastern part, and a third in the western part ranging also
onto adjacent farmland farther west. Most observations were made
on this latter group, which consisted of approximately eight pairs.
The members of each pair usually stayed near together, or at least
maintained contact by sight or sound. The whole group also tended
to move about as a unit, and each individual maintained contact
with the main group by occasional calls and answers, even though
at the time the individual might be foraging independently, hun-
dreds of yards from its companions. The precise limits of activity
for this group were not determined, but the area encompassed by
their usual activity was measured roughly as 3000 feet from north
to south and 2000 feet from east to west. It consisted of two
wooded hills, with an intervening valley of fields and pasture, 300
yards across. The jays spent most of their time on one or the
other of the wooded hills, crossing the intervening valley fre-
quently, and pausing en route to perch in high isolated trees or
groves. Occasionally they foraged along creekside thickets in this
predominantly open area. Jays were often seen to fly M to % mile
from one hillside to another.

With the onset of the breeding season, flocks such as the one
described did not disintegrate completely. Although largely pre-
occupied with its mate and nest, each individual continued to par-
ticipate to some extent in group activities. Two or more pairs often
joined each other in foraging or maneuvers which were for the
most part amicable. Each year jays nested in the grove near the
Reservation headquarters. On several occasions a jay incubating
at this place was seen to fly off its nest to a hilltop 1150 feet west
in response to the calls of others on the hilltop, and it joined a
group there. The calls to which it responded were not indicative
of any unusual disturbance. In the process of nest building, jays
were seen to bring sticks the full distance from this same hilltop,
but more often twigs were obtained nearer the nest site, sometimes
from the same tree where the nest was situated. After a fledgling
had left the nest, but was still unable to follow the adults, the latter
were seen to bring food regularly from a hillside approximately 400
feet from the young bird's hiding place beneath the nest tree. In

238 Unr-ersity of Kansas Publs., Mus. Nat. Hist.

June, 1956, an adult was seen to bring food to a nest in this same
tree from a distance of 900 feet.

In die nesting season instances of intolerance observed from time
to time, did not seem to involve defense of a well defined territory.
However, the nesting jays were intolerant of other pairs courting
or lingering in the nest tree or in nearby trees. On May 11, 1955,
while incubation was in progress in one nest at the Reservation
headquarters, a pair of jays came into a large elm 90 feet from the
nest tree, and were met there by the resident pair. The beginning
of the encounter was not seen. When noticed, all four jays were
mo\dng about through the tree rapidly, uttering low calls. After
several minutes of maneuvering, they shifted to the nest tree where
the scolding and movements became more animated. A defending
jay would dart at an intruder and follow it persistently for several
seconds. A variety of squawks, squalls, and scolding sounds were
uttered. Finally the intruding pair was put to rout, and with
raucous calls they flew off together headed northeast.

In time of stress, in the presence of a natural enemy, nesting jays
were completely tolerant of neighboring pairs, and two or more
pairs might combine in mobbing it. On June 7, 1955, excited call-
ing drew my attention to a large black rat snake which had climbed
to a jay's nest and had begun to eat the young. Within a few
seconds the scolding adult jay was joined by its mate, and both set
up a frantic clamoring. They closed in, and actually attacked the
snake, striking it many times and staying within a foot of it through-
out most of the encounter, though leaping back warily at each
movement it made. After approximately one minute a second pair
of jays arrived. They participated in the heckHng, but did not
attack, and stayed at a safe distance, several feet away. They left
after approximately two minutes, while the snake was still on the
nest.

Corvus brachyrhynchos Brehm

Crow

Status. — Common resident; those that breed on Reservation re-
main in vicinity throughout year, with influx of migrants from
farther north in winter.

Habitat. — During annual cycle crows use every available habitat,
exploiting any available food source. In general, however, they
probably utilize open fields more than woodland, and cultivated
fields in preference to pastureland. All nesting sites discovered
were in thick woodland.

Home Ranges of Vertebrates 239

Movements. — Resident crows generally kept apart from the large
flocks of wintering crows, and the two groups were notably diflFerent
in habits during the part of the year when both were present. In
autumn the population of migrant crows moving in for the winter
built up gradually over periods of weeks, as numbers were aug-
mented by new arrivals. In several different years flocks of re-
cently arrived wintering birds were first noticed on the following
dates: October 17, 1950; October 3, 1951; September 18, 1952;
September 14, 1953; October 26, 1957. The wintering crows were
especially gregarious. In roosting they congregated in flocks which
sometimes had thousands of individuals. Each morning there was
partial dispersal of the roosting aggregation, and the flocks of vary-
ing sizes scattered over an area as large as ten square miles. As the
crows proceeded with their morning foraging, the large flocks dis-
integrated into progressively smaller units. In late afternoon the
crows became increasingly restless, tending to gather at rallying
points such as tall trees on hilltops. Usually before sunset these
groups began homeward flights toward the roosting area, strung
out in long straggling files. Roosting procedure was much influ-
enced by the weather. Rain, snow, and wind interfered with the
normal foraging activities, and caused the crows to congregate in
the roosting area long before dark, as early as 3 P. M. on midwinter
days. Prolonged maneuvers preceded the actual roosting, as hun-
dreds or even thousands of individuals milled about, calling, flying
and perching together. These maneuvers were notable for their
seeming aimlessness and lack of leadership, often with indecision
up to the last moment regarding choice of a roosting site. Often the
main group would split, one division moving away hundreds of
yards or more to a different roosting area. Movement from one
group to the other might begin with a lone individual or a pair, fol-
lowed by a trickle and then a steady stream and finally a mass
movement as those remaining merged with the rival flock. Oc-
casionally there might be simultaneous interchange between two
such large groups, with uncertainty as to which would eventually
be drained into the other. Actual roosting sometimes occurred
soon after sunset but in other instances it was delayed until almost
total darkness. Roosts were in dense thickets of osage orange, or
less frequently, in thick honey locusts. The crows flew down
rapidly from their hovering over the roost or from tall trees nearby,
into the thicket.

Many of the wintering crows seemed to be in pairs as they for-
aged and moved about during the day, sometimes within a group

240 University of Kansas Publs., Mus. Nat. Hist.

and sometimes more or less independently. In the evening flight
toward the roost, pair-associations were especially noticeable. In
maneuvers over the roost or nearby, pairs were even more readily
discernible. In a hovering flock individuals would often swoop at
others nearby. The one followed would bank sharply downward
in a long swoop, and then carried on by its momentum, would rise
abruptly to approximately its former elevation, closely followed by
the partner, which maintained appro.ximately the same spacing
throughout the maneuver. In some instances three individuals
seemed to be associated; such groups perhaps included a young
of the previous nesting season still remaining with its parents.

Throughout April the numbers of wintering crows dwindled, but
usually some stayed on into May. The latest dates when sizable
roosting aggregations were seen in several different years were
May 4, 1952, May 11, 1953, and May 21, 1956. Crows were known
to come for as much as three miles to roost. It is doubtful whether
any came from much farther. The wintering flocks varied greatly
in numbers from year to year, but it was not determined whether
these variations represented widespread trends in population or
whether they resulted merely from shifts to other roosting areas.
A roosting aggregation estimated to contain at least 2000 crows was
first discovered on December 25, 1950, in a dense osage orange grove
a quarter mile southeast of the southeast comer of the Reserva-
tion. By February 8, 1951, this aggregation was roosting a half-
mile farther east. In October, 1951, both these sites were utilized,
and smaller groups that spht off from the main aggregation roosted
at greater distances. The crows seemed reluctant to enter any
roost, and were inclined to panic at any alarm. On several occasions
when it was nearly dark, after crows had already settled on the
roosts, they were heard cawing in excited confusion, and were seen
erupting in all directions, flying low and rapidly, and continuing on
out of sight. Presumably they then roosted singly or in small
groups far from the communal roost, which was subsequently
shifted to a new site. On the roosts they were especially subject to
predation by a variety of flesh-eaters, notably the horned owl.
Evening flights toward the roost were often interrupted when one
of these owls hooted, inadvertently drawing attention of the crows
to itself. As the owls were most vocal between sunset and dark, this
mobbing was an almost daily occurrence. A crow having spied
the owl, swooped down toward it cawing excitedly and causing
others streaming past to join it. Soon hundreds might congregate to
heckle the owl, but they continued on their course within a few
minutes.

Home Ranges of Vertebrates 241

In spring long before the wintering crows left, the resident crows
began nesting. Each pair selected a nest site in thick woodland
and spent much of each day in the vicinity. The onset of the breed-
ing season was indicated by the peculiar squalling calls in the
woods where courting pairs had settled to nest. Over several years
these calls were first heard on March 25, 1953, March 22, 1954, April
18, 1955, March 31, 1956, and April 26, 1957. The nesting crows
were non-territorial, and were social in many of their activities.
Foraging was carried on chiefly at a distance from the nests, in open
land, and several pairs from the same nesting area or from different
areas, might mingle freely in their search for food. In several dif-
ferent years, for much of the nesting season, a small patch of al-
falfa in a field west of the Reservation was observed to be a favorite
feeding ground. Insect prey, perhaps scarabaeid beetles, seemed
to be abundantly available at this place, which was approximately
% mile south of a slope covered with a dense grove of osage orange.
Crows that nested in the grove spent much of their time in the
alfalfa patch, and made the round trip to it and back many times
each day.

Often two or three or four pairs of crows nested in the same part
of the woods, within a distance of 100 yards. An individual might
guard the entire nesting area, and spread the alarm at any dis-
turbance. In the summer of 1952, in a 200-acre woods in the western
and northern part of the Reservation, five distinct nesting areas
were noted, with a total of approximately 20 adult crows.

Panis atricapillus Linnaeus

Black-capped Chickadee

Status. — Abundant resident, present on all parts of the area where
trees or tall shrubs are present.

Habitat. — Forest.

Much of the activity of this species is in American elms, these
being the most abundant trees of the area, but clear-cut preference
is not evident, and all available species of trees and large shrubs
are used. The population of chickadees tends to concentrate along
edges of woodland, especially where the adjacent open land is in an
early stage of succession, with coarse weeds such as sunflower and
giant ragweed, their seeds providing a food supply. Presumably
the presence of tree holes for roosting places is a limiting factor.

Movements. — Chickadees are social at all times of the year, and
are nearly always seen with a group, or, in the breeding season, a
mate. MOd sunny days in late winter initiate pairing and terri-

242 University of Kansas Publs., Mus. Nat. Hist.

torial behavior. Territorial calling was first heard on February
11, 1952, January 26, 1953, and January 29, 1954. By early Febru-
ary', pairs may separate from the flocks and keep to themselves,
temporarily, but cold, wet or windy weather causes recombination
into groups. On March 20, 1952, a group of six were foraging to-
gether in a small ragweed patch, with no indication of pairing or
of hostility, except that frequently they gave calls "zr-wick-ick"
usually associated with territoriality. In early March trios of chicka-
dees have been observed in prolonged quarrels and maneuvers,
as tu'o competed for the same mate, each competitor scolding
persistently and striving to keep between the mate and the op-
ponent.

In late March, April, and May, 1953, the establishment of a
territory, and part of the nesting cycle were observed in a pair in
the vicinity of the headquarters. On the morning of March 26 the
pair was observed moving about silently in what was later recog-
nized as their territory. One entered a cavity in a dead snag
where bluebirds were nesting, and was instantly routed by them.
On March 30, both chickadees were in the top of an elm 150 feet
farther north, and were staying within a radius of a few yards.
Catkins of the elm provided an abundant food supply. An in-
clined dead limb approximately two inches in diameter had a
decayed spot near its end, and the chickadees were persistently
working on this trying to enlarge it into a cavity for nesting. As one
pecked at the soft wood, removing small fragments, the other
would forage nearby. At frequent intervals the one at work was
relieved by its mate, and took a turn at foraging. Wlien another
pair flew into a large elm 80 feet away, giving territorial calls, the
pair under observation promptly flew toward them and engaged
them in a quarrel which consisted chiefly of maneuvering at close
quarters in the top of the tree, with much scolding. The commo-
tion attracted a third pair and then a fourth, and for a quarter
hour the scolding birds milled about through several nearby trees,
including the one where the nest cavity was located. After they
dispersed, the pair under observation promptly resumed work on
the nest cavity. Later the same day another territorial melee
similar to the first one was observed. It seemed that territorial
boundaries were as yet ill-defined, and individuals that attracted
attention in intermediate areas were harrassed by others until
they retreated, or until the quarrel was terminated by mutual with-
drawal. If such quarrels were prolonged, they usually attracted

Home Ranges of Vertebrates 243

other pairs. Many three-pair and four-pair quarrels have been
witnessed in late March and early April.

On the morning of April 2 work was still in progress on the
same nest cavity, but several times that day one of the chickadees
was seen to fly down to a five-foot log leaning against the side of
a building and peer in a cavity near its upper end. This was ap-
proximately fifty feet northeast of the nestsite originally chosen.
On the morning of April 3 the pair had definitely abandoned the
original nest cavity, on which they had made little progress in
four days of excavation. They were visiting the new cavity every
few seconds. Although it appeared to be of ample size, with an
entrance VA inches in diameter and a depth of several inches, the
chickadees spent most of their time from April 3 through April
9 enlarging the cavity. Carrying of nesting material was first
observed on April 13 and last on April 23. The carcass of an
opossum approximately 100 feet from the nest was the source of
most of the nesting material. Many times the chickadees were seen
flying with billfulls of hair from the carcass to the nest. On other
occasions they visited a blackberry thicket some 200 feet south of
the nest, and brought back tufts of cottontail hair. The total area
covered by this pair during nest building and incubation was com-
puted as 2.3 acres, and was a little more than 600 feet in length.
Actually the outlying parts of this area were visited rarely and they
overlapped broadly the areas of at least two other pairs. Activity
was concentrated in two elm trees within a 70-foot diameter, an
area of approximately .1 acre. This territory was remarkably small
compared with those that Odum (1942: 526) observed and mapped
on the Huyck Preserve near Rensselaerville, New York, which
ranged from 8.4 to 17.1 acres (average 13.2). Odum found that
the size of the territory and the vigor of its defense decreased as the
nesting season progressed.

In late May, 1953, a pair of adults with fledged young were ob-
served and followed several times within a half acre area of wood-
land edge, suggesting that at this stage, with varied and abundant
food supply, and while fledglings are still unfamiliar with the ter-
rain, family groups may confine their activities to relatively small
areas.

In winter, individuals cover relatively large home ranges, usually
associated as members of a flock, and often associated, although
more loosely, with other kinds of small birds. In few instances
were a sufficient number of records obtained, for any one individual,

244 UNIVEBSITi' OF KANSAS PuBLS., MuS. NaT. HiST.

to permit outlining a home range with any degree of completeness.
One such home range, plotted on the basis of nine capture points,
was a pentagon of 19.1 acres. However, the average distance
between successive captures for a series of individuals provides the
best basis for calculating size of home range in \\'inter. A total of
56 such movements for 21 different individuals averaged 583 feet.
A home range with a radius of this length would cover 24.4 acres.
Odum (op. cit. :529) found larger home ranges in New York, with
an average size of 36 acres (21 to 55). For the individuals in my
study, successive records were obtained within a few weeks at most
and it is unhkely that extensive shifts in range occurred within the
period that an individual was trapped. In six other instances con-
secutive records of individuals were in different winters, and for
these, distances averaged 800 feet, indicating that there is generally
some reshuffling of ranges after the summer interval.

In winter mixed flocks of small birds often forage together, and
three kinds or even four or five kinds are often found in close asso-
ciation. The black-capped chickadee is perhaps the pivotal mem-
ber of such associations. At least it is usually the most conspicuous
species, and being less specialized than some other kinds in its
foraging habits, it may function as an intermediate link to hold
them together. Wing (1946:511) listed nine species-combinations
in such winter aggregations. He stated ". . . it seems to me
that the company sticks together day after day." At times aggrega-
tions have been followed by me for hundreds of yards, and have
been noted to change the rate of travel or to reverse directions
abruptly without losing their unity. Nevertheless, it is obvious
that the associations are temporary. On many occasions when the
travelling groups have been followed, individuals or species have
dropped out after a short time. Actually, the associations are
ephemeral and change rapidly. The tufted titmouse is the most
consistent associate of the chickadee. For both parids, seeds of
giant ragweed are a favorite winter food, and mixed groups often
forage together where there are ragweed patches near the edge of
woodland. Instances of incompatibility between the two species
are rare and have been observed only when a titmouse and chick-
adee were competing for a roosting place, in a cavity of a hollow
limb. In mixed groups there are almost always more chickadees
than titmice. The chickadees tend to forage more rapidly, and to
cover larger areas, and sooner or later they leave the titmice behind.
Foraging habits are somewhat similar, but at times the titmice do

Home Ranges of Vertebrates 245

much of their foraging on the ground, while chickadees rarely come
to the ground. Golden-crowned kinglets appear on the area ir-
regularly, but are frequently in association with chickadees. In
such mixed groups, the kinglets tend to move more rapidly and
leave the chickadees behind. Downy woodpeckers and brown
creepers also frequently forage in association with chickadees, but
both kinds move about more slowly than the chickadees and tend
to be left behind. Juncos and myrtle warblers at times associate
with chickadees but less persistently than the other species men-
tioned.

Parus bicolor Linnaeus

Tufted Titmouse

Status. — Common resident in woodland.

Habitat. — Forest, with tendency toward preference of oak-hick-
ory-elm type especially in summer; similar to the chickadee in
general habitat and in preference for edge situations which provide
a food supply of weed seeds, and are frequented in winter especially.

It di£Fers from the chickadee in doing much of its foraging on
the ground, and it often forages on exposed soil in sheltered spots
such as cut banks of gullies at the edge of the woods. Like the
chickadee, this species requires tree holes for roosting and nesting,
and in some situations the absence or scarcity of holes may consti-
tute a limiting factor.

Movements. — An individual titmouse ordinarily remains year-
round in the same general area, varying its home range according
to the seasonal availability of food. Changes in population density
and in intolerance according to season, are also important factors
controlling the size of an area occupied by an individual.

Territorial calling usually is initiated by unseasonably warm
weather in January. For example, on January 4, 1956, a clear day
with temperature in the mid-fifties, singing was heard several times.
On January 11, 1951, an unusually mild day (74° F. ), singing was
frequent. Individuals well spaced through the woodland were
calling and answering each other at regular intervals. Such early
territorial activity was quickly terminated by the return of in-
clement weather. However, by the end of January or early Febru-
ary, territorial calling was more easily evoked, and might occur on
clear mornings even when air temperature was at or near freezing.
In February territorial behavior became increasingly prominent.

At times of year other than the breeding season, titmice are

246 Univ'ersity of Kansas Publs., Mus. Nat. Hist.

somewhat social in their habits. In summer, family groups of
fledged young and their parents, move about together. In autumn,
titmice are usually found in associations of two to eight individuals.
In most instances these, too, are perhaps family units that have
maintained their identity since the nesting season. However, some
of the larger groups must include either successive broods reared
by the same parents in the course of the season, or composite as-
semblages from different families. At all times of year there is some
intolerance for individuals other than the mate or the social group.
Seeds of osage orange made available by fox squirrels tearing apart
the "hedge balls" constitute an important part of the diet in winter,
especially when the ground is snow-covered. An isolated osage
orange tree in a field near the headquarters was a favorite feeding
place in February, 1952, because of its heavy crop of fruit. On
February 6, a group of three titmice, recognizable by their color
bands and often seen together, were followed to the tree. After
they had fed there for several minutes, two others came to the
tree from another direction. Immediately one of the original group
darted at one of the newcomers and chased it rapidly from branch
to branch for several seconds, causing it to leave, and it flew to
another tree 150 feet south. In the next few minutes it returned
time after time to the osage orange tree, but each time it was soon
driven off by the hostile member of the original group. When
driven away it retreated to a tree 150 feet south, or to one 50 feet
north or a third 250 feet west. Several times, flying toward the
osage orange tree it was intercepted by the defending titmouse and
turned back. The pursuit was accompanied by much scolding and
calling by both titmice. Hostility was notably lacking in the other
three individuals.

Outright fighting has not been observed, but territorial encoun-
ters are frequent in spring. These may involve two individuals of
different pairs, or may involve as many as three pairs simultaneously.
With loud and persistent scolding, the opponents would hop and
flutter from twig to twig, within a few inches of each other, each
attempting to block the other's progress into its own territory.
Such quarrels might take place frequently at about the same place
over periods of days, when some attraction such as a potential
nest site or food source happened to be situated near the territorial
boundary.

Size of territory is subject to much variation. In late February,
March, April, and May, when territorial calling is prominent, indi-

Home Ranges of Vertebrates 247

viduals in adjacent territories answer each other regularly. In 24
instances approximate distances between such calling titmice on
adjacent territories were recorded. These distances ranged from
190 feet to 900 feet, averaging 484. Assuming this figure represents
the average distance between centers of adjacent territories, the
average territory would cover 4.2 acres. In April and May, the
nesting season, individuals or pairs were often kept under observa-
tion for periods up to several hours. For 19 such observations the
most distant points reached by individual titmice ranged from 250
to 950 feet apart and averaged 400 feet. Assuming that the latter
figure represents the average diameter of a territory, the territory
would average 2.9 acres. This is a minimum figure, because an
individual watched for a relatively short time could not be expected
to cover the full extent of its territory. A typical territory probably
is somewhere between the two average figures obtained (4.2 acres
and 2.9 acres ) or about 3/2 acres. One pair observed frequently in
June and July, 1952, while they were feeding young, covered an

Fig. 17. Winter range of a tufted titmouse, recorded at 26 places from
December 21, 1954, to January 20, 1956. A relatively small inner area shown
enclosed by crosses, comprised the usual home range, but occasionally the
bird wandered farther. All parts of this winter range were used also by

many other titmice.

248 University of Kansas Publs., Mus. Nat. Hist.

elliptical ten-acre area, approximately 600 feet in length and 220
feet in breadth. The young were banded on June 29, when they
were nearly fledged. Although still unable to fly, they had remark-
able climbing ability. When released on the ground each would
make for the base of a tree, and gripping the rough bark firmly,
would walk up the vertical trunk, fluttering to maintain its position.
Three days later all tlie young had left the nest and were in the
troetops, all within a 50-foot radius some 200 feet from the nest.
Although still unable to fly strongly, the fledglings seemed to be
at ease, as they were preening and frequently fluttering from branch
to branch. The adults were foraging in different directions, and
frequently returning to deliver food.

In winter, individuals and groups range more widely than in
spring and summer. Average size of the minimum home ranges
measured varied according to numbers of marginal locations, as
follows: 4 points (8), 10.4 acres; 5 points (14), 10.5 acres; 6 points
(7), 13.9 acres; 7 points (2), 11.3 acres; 8 points (2), 19.7 acres.
Fig. 17 shows the known range of the individual for which the
largest number of records were obtained. For 16 individuals each
live-trapped in seven or more locations, the areas encompassed
varied from 43.9 to 3.2 acres but were mostly in the range, 9 to 15
acres and averaged 14.4, These are minimum home ranges in most
instances, but several for which the higher figures were recorded
may have shifted within the time span of the records. For 34
other titmice that were live-trapped fewer times, a total of 80
movements recorded between consecutive points of capture, aver-
aged 466 feet, indicating home ranges of 15.5 acres.

Sitta carolinensis Latham
White-breasted Nuthatch

Status. — Rare and irregular visitor.

Habitat. — Typically oak woodland.

Movements. — One was recorded on the area in 1949, by R. W.
Fredrickson. The only one recorded by me on the area was first
seen on November 24, 1955. It was seen and heard frequently in
the next three months, and was color-banded in December. Its
isolation from others of its species together with the fact that it
was banded, made individual recognition easy. It was often ob-
served and followed, and was live-trapped several times. Its known
range was a woodland slope of northwest exposure, with mixed
growth, chiefly of American elm, chestnut oak, and shagbark

Home Ranges of Vertebrates 249

hickory. It showed no definite preference for any one kind of tree,
but seemed to forage on any kind that happened to be in the
vicinity. On several occasions it was seen in a large elm at the
Reservation headquarters. It usually spent many minutes in a
tree before moving on to the next. On one occasion it was followed
on a circuitous course 740 feet between most remote points, and,
on another occasion, 500 feet. The hexagonal area encompassed
by the records over the three-months period was measured as 37.1
acres, of which approximately one-sixth was open field, the remain-
der woodland. The area was approximately 2000 feet long. The
nuthatch may not have ranged over all of this simultaneously as
only the later records were in the southern half of it. Usually this
nuthatch was seen moving about alone, but at other times it was
associated with creepers, chickadees, and titmice.

Sitta canadensis Linnaeus
Red-breasted Nuthatch

Status. — Rare and irregular winter visitor.

Habitat. — Coniferous forest.

Movements. — R. W. Fredrickson recorded this nuthatch from the
Reservation in 1949 or 1950. On September 11, 1954, one was
calhng near the headquarters, and on October 8 one was heard
700 feet northeast. On October 22, 1954, two were heard in wood-
land at a hilltop just west of the Reservation boundary. Probably
different individuals were involved in each instance. There are
no conifers on the Reservation, and the only evergreen trees
present are red cedars (Juniperus virginiana) of bush size. Win-
tering nuthatches therefore usually do not stop on the area, and the
fact that several were recorded in the autmn of 1954 suggests an
unusually heavy wave of migration then. Another was seen on
October 1, 1957.

Certhia familiaris Bonaparte
Brown Creeper

Status. — Moderately common in winter.

Habitat. — Woodlands throughout the entire area; most often seen
in denser type of woods but little preference for any particular kind
of tree has been noted.

Movements. — Creepers are present on the area each winter.
However, because of their inconspicuousness, secretive habits, and
small numbers, they are not regularly observed. Early autumn rec-

250 Uni\'ersity of Kansas Publs., Mus. Nat. Hist,

ords are October 13, 1948; October 23, 1955; and October 14, 1956.
Latest spring records are April 9, 1953; April 6, 1954; April 4, 1956.

An individual was often seen frequently over periods of days or
weeks in tlie same tree. Most commonly each individual was soli-
tary but sometimes two were seen together and one might follow
the other as they shifted from tree to tree. In early February, 1955,
a group of at least four roosted together regularly on the stoop of the
laboratory building. In late afternoon, or at any time in the day
when particularly inclement weather, such as sleet, snow, or high
wind, interfered with normal foraging, the members of this group
would suddenly appear on the outer wall of the building, and
climbing up the rough siding, would cluster in a compact mass at
the top. The well protected spot selected for this aggregation was
only a few inches from the comer of the door at the entrance of the
building, and persons passing in and out often disturbed the creep-
ers. Probably as a result of this frequent disturbance, the creepers
deserted this roost in favor of one in the bole of a hollow elm 140
feet northeast of the building. Somewhat similar observations on
roosting were recorded by Stone (1950:391) at Schenectady, New
York, on February 27, 1949. He observed two come to roost at 4:15
P. M., climbing stucco siding to the point where a chimney met the
overhang of a roof, partly sheltered by an old mud-dauber's nest.
The birds roosted snuggled together, with their bills piercing the
wood of the roof. According to Stone's informants, the birds had
been using this roost for more than a month.

On most occasions the creepers seen on the Reservation were as-
sociated with other small insectivorous birds, including black-
capped chickadees, tufted titmice, downy woodpeckers, and golden-
crowned kinglets. These associations seemed to be extremely loose
and fluid. Because of differences in rate of foraging, the birds of
one species might soon move out of sight and hearing of the others.
Each kind was attracted to the group whenever it saw or heard them.

Individual creepers were often followed for an hour or more in
order to determine the extent of the area covered in their daily
movements. The creeper was usually soon lost to the observer. Its
inconspicuousness, and faintness of its utterances, which were
audible only at close range, made it especially difficult to follow.
On November 30, 1957, one was followed from the creek near the
Reservation headquarters through woodland on a north slope to a
hilltop approximately 800 feet from the starting point, and then part
way back down the slope.

Home Ranges of Vertebrates 251

Troglodytes troglodytes (Linnaeus)
Winter Wren
Status) — Rare winter migrant.

Habitat. — Well sheltered places such as thickets and ravines are
required.

Movements. — One was seen on the Reservation in December,
1948, but none has been recorded since then.

Troglodytes aedon Vieillot

House Wren

Status. — Present each year only briefly after arrival from north-
ward migration in spring, and before departure in autumn, but
there are no breeding records from the Reservation.

Habitat. — Mainly woodland edge, especially where there are
thickets, gullies with exposed tree roots, or rock ledges, providing
an abundance of niches and crannies such as this species requires.

Although numerous singing males in apparently established ter-
ritories are present in early summer, they soon leave the area. It
is not fully evident what habitat deficiencies cause this exodus;
scarcity of low cavities of the type required for nest sites is prob-
ably one factor but it cannot wholly explain the failure of these
wrens to breed on the area.

Movements. — Over a six-year period, 1952 through 1957, wrens
have first appeared in April, on the 28th, 23rd, 8th, 15th (approxi-
mately), 25th and 10th. Each year in a period of a few days after
the first one was noted, numbers increased rapidly, and the song
was heard frequently. Singing males seemed to be settling in
breeding territories but within about three weeks all were gone,
ordinarily. In 1956, an exceptionally late one was heard singing
on June 2, but none had been heard previously since May 19. None
has been noted on the area in July or August. In September there
is an influx of wandering house wrens, perhaps already started on
their southward migration. They were first noticed on September
14, 1948, September 8, 1953, September 11, 1954, September 20,
1955, and September 14, 1957. Such individuals may settle for
periods of days. From September 15 to 18, 1953, a wren thought
to have been the same individual was seen repeatedly in a weedy
gully within a distance of approximately 100 feet, and it would turn
back when driven to the hmits of this small range.

252 Universiti' of Kansas Publs., Mus. Nat. Hist.

Thryothorus ludovicianus (Latham)
Carolina Wren

Status. — Resident in woodland; because of its specialized require-
ments, total population amounts to few pairs or family groups. Pop-
ulations fluctuate noticeably, and, as this locality is near north-
western extreme of geographic range, intolerance to cold and food
scarcity occasioned by winter storms are perhaps important limiting
factors.

Habitat. — Woodland and "edge" where there are streams or
gullies with steep, cut banks, and exposed tree roots are preferred.
Both spreading of the forest and severe eroding of gullies presum-
ably have improved habitat conditions for this wren.

Movements. — Carolina wrens have been observed in pairs at all
times of year. The pair-associations and the territories occupied
are probably permanent for as long as both birds survive. These
wrens sing more consistently throughout the year than do any other
birds on the Reservation. On sunny days, even in midwinter, song
may be heard frequently. The territory of a pair may be so remote
from those nearest it that ordinarily the neighbors are neither seen
nor heard. In most observed instances territories were well sep-
arated, but some were adjacent. Such territories were in contact
for only a small part of their circumferences, because the two pairs
ranged along different stretches of the same gullies and their ter-
ritories were placed end to end. When numbers were relatively
low, territories were usually not adjacent and a pair might take
over the former territory of an adjacent pair, in whole or in part.
In six instances in which different adjoining males were heard
answering each others' songs, distances between them, in feet, were
approximately: 1300, 1200, 1100, 900, 800, and 580.

The gully of the intermittent creek near the Reservation head-
quarters provided the axis for a territory which was occupied
through most of tlie period covered by my observations. Over a
six-year period the occupants were often replaced by newcomers as
shown by the disappearances of several banded individuals. The
total number involved is unknown but probably on the average
each pair was replaced at least once annually. Nevertheless this
territory tended to maintain its general conformation. At one end
a broad dike across the ravine, and tlie pond behind it, formed a
territorial boundary seldom crossed by the wrens. In the other

Home Ranges of Vertebrates

253

Fig. 18. Territory of a pair of Carobna wrens, as revealed by the male's
singing stations, in 1953, 1954, and 1956. Each year the territory extended
along a wooded ravine. In 1953, when by far the most complete records
were obtained, activity was most concentrated in the relatively small central
portion. In 1953 the wrens occasionally crossed the narrow east end of the
field to the woodland edge thicket on the north side, while in other years
they kept on the south side of the field.

254 University of Kansas Publs., Mus. Nat. Hist.

direction, where the gully emerged from woods into almost flat,
open fields, and abruptly changed in aspect, with lower less eroded
banks, the habitat became less favorable, to the exclusion of the
wrens. In 1952, 1953, 1954, and 1956 the length and area of this
territory were measured as 1100 feet, 5.8 acres (from 6 points);
1150 feet, 9.2 acres (from 9 points); 1150 feet, 3.9 acres (from 9
points); and 1250 feet, 7.6 acres (from 10 points), respectively.
Singing and other activities of the wrens tended to be concentrated
in a relatively small central area of blackberry thickets, logs and
steep eroded banks with exposed tree roots. The gully along which
this territory was situated was in woodland edge. A hillside ex-
tension of this same woodland converged to join that of the gully
near the pond ( Fig. 18 ) . A grassy field lay between. In 1953 the
wrens often crossed the narrow end of this field to the edge of the
hillside woodland. A distance of about 250 feet was the maximum
that they essayed to cross, and even then they were likely to pause
en route in isolated trees or shrubs. Failure to cross the field where
it was wider emphasized the secretive habits of these wrens.
Brewer (1955:141) recorded average areas of only .3 acre (.12 to
.61) covered by three Carohna Mo-ens in a swamp thicket habitat
in southern Illinois.

Chapman ( 1947 ) who observed several nesting pairs in La Salle
Parish, Louisiana, noted that the feces of the young were dropped
by the parents 50 to 100 feet from the nests. Coin and Coin
(1954:59) noted that in carrying any but minute objects to the nest,
the wrens would climb to an elevated perch to gain altitude before
taking ofiF. They cited instances of wrens climbing 14 feet to fly
44 feet, and 16 feet to fly 49 feet.

Wrens were absent from this area from mid-February to late
June in 1952; from mid-May until late December in 1954; for many
weeks in late winter and again in early summer in 1955. These
absences from the area of favorable habitat were correlated with
general reductions of the population, which perhaps resulted from
the decimating effects of severe winter storms.

Trautman (1940:318) at Buckeye Lake, Ohio, noted that in un-
usually cold weather Carolina wrens often shifted from their regular
territories in partly exposed situations to neighboring woodland.

In late August and early September, 1953, the two adults in the
territory near the Reservation headquarters were accompanied by
three fledglings. When first observed, on August 19, this group
stayed in a thicket area only a few yards in diameter, but within a

Home Ranges of Vertebrates 255

few days the group was moving about over the entire territory, still
tending to remain near together.

In the winter of 1955-1956 the resident pair roosted huddled to-
gether in an old phoebe nest under the eaves of the garage at the
Reservation headquarters which was near the center of the terri-
tory. The previous summer they had nested successfully in an
open can lying in a cardboard carton inside this same building.

Cistothorus platensis (Latham)
Short-billed Marsh Wren

Status. — Rare migrant.

Habitat. — Usually associated with rushes, reeds or tall grass in
riparian situations.

Movements. — On April 9, 1957, one was seen at the small pond
on the Rockefeller Tract. Cover was limited to a strip of low smart-
weed approximately 15 feet wide and 100 feet in circumference.
Several times the wren permitted my approach to within 15 feet
before making a short flight. It was seen several times that day
and again on the following morning, but late in the second day it
had disappeared.

Mimus polyglottos (Linnaeus)
Mockingbird

Status. — Occasional wanderer, chiefly in spring.

Habitat. — Open situations, with groves or scattered trees are
favored. Deficiencies of the Reservation as a habitat probably result
from the too dense growth of vegetation — both the herbaceous
cover in the more open situations and the trees and shrubs in the
woodland.

Movements. — My only definite records are for April 27 and 28,
1956. On many occasions mockingbirds have been seen perched
on telephone wires or in roadside thickets adjoining pastures or
cultivated fields within half a mile of the Reservation.

On April 10, 1957, one was observed on the Rockefeller Tract.
Its behavior suggested that it was still on migration. It had stopped
at a fence comer where there was a high mound of earth with
weeds and trash providing shelter. Mostly it perched silently, with
no attempt at foraging. About 15 minutes after first seeing it, I
heard it sing briefly. After about an hour, it became more animated,
and from time to time, made short, quick flights of 15 to 20 feet to

7—3117

256 University of Kansas Publs., Mus. Nat. Hist.

tlie ground nearby to catch insects. After several such sallies, the
bird flew to the nearby fence and then struck out north northeast
and flew out of sight across open rolling fields.

Dunietella carolinensis (Linnaeus)
Catbird

Status. — Moderately common spring migrant and temporary resi-
dent in early summer.

Habitat. — Thickets, blackberry, elderberry, dogwood, and sap-
lings of honey locust and osage orange, mainly at woodland edge.

Movements. — Arrival dates for several diflEerent years were:
April 30, 1951; May 1, 1952; May 9, 1953; May 6, 1954; April 28,
1955; May 6, 1956; and May 6, 1957. In 1956 several were seen at
Lone Star Lake on April 29, more than a week before their appear-
ance on the Reservation. Pairs have often been seen together in
the latter half of May, along creekside thickets between the head-
quarters and the pond, and elsewhere on the Reservation, but none
has ever been seen or heard on the area in June. They find habitat
conditions unfavorable and move elsewhere. On July 7, 1954, and
July 25, 1952, individuals were heard calling in thickets near the
headquarters. There are several records for August and even more
for September, probably resulting from a dispersal after the nesting
season. Late records, probably of individuals already started on
migration, are: October 3, 1951; October 4, 1953; and October 1,
1956.

Toxostoma rufum (Linnaeus)

Brown Thrasher

Status. — Occasionally heard and rarely seen in summer; one or
more pairs may breed on Reservation, but as yet no nests have been
recorded.

Habitat. — On the Reservation thrashers have been heard and
seen mainly on a relatively small area of a few acres, a soutli slope
with trees mostly of honey locust and osage orange, the larger ones
scattered, but with dense thickets of saplings of these species, and
of dogwood, prickly ash, and blackberry patches. For two miles
south of the Reservation, the county road is bordered by dense
thickets of osage orange, and many pairs of thrashers nest in these
thickets.

Movements. — Thrashers arrive on the Reservation each spring,
and are seen and heard frequently over a period of days or weeks

Home Ranges of Vertebrates 257

thereafter. For several years, first appearances in spring were April
26, 1951; April 18, 1952; April 23, 1953; April 10, 1954; April 20,
1955; May 3, 1956; and April 24, 1957. So far as known, thrashers
have not nested on the Reservation. Lack of open areas, with suit-
ably short vegetation adjoining thickets, probably prevents their
establishment. In 1951, 1952, and 1953 thrashers were noted only
in April. In 1954 thrashers appeared earlier than in other years,
and were seen more often than in the other five years combined.
They were seen repeatedly in June, August and September.

Turdus migratorius Linnaeus

Robin

Status. — Resident in general area, but of erratic occurrence on
Reservation, where evidently occurs only as migrant; in summer
rarely seen; those present from time to time do not stay for more
than part of day, and they seem to be dispersing juveniles or in-
dividuals that are wandering, having failed to estabHsh breeding
territory, or having completed breeding cycle.

Habitat. — Robins that stop on the Reservation use nearly all
parts of the area, at different times, depending on the season and
the local distribution of the food supply. Grape {Vitis vulpina) is
the one chief food supply that attracts robins to the area at the
time of the fall migration. Grapevines are most abundant on the
cooler, moister, north slopes, and in these situations robins con-
gregate. In early fall, with the first wave of migration, robins may
feed avidly on pokeweed fruits in open fields and edge situations.

Movements. — Robins have been recorded every month of the year,
but there is only one record for June. In July, August, and early
September they have been seen only occasionally, wandering singly
or in small groups. In early autumn there is a sudden influx of
migrants. Robins were first noticed in large numbers on October
11, 1948; October 25, 1950; October 13, 1951; September 13, 1952;
October 5, 1953; October 5, 1954; October 7, 1955; October 24, 1956;
and September 21, 1957. Through much of October, while weather
remained warm and leaves were still unshed from deciduous trees,
the robins tended to stay in well shaded places in thick woods, and
did not wander over the area extensively. At times in late October,
November, and December, the species may have exceeded in num-
bers all other kinds of birds on the Reservation. Numbers were
dijfficult to estimate because of the frequent and erratic movements.
Throughout the colder part of the winter robins were seen much

258

University of Kansas Publs., Mus. Nat. Hist.

less frequently, and at times, in late December, January, and early
February, none was present on the area. In early April robins,
usually single or in small groups, and obviously migrating, were seen
more frequendy than in February or March. On April 11, 1956, a
pair was seen near the Reser\'ation headquarters, the male singing
frequently in several favored perch trees. Foraging was chiefly on
a bare area where the soil had been turned in the installation of a
pipe hue the previous October. After remaining in the vicinity for
two weeks, this pair disappeared.

Hylocichla mustelina (GmeUn)
Wood Thrush

Status. — Moderately common summer resident in woodland.

Habitat. — Wood thrushes on the Reservation forage on leaf litter
in low, damp situations where there is a large amount of shrubby
or herbaceous undergrowth.

Movements. — Dates of arrival on the Reservation in several dif-
ferent years were: May 5, 1950; April 20, 1951; April 30, 1952;
May 5, 1953; April 25, 1954; May 7, 1955; April 26, 1956; and May
9, 1957. In 1953 they were seen and heard in abundance at Miami
County State Lake two days before they appeared on the Reserva-
tion 50 miles farther north. Singing was heard chiefly in May and
early June. Later in the season singing was more subdued, and was
largely confined to short periods around dawn and dusk, and just

Fig. 19. Territory of a wood thrush, as revealed by singing
stations of male (dots) in early summer of 1953. The territory
was in woodland edge along a streamside thicket. Occasionally
the thrush crossed the narrow east end of the field to the edge
of the woodland on the north side of the field. In the absence
of adjoining pairs, the thrush sometimes visited outlying stations,
as shown here, beyond the usual limits of its territory.

Home Ranges of Vertebrates 259

after showers. On May 12, 1952, several wood thrushes were
foraging within a 50-foot radius, and it seemed that territoriahty
was not yet well established. Usually within a few days of first
arrival a pair was established along the creek south and east of the
Reservation headquarters, to the exclusion of others. In 1953 the
pentagonal area within which the singing male ranged was meas-
ured as 12.6 acres. Actually only small parts of this area were
regularly used. The territory proper, within which the male moved
about as he sang, consisted of two separate parts, each several times
longer than wide. The combined area was computed as 1.4 acres
(1.05 and .35), both along woodland edge, with an intervening field
150 feet across. (Fig. 19.)

Hylocichla guttata (Pallas)

Hermit Thrush

Status. — Uncommon migrant in spring and autumn; occasional
in winter.

Habitat. — Woodlands, thickets and edges, with fairly open
ground for foraging.

Movements. — In 1954 hermit thrushes arrived on the Reserva-
tion around mid-October. Although they were seen on only a few
occasions, one or more were heard each morning for almost a
month, calling in thickets just south of the Reservation headquar-
ters. On December 25, 1955, one was seen in a thicket of wdld
plum ( Prunus americana ) at the edge of a field.

Hylocichla ustulata (Nuttall)
Olive-backed Thrush
Status. — Rare migrant.
Habitat. — Boreal forests.

Movements. — R. W. Fredrickson recorded one on May 7, 1949,
and several on May 11, 1950.

Hylocichla minima (Lafresnaye)
Gray-cheeked Thrush
Status. — Occasional migrant.
Habitat. — ^Woodland.

Movements. — On April 11, 1956, two were seen on a brushy east
slope in woodland dominated by American elm ( Ulmus americana).
Each time the observer approached, they flushed and moved farther

260 University of Kansas Publs., Mus. Nat. Hist.

up the slope, keeping near together, and continuing for more than
100 yards in the same direction, until finally they were lost. The
association suggested that these thrushes were already paired.
However, Dilger (1956:345) found that in this genus the male
arrives first to establish a territory, and reacts to the female's ar-
rival with hostiht)' and pursuit, which lasts for several days, grad-
ually subsiding as the male becomes tolerant of the female and
the pair-bond develops.

Hylocichla fuscescens ( Stephens )

Veery
Status. — Rare migrant.
Habitat. — Northern deciduous forests.
Movements. — R. W. Fredrickson recorded one on May 15, 1950.

Sialia sialis (Linnaeus)
Eastern Bluebird

Status. — Resident and common in general area, but of somewhat
erratic occurrence on Reservation; habitat conditions most favorable
in winter, worsening in late spring as growing season progresses,
to detriment of those individuals that attempt to breed on area.

Habitat. — Parkland, with medium to large trees for perches,
and short grass or other low ground vegetation.

Open groves of walnut and honey locust, especially on hilltops
where soil is shallow and grass is short, provide preferred situa-
tions in winter. In summer, as brome and other grasses grow longer,
and mature, bluebirds are handicapped in their foraging and they
tend to shift from the Reservation to fallow fields, stubble or grazed
pastures where, evidently, the shorter vegetation facilitates the de-
tection and capture of ground-living insects.

Movements. — Bluebirds are present on the area throughout the
year, but their numbers and local distribution vary. Regularly
each year a pair established a territory in the vicinity of the Reser-
vation headquarters, and invariably their nesting attempt failed.
Each year singing began on clear, mild days of late winter. In the
six-year period, 1952 through 1957, singing was first heard on the
following dates: March 16, February 28, February 14, March 11,
March 22 and March 12. As singing begins winter flocks break
up into pairs, and each sets up a territory. Disintegration of the
flocks is gradual, and may be temporarily halted by inclement
weather. For periods of weeks the birds may spend part of each

Home Ranges of Vertebrates 261

day on the territory and the remainder with the flock. On March
24, 1952, nearly a month after first singing, four bluebirds flying
high together, headed south southeast, came to the field near the
headquarters, and the two j^airs Ht in trees approximately 100 feet
apart. They remained most of the day, the males singing from top-
most branches of trees, and tending to keep to difFerent parts of
the field. However, from time to time all four gathered in the
same tree with no show of hostility. It seemed that all four were
members of the same flock.

In early spring the usual procedure was for the birds to remain
with the flock for the main foraging period in the morning, and
visit tlie territory later in the day. Several times in early spring
small flocks have been seen fluttering about trees with hollow
branches that oflFered potential nest sites. In such instances there
was much scolding and quarreling, and occasional short pursuits.
Territories seem to be selected originally without regard for the
presence of possible nest sites. These are investigated later. In
1953, 23 days elapsed between the time that singing started and
the first record of the birds carrying nesting material; in 1954 this
interval was 54 da vs.

In 1952, 1953, and 1954 the movements of bluebirds that were
nesting in the headquarters area were observed in detail and terri-
tories were mapped (Fig. 20). Nest sites were different in each
year but all three were within a hundred-foot radius. Location and
conformation of the territory varied from year to year, but its size
was approximately the same. Greatest diameter varied from 850
to 950 feet. Outlying areas constituting a large part of each terri-
tory were used relatively little and activity was concentrated in a
smaller central part that was only about half as broad and half as
long as the entire territory. Because the territory studied did not
adjoin others, it may have been somewhat expanded in each year
that it was observed, and the central part where activity was con-
centrated may have been more typical of the size in areas where
the birds are subject to territorial pressure from neighbors. In the
three years territories and their more intensively used central por-
tions were measured as follows: 1952, 5.4 acres and 1.9 acres; 1953,
8.6 acres and 1.5 acres; 1954, 7.0 acres and 1.4 acres.

In 1952 and 1953 fate of the nest was unknown. In 1954 the
upright log which contained the nest cavity blew down in a storm.
In 1955 the nest was robbed by a black rat snake ( a probable fate
also in the years for which no information was obtained ) . The day

262

University of Ka.nsas Publs., Mus. Nat. Hist.

Fig. 20. Territory of a pair of bluebirds, as revealed by singing
stations of the male, in 1952, 1953, and 1954. Each year the area o?

Principal activity was concentrated in an elm grove at the Reservation
eadquarters, but the distance and direction of outlying stations varied
from year to year. Invariably nesting attempts in this area failed.

Home Ranges of Vertebrates 263

after destruction of the nest the bluebirds deserted their former
territories and did not return again that season. Failure to re-nest
may have resulted in part from the deterioration of habitat condi-
tions as the season progressed, and herbaceous vegetation grew tall
and rank. Disturbance of the natural vegetation in the headquar-
ters area of more than an acre, by trampling and cutting, seemed
to render it more favorable for nesting than other parts of the Res-
ervation that were like it.

In two different years shortly after failure of the nest and dis-
appearance of the pair in the headquarters area, bluebirds thought
to have been the same, were seen in May, 1953, and May, 1955,
frequenting an area 660 yards north at the edge of the Reservation
where it adjoined a mowed field. On July 4, 1954, a family group
appeared in the headquarters area and spent much of their time
there through July, August and early September. Sparseness of
vegetation resulting from drought that spring, and unusually heavy
cutting of grass and weeds in the yard and clothesline area, prob-
ably rendered this place suflSciently favorable for occupancy by
the birds.

Laskey (1940:189) who made an extensive study of the nesting
of bluebirds in Nashville, Tennessee, stated, "Apparently boxes
should be several hundred feet apart to allow suflBcient territory for
each breeding pair. They should be set out in winter because
nest sites are investigated by bluebirds on mild days throughout
the cold season, and territorial defense starts in early spring."

Polioptila caerulea (Linnaeus)

Blue-gray Gnatcatcher

Status. — Common summer resident; many pairs breed on Reserva-
tion each year.

Habitat. — Gnatcatchers live in woodland, with preference for
more open, drier situations. Large honey locusts in fields near the
edge of woodland provide the favorite sites for nesting and forag-
ing. Large elms are the second choice. On south slopes of xeric
aspect, with scrubby growth of chinquapin oak, osage orange, and
dogwood, gnatcatchers may be abundant. They are least often
in oak-hickory woodland with a dense leaf canopy.

Movements. — Earliest recorded dates for gnatcatchers, over the
years, 1950 through 1957, were in April each year: the 11th, 23rd,

264 University of Kansas Publs., Mus. Nat. Hist.

16th, 3rd, 10th, 15th, 11th, and 23rd. Each year within a few days
after the first arrivals the species becomes common. Because of
their brisk and frequent movements, often in low or open trees, and
their almost constant calling, individual gnatcatchers could be
kept under observation more easily than most other kinds of birds,
and they were frequently followed. On May 6, 1950, a pair was
observed nest-building in the headquarters area. On the following
day this nest was completed, having been built mainly or entirely
within a 24-hour period. On May 8 the same pair was nest-building
again approximately 100 feet from the recently finished nest, which
had been abandoned for undetermined reasons. One gnatcatcher
visited the first nest and took material from it to be incorporated
in the new nest. In mid-July, 1952, a pair was observed feeding
young in a nest just south of headquarters. Foraging was chiefly
within 50 to 100 feet of the nest, either in the same large American
elm or in three adjacent trees, two elms and a walnut. In a period
of an hour and a half (3:30 to 5:00 P. M. on July 13), the Uvo par-
ents made a total of 57 trips to the nest to feed their young. The
total area encompassed by this pair in foraging for their young was
measured as 1.2 acres, but some outlying parts were rarely visited.
On July 6, 1953, another pair was observed feeding nestlings,
foraging in a willow grove 70 to 120 feet from the nest. In late
May and June, 1952, a pair observed on several occasions covered
a territory of 2.2 acres. This territory was along a wooded ravine,
with a grove on one side and several scattered trees in a field on
the other. In May, 1952, a pair was observed in a large honey
locust in a hilltop field. They seemed to spend most of their time
in this one tree, but occasionally crossed to nearby woodland.

Gnatcatchers have rarely been seen after July. The latest record
is September 4, 1953.

Rcgulus satrapa Lichtenstein

Golden-crowned Kinglet

Status. — Regular winter visitor but variable in numbers. Unlike
some other wintering birds these kinglets do not seem to settle in
one place to spend winter but are inclined to wander. As result,
numbers present at any one time vary from none at all to dozens or
even much higher numbers.

Habitat. — Woodland, especially of northern conifers, but on
Reservation in winter, most typically where trees are low and
dense, with vine tangles.

Home Ranges of Vertebrates 265

Movements. — Golden-crowned kinglets have most often been
first noted on the area in October (25th in 1950; 25th in 1951; 18th
in 1952; 14th in 1953; 27th in 1957). They have also been recorded
in November, December and January, but never in late winter or
spring. Those seen were usually in flocks of three to fifteen, and
they were often associated with brown creepers, black-capped
chickadees, tufted titmice, or downy woodpeckers, or with two or
more of these kinds simultaneously. Kinglets were never seen on
successive days in the same place. Either because they habitually
wander, or because habitat conditions were unfavorable, those that
came to the area never stayed long.

Regulus calendula (Linnaeus)

Ruby-crowned Kinglet

Status. — Regularly present for short periods in spring and
autumn on migration.

Habitat. — On the Reservation the ruby-crowned Idnglet most
typically frequents edge situations, working through willow, dog-
wood, redbud, and saplings of elm and other trees.

Movements. — Ruby-crowned kinglets are recorded on the Reser-
vation each spring (March 30 to May 16), and autumn (Septem-
ber 17 to October 27). Unlike golden-crowned kinglets, they are
solitary, and individuals tend to settle for periods of days in wood-
land edge thickets, along ravines or beside the pond. Although
none was individually marked, kinglets of this species were often
seen on successive days in the same tree or thicket, and those kept
under observation for varying periods usually remained within a
radius of a few yards, despite the fact that they were almost con-
stantly in motion. In a rocky situation just south of the old quarry,
in a thicket of tall elm saphngs mixed with redbud and dogwood
with tangles of vines, ruby-crowned kinglets were seen much more
regularly than at any other one place. Habitat conditions were so
favorable in this situation that it invariably attracted some of those
migrating across the area.

Bombycilla cedrorum Vieillot

Cedar Waxwing

Status. — Occasional transient.

Habitat. — On Reservation, woodland and edge situations.

Movements. — Definite records are November 2, 1948, November
5, 1952 (flock of ten feeding on bittersweet berries), November

266 University of Kansas Publs., Mus. Nat. Hist.

22, 1955 (flock of 15), December 20, 1955, December 26, 1949, and
in May, 1956, and April and May, 1957. Unlike those recorded in
November and December, which were obNaously wanderers, the
flocks seen in spring stayed in the same vicinity for several days.
In 1956 the flock was first noticed at 5 P. M. on May 23, in the wil-
low grove beside the pond. The birds were feeding avidly on the
chrysomelid beetle larvae that were abundant on the willow leaves,
often hovering momentarily to reach these larvae. There were
approximately 30 waxwings in the entire flock and they were scat-
tered through the grove, but tended to keep together in small groups.
This flock was seen frequently in the next four days at the willow
grove. On several occasions the flock temporarily left the vicinity,
all flushing nearly simultaneously, prepared by the rapid lisping
notes uttered for several seconds before the take-off. On such oc-
casions they passed out of sight in high rapid and direct flight, but
sometimes returned within a few minutes, obviously having not
stopped elsewhere.

Putnam (1949:148) described social nesting habits of the cedar
waxwing; each pair's territory consisted of a nest site, a guarding
perch, and a small amount of space (225, 270 and 1100 square
yards ) . The shortest distance observed between nests was 38 feet.

Lanius ludovicianus Linnaeus

Loggerhead Shrike

Status. — Resident on nearby areas, but present on Reservation
only as rare visitor.

Habitat. — Most typically closely grazed pasture or short-grass
prairie with scattered thorny trees.

Movements. — On the Reservation the chief habitat deficiency
seems to be the rank growth of herbaceous vegetation, preventing
the shrike from seeing its prey on the ground. Shrikes are notably
sedentary in habits, and in some instances at least, stay for the
entire year within the confines of the small territory. On only two
occasions was a shrike definitely recorded from the area, and in
each instance it was foraging on adjacent land and did not venture
far onto the area.

Through 1954, 1955, and 1956 a pair of shrikes, perhaps the same
individuals, were often seen perched on wires along the county road
just south of the southwest corner of the Reservation section. The
maximum range along the county road was approximately half a
mile, but on most oc;casions they were seen within an interval of

Home Ranges of Vertebrates 267

about 200 yards. The extent of the territory in other directions
was undetermined.

Throughout 1957 a shrike was seen frequently on the Rockefeller
Tract. It kept to the pasture, the vicinity of the buildings, and the
telephone line along the road. Occasionally it was accompanied by
a second shrike. Its known range was approximately 25 acres, but
at times it could not be found within this area.

Sturnus vulgaris Linnaeus
Starling

Status. — Appears on the Reservation chiefly as migrant in spring
and autumn; occasionally as wanderer at other seasons.

Habitat. — Requirements including that of hollows for nesting
sites and that of open situations permitting easy walking, are not
well satisfied on Reservation; therefore starlings never have stayed
long.

Movements. — In autumn, especially, spectacular migrations of
starlings over the area occur, and the total numbers probably exceed
those of any other species. Early season records of migration are:
September 16, 1951 (several small groups, stopping to feed in
thickets); September 27, 1953 (flock of twelve); September 20,
1954 (a single immature bird); September 30, 1956 (several
hundred). These were forerunners of the main migration, which
was usually at its peak in approximately the last week of October,
and in the first week or two of November. In this period many
thousands might pass over in one day, and migrating flocks might
be in sight almost continually. Occasional compact flocks of from
less than a dozen to several hundred individuals, maneuvered,
perched, and fed in close formation. One such flock having several
thousand starHngs remained on the Reservation most of the day
on October 28, 1957. The largest flocks flying high over the area,
were strung out on a broad front and straggling in loose formation.

On their northward migration each spring starlings were far less
conspicuous. Earliest record is February 22, 1956, when several
flocks passed over, flying high, headed northeast. On March 8,
1954, one paused briefly in a tree near the headquarters before
continuing on north.

On March 20, 1953, a male was observed quarreling with a pair
of bluebirds, in an elm having a hollow limb. This male starling
soon flew northwest and did not return. On April 1, 1952, a male
was seen moving about slowly, investigating the same nest cavity

268 University of Kansas Publs., Mus. Nat. Hist.

and others in nearby trees. On March 28 and April 17, 1954, a male
was investigating cavities in trees near the headquarters. In the
last four days of March and throughout early April, 1955, a pair
was seen frequently in the headquarters area, where they seemed
to be preparing to nest. In April, 1956, a male was seen quarreling
with a pair of red-bellied woodpeckers near tlie cavity where they
had begun nesting, but he gave up and flew away headed north-
east. On May 3, 1956, a pair seemed to be preparing to nest near
the headquarters. So far as known, all these nesting attempts were
abortive, and the starlings soon left the area.

On the Rockefeller Tract many starlings were present throughout
the summer. Their activities were concentrated about the house,
barn and sheds, and an elm witli hollow Hmbs in the yard. The
relatively open habitat made this area more favorable than the
Reservation. In their foraging the starlings often flew half a mile
or more to newly plowed fields, pastures, or other areas that were
especially favorable for finding food.

Vireo griseus (Boddaert)
White-eyed Vireo

Status. — Moderately common summer resident.

Habitat. — Brush thickets, scrubby woodland, and woodland edge.

The species of plants present seem unimportant as long as the
growth habit, with abundance of stems and leaf surface for foraging
within a few feet of the ground surface is fulfilled. Dogwood,
dwarf oak, plum, blackberry, elm and hackberry saplings, and
poison ivy are among tlie plants most frequently used on the Reser-
vation. Several or many pairs of these vireos nested on the Reser-
vation in most summers of the past few years. The numbers have
varied noticeably from year to year.

Movements. — Arrival dates have been notably variable: May 24,
1952; May 6, 1953; April 23, 1954; May 3, 1956; May 3 (approxi-
mately), 1957. In 1955 none was seen. Latest dates of record were
August 31, 1952; October 5, 1953; July 30, 1954; September 16,
1956; and September 8, 1957. In 1952 and 1953 pairs (or perhaps
the same pair) nested near the headquarters, llie nest was not
found in either year, but tlie male vireo was a consistent singer,
and his course could be easily followed as he moved about through
dense vegetation where he could be seen only with difiiculty. In
1953 a bob-tailed fledgling was found in the southwestern part of
this territory, and it could not have travelled far from the nest.

Home Ranges of Vertebrates

269

The nest of a second pair 850 feet farther west along the same ravine
was kept under observation for part of the period of incubation
and nestling development. When this latter nest was destroyed by
a predator, two cowbirds, nearly fledged, were the only remaining
occupants.

Fig. 21. Territory of a white-eyed vireo in early summer of 1953, and in
early summer of 1954. Dots indicate locations where singing male was
recorded. In both years the territory covered the same general area, dif-
fering only in details. It consisted cliiefly of streamside thicket along wood-
land edge, but included also part of an elm grove in the narrow end of a
field, and a strip of woodland edge thicket on the north side of this field.

The territory near headquarters was, in both 1953 and 1954,
crescent-shaped, following up the ravine and edge of the woods,
northwest and crossing the narrow, brushy east end of the meadow
to the woodland on the opposite side. In 1953 the territory was
measured as 6.5 acres, and in 1954 as 5.4 acres; the areas corre-
sponded throughout most of their extent (Fig. 21). In every in-
stance territories observed by me were much larger than the .33
acre territory listed by Brewer ( 1955 : 141 ) .

270 Univeesity of Kansas Publs., Mus. Nat. Hist.

In 1955 no white-eyed vireos were heard or seen anywhere on the
Reservation, and numbers must have been exceptionally low. In
1956 and 1957 the territory near the headquarters was not occu-
pied and these vireos were heard elsewhere only a few times in the
entire season. Their numbers were still low on the Reservation.

Vireo bellii Audubon
Bell Vireo
Status. — Moderately common summer resident.
Habitat. — Brush thickets in open places. Wild plum perhaps
provides the optimum habitat. Wild cherry, dogwood, fox grape,
crab-apple, and stands of saplings, of elm, honey locust, and syca-
more also are used.

Movements. — In 1948 and 1949 these vireos were observed oc-
casionally in roadside thickets at the west edge of the Reservation.
Subsequently none was seen until 1953, but thenceforth they were
present each summer, chiefly in thickets that had sprung up since
discontinuance of grazing in 1949, and were not yet suflBciently
developed to provide adequate cover in the period 1950-52, when
no Bell vireos were present. Earliest records for a four-year period
were May 5, 1954; May 10, 1955; May 13, 1956; and May 8, 1957.
The two latest records are August 17, 1953, and August 23, 1954.

Two wild cherry trees ( Prunus serotina ) at the north edge of the
Reservation, bordering a wheat field, constituted the focal point
of a vireo's territory each year from 1953 through 1956. The sing-
ing male spent most of his time in the t\vo adjacent trees, in an area
only about 60 x 20 feet, but occasionally he ventured to nearby low
thickets along the boundary fence and at the edge of the field. In
1954 this territory was measured as .4 acre. Another territory oc-
cupied each year was in a gully having a thicket of wild plum, su-
mac, prickly ash and saplings of elm and honey locust. In the
summer of 1954 the singing vireo was recorded and followed
many times and its territory seemed to be co-extensive with this
thicket, which was approximately 620 feet long and averaged 50
feet wide (.6 acre). By 1955 thick stands of saplings had grown
up in nearby parts of the field west of this gully, and the vireo
ranged through them being no longer closely confined to the gully.
By 1956 this new thicket area seemed to be the center for a terri-
tory, and the vireo was followed about over an area approximately
150 X 60 feet. Another individual, separated from this one by an
open meadow 150 feet across, was followed for 450 feet along a
woodland edge thicket.

Home Ranges of Vertebrates 271

A territory in a brushy field, in June, 1955, was measured as one
acre, but a central area comprising about one-fourth of the total
was used much more intensively than the remainder. Knowledge
of the territory of these vireos was gained almost entirely by fol-
lowing the song. As a bird moved about through thick brush or
flitted from clump to clump it was glimpsed only rarely. Females,
being relatively silent, were observed on only a few occasions.

Hensley (1950:243) recorded a territory of 3.1 acres for a pair
in Piatt County, Illinois. The area was grassland, with scattered
trees and shrubs, and it was more open than the areas where my
own observations were made. Hensley observed young 30 feet
from the nest on the day they left it, and five days later observed
the brood congregated 300 feet from the nest.

Vireo flavifrons Vieillot
Yellow-throated Vireo
Status. — Rare migrant or transient.
Habitat. — Deciduous forest.

Movements. — The only record from the Reservation is of one re-
corded by R. W. Fredrickson on May 6, 1950.

Vireo solitarius (Wilson)

Solitary Vireo

Status. — Rare migrant or transient.

Habitat. — Forest, usually of conifers.

Movements. — The only record from the Reservation is of one
seen May 3, 1952, by R. W. Fredrickson.

Vireo olivaceus (Linnaeus)
Red-eyed Vireo

Status. — Regular summer resident; except for the field sparrow
is perhaps most abundant species of this category.

Habitat. — Deciduous forest, occurring throughout the woodland
on the Reservation except that of more xeric type, as represented
on south slopes.

The trees most regularly used, in approximately their order of
importance, include American elm, shagbark hickory, black walnut,
chestnut, oak, sycamore, black oak, red oak and coffee tree. Trees
that are rarely used are honey locust, osage orange, red haw, dwarf
oak and wild plum. There is scarcely any overlapping of habitat
between the red-eyed vireo and the two other species. Bell and

272 University of Kansas Publs., Mus. Nat. Hist.

white-eyed, that nest on the Reservation. However, the warbhng
and red-eyed overlap broadly in habitat preferences.

Movements. — The arrival dates recorded for several different
years were: April 26, 1951; April 25, 1952; April 30 (approximately),
1953; April 26, 1954; April 21, 1955; May 2, 1956; April 24, 1957.
Latest records of occurrence were: August 31, 1952; October 5,
195-3; and September 6, 1956. In May and June, especially, the red-
eyed vireo is a persistent singer. An individual foraging through
the treetops might seldom be visible from the ground, but its
course could be followed by its frequent bursts of song. Such a
bird might move across its small territory several times in the course
of an hour. An individual that lived on the wooded slope just
southeast of the Reservation headquarters in 1955 was often fol-
lowed and observed in May and June. The total area encompassed
by its records was measured as 2.5 acres, but included many outly-
ing points. Most of its movements were within a smaller area of
elliptical shape, measured as .5 acre.

In 31 instances distances between singing males in what seemed
to be adjoining territories, were measured, and these ranged from
700 feet to 80 feet, averaging 346. Territories averaging approxi-
mately 2.1 acres are indicated.

Vireo gilvus (Vieillot)
Warbling Vireo
Status. — Transient and temporary resident in late spring.
Habitat. — Groves at edge of woodland or near it. In almost all
instances these vireos were in large American elms.

Movements. — The species has been recorded on the Reserva-
tion only in May and in the last week of April: May 6 to 18, 1953;
May 3 to 31, 1954; April 25 to May 12, 1955; May 2 to 7, 1956; and
April 29 to May 26, 1957. For periods of days an individual might
be heard at about the same place, as if it had settled on a small
territory. Often such a bird tended to stay within one tree. Habi-
tat conditions must have been deficient in some respect, as such
birds never remained more than a few days.

Mniotilta varia (Linnaeus)

Black and White Warbler

Status. — Known from the Reservation only as transient and
temporary resident in spring, although there are breeding records
for northeastern Kansas.

Home Ranges of Vertebrates 273

Habitat. — Deciduous forests.

Movements. — On April 12, 1951, one was seen moving about
with a group of chickadees as it flitted from trunk to trunk. Other
specific records are: May 17, 1951; May 8, 14, and 15, 1953; and
May 14, 1954. On May 8, 1953, one was seen to fly at another and
chase it from the grove south of tlie Reservation headquarters.

Vermivora peregrina (Wilson)

Tennessee Warbler

Status. — Migrant and temporary resident in late spring; perhaps
crosses area on southward migration in autumn also, but no specific
records have been obtained.

Habitat. — Northern coniferous forests in the breeding season and
tropical forests in winter. During brief sojourn on Reservation
seems to prefer large American elms.

Movements. — Because of my unfamiliarity with this species and
the brevity of its stay on the Reservation, I have overlooked it in
most years, but probably it is present regularly, on migration. On
May 14, 1954, many were present within a small area of an elm
grove south of the headquarters, and the song could be heard al-
most continually. Chasing was observed from time to time, and
there was a distinct tendency for each individual to keep to his own
part of a tree, excluding others. On May 20, 1954, further observa-
tions at the same place heightened this impression. An individual
observed for 15 minutes foraging rapidly through the crown of a
tree stayed witliin a space perhaps 20 feet across, tending to make
a regular circuit as it hopped, flitted and sidled through the leaf
canopy, moving only a few inches at a time, and frequently picking
off minute food items. It was covering approximately the same beat
as one of those observed six days earlier, and quite possibly was
the same.

Vermivora celata (Say)

Orange-crowned Warbler

Status. — Migrant and perhaps temporary resident in spring.
Probably crosses area on southward migration in autumn but no
records have been obtained at that season.

Habitat. — Boreal forests in the breeding season; chiefly Gulf
States in winter.

On the Reservation these warblers have been seen in woodland
edge — American elms with vine tangles of grape, Virginia creeper,

274 University of Kansas Publs., Mus. Nat. Hist.

and poison ivy. On April 19, 1954, these warblers were heard sing-
ing at several places and chasing was seen.

Vermivora ruficapilla (Wilson)
Nashville Warbler
Status. — Rare migrant.
Habitat. — Woodland and thickets.

Movements. — Richard W. Fredrickson saw several on the Reser-
vation on April 30, 1949, the only definite date of record.

Dendroica petechia (Linnaeus)

Yellow Warbler

Status. — Regular spring migrant and temporary resident; not
kno\vn to nest on area, which however, is within breeding range.
Habitat. — Deciduous forest.

Movements. — Heard and seen frequently each year in late April
and May. In 1954, for example, it was recorded from April 27 to
May 18. In May for five days in succession one was seen at the
same place in the elm grove south of the headquarters, and it was
suspected that this was the same individual.

Dendroica coronata (Linnaeus)

Myrtle Warbler

Status. — Regular migrant in spring and autumn.

Habitat. — Northern coniferous forests in breeding season, and
southern deciduous forests in winter.

Movements. — This is the most conspicuous of migrating warblers
on the area, in both spring and autumn.

Fall migration is concentrated in October, especially the first
week, and spring migration is chiefly in late April and early May.
In 1948 myrtle warblers were recorded on October 13 and 14. On
October 13, 1950, several newly arrived migrants were seen in
company with white-throated sparrows. At noon on October 11,
1951, many were noticed in elms in the headquarters area, the van-
guard of a migration wave. None had been seen earlier, but they
were present thenceforth through most of October. In 1953 the
first one was seen on October 4, but by the next day they were
present in abundance. In 1956 two were seen on October 2, and
in 1957 the first record was on October 4. Spring records are April

Home Ranges of Vertebrates 275

23, 1953; April 23 and 27, 1954; April 25 and 27, May 8, 22 and 25,
1956; and April 23, 24, and 26, 1957. The species has been recorded
on the area only in April, May and October.

Dendroica virens (Gmelin)

Black-throated Green Warbler

Status. — Rare migrant.

Habitat. — Chiefly northern coniferous forest in nesting season.
Movements. — Seen on several occasions in late spring. No defi-
nite date recorded.

Dendroica fusca (MiiUer)
Blackburnian Warbler

Status. — Rare migrant.

Habitat. — Boreal forest in nesting season; tropical forests in
winter.

Movements. — The only recorded occurrence is of a male seen on
May 15, 1954.

Dendroica striata (Forster)

Black-poll Warbler

Status. — Uncommon migrant and temporary resident in spring.

Habitat. — Northern coniferous forest in nesting season; tropics
in winter.

Movements. — Black-poll warblers were recorded on May 18 and
19, 1953, and May 14 and 15, 1954. The 1953 records probably
both pertain to tlie same individual, as both were in the same tree,
an American elm in a field near the edge of the woods. The warbler
was silent, and moved deliberately, usually running along small
branches rather than flitting. It was in the same part of the tree on
the two successive days and on the second day while it was under
observation, it spent more than an hour there.

Seiurus aurocapillus (Linnaeus)

Oven-bird
Status. — Rare migrant.
Habitat. — Deciduous forest.

Movements. — My only definite record is of one seen on April
30, 1951.

276 UNivxEsiTi' OF Kansas Publs., Mus. Nat. Hist.

Seiurus noveboracensis (Gmelin)
Northern Water Thrush
Status. — Rare migrant.
Habitat. — Nortliern forests, in swampy places.

Movements. — The only record for the Reservation is of one seen
April 30, 1949, by R. W. Fredrickson.

Seiurus motacilla (Vieillot)
Louisiana Water Thrush
Status. — Rare migrant.
Habitat. — Deciduous forest.

Movements. — The only one recorded by me was seen on May 5,
1952. It was called up by squeaking.

Oporornis formosus (Wilson)
Kentucky Warbler

Status. — Moderately common summer resident.

Habitat. — Occurs throughout most of woodland on Reservation,
but prefers mesic situations, with abundant leaf litter, and with
dense understory vegetation to provide concealment for nest.

Movements. — Earhest recorded occurrences of Kentucky warblers
in several different years were: April 21, 1952; April 26, 1954; April
28, 1955; May 3, 1956; and April 26, 1957. Arrival was timed when
woody vegetation had leafed out, and the individuals first recorded
were singing males, which were seen rather than heard. Establish-
ment of territories begins as soon as the males arrive; whether fe-
males reach the area simultaneously, or later, is not known.
Throughout their summer sojourn on the area, however, Kentucky
warblers are rarely seen. Information concerning their movements
has been obtained chiefly from the song, and therefore is based on
the males, which are persistent singers in May and June.

In 1952, 1954, 1955 and 1957, a pair nested in the headquarters
area. Conformation of the territory varied from year to year. In
1952 the territory was measured as 4.3 acres, and it centered on a
south-facing slope of thorny woodland with dense undergrowth.
A small, disjunct segment of this territory was along a ravine in
woodland edge 100 feet south across a meadow from the main por-
tion. Ill 1954 the entire territory, measured as 3.4 acres, was along
this ravine in the woodland edge, and it overlapped only the small

Home Ranges of Vertebrates

277

disjunct portion of the 1952 territory. The 1955 territory was
measured as 3.9 acres, most of which corresponded with the 1954
territory, but .5 acre was separate, on the north side of the meadow,
within the area of the 1952 territory.

Fig. 22. Territory of a pair of Kentucky warblers in
May, 1955, as revealed by singing stations of the male.
The territory was in woodland edge thicket. A small part
of the territory was separated from the remainder by the
narrow end of a field. Arrow indicates the place where
the warbler was seen to cross.

In May, 1956, a male that was singing persistently in dense woods
on a northwest-facing hillside, was followed and observed fre-
quently. A large elm, that was dead and bare, was the focal point
of his activity and he returned to it regularly after frequent short
forays in diflFerent directions. Unlike most other individuals that
were noted, this one did most of his singing from high perches in
the trees. His territory was only one acre.

Geothlypis trichas (Linnaeus)
Yellowthroat

Status. — Summer resident; many establish territories in early
summer and some may breed on the Reservation.

Habitat. — The swampland habitat most typical of this species
has been in some years totally lacking on the Reservation when the
pond was dry and in other years at best was limited to much less
than an acre in extent, in Typha patches at the edges of the pond.
Nevertheless, many yellowthroats arriving on migration in late
spring, settle on the area temporarily in atypical habitats. Some
have been found established in grassy fields, both in bottomland
and on hilltops. Others have been seen in woodland edge thickets,
and still others in woodland, where undergrowth of gooseberry and

278 University of Kansas Publs., Mus. Nat. Hist.

coralberry was plentiful. Just west of the Reservation boundary
a small alfalfa field provided the territory for one of these birds in
1949.

Movements. — Yellowthroats were seen and heard most often in
May upon their arrival from their northward migration. Earliest
recorded dates on the Reservation for several years are: May 14,
1950; May 17, 1952; May 9, 1953; April 26, 1954; May 4, 1955;
May 3, 1956; and May 8, 1957. The singing males established
themselves temporarily in a variety of situations, such as: fields of
brome grass; willow grove beside the pond; thickets at field border;
dry, south-facing slope with thorny woods and thick undergrowth;
alfalfa field. In June and early July they were seen or heard rela-
tively seldom, and so few were noted after mid-July that it seemed
they must have moved away from tlie area except for occasional
stragglers. In 1953 one was seen repeatedly at the pond and near
it in late July and early August. In 1955 one was seen at a fence
along the edge of a field of tall grass. Only one territory of a yel-
lowthroat was measured. It was 3.4 acres, based on only 5 mar-
ginal points. Perhaps the relatively large size of this territory was
due to some lack in the habitat.

Stewart (1953:105) recorded territories of ten monogamous
males which averaged 1.26 ± .12 acres (.8 to 1.8), while one poly-
gamous male occupied a territory of 3.4 acres. Brewer (1955:141)
recorded five territories in a swamp habitat in southern IlHnois,
with an average of .56 acre (.24 to 1.09).

Icteria virens (Linnaeus)
Long-tailed Chat
Stottis. — Regular summer resident but in variable numbers.
Habitat. — Limited to situations where there are dense thickets
such as blackberry, and rank weedy growth.

Movements. — Over a period of several years earliest dates of
record on the area for chats were: May 9, 1953; May 12, 1954; May
14, 1955; May 18, 1956; and May 10, 1957. Most records are for
May and June, but probably chats remain through the nesting
season. Because of their retiring habits and the dense cover of
their habitat, they might have easily escaped attention after the
singing and use of prominent perches associated with establishment
and defense of territory stopped.

In May and June, 1955, a singing male was heard and followed
several times in a hilltop field. A sapling thicket with crabapple,
American elm, osage orange, honey locust and other woody plants,

Home Ranges of Vertebrates 279

up to ten feet high, provided such dense cover that the chat was
rarely glimpsed. The thicket area was approximately .9 acre in
extent and was well separated from other suitable habitat. The
chat's territory seemed to be co-extensive with the thicket. In May
and June, 1956, a chat was followed as it moved about and sang,
along a gully bordered with thickets of wild plum and various other
shrubs and saplings. On both sides of this gully there were formerly
cultivated fields, but on one side woody vegetation, chiefly of elm
saplings, was invading the field. The chat used this thicket also,
though preferring the denser growth adjacent to the gully. The
territory encompassed by its movements while singing was 575 feet
long and 230 feet wide at the broadest point, covering approximately
two acres.

Brewer (loc. cit.) recorded four yellow-breasted chat terri-
tories that averaged .33 acre (.14 to .71).

Wilsonia pusilla (Wilson)

Wilson Warbler

Status. — Occasional migrant.

Habitat. — Boreal forests in breeding season; tropics in winter.
Movements. — The only definite record is of one seen in the elm
grove near the headquarters on May 19, 1953.

Wilsonia canadensis (Linnaeus)

Canada Warbler

Status. — Occasional migrant.

Habitat. — Boreal forests in breeding season, tropics in winter.

Movements. — The only definite record is of a male seen on May
15, 1953. It was working through an elm grove near the headquar-
ters in the morning, and shortly before dusk one (presumably the
same) was seen at the edge of the woods 100 feet farther south.
It was foraging by catching flying insects, darting out in pursuit at
intervals of a few seconds.

Setophaga ruticilla (Linnaeus)

American Redstart

Status. — Rare migrant.

Habitat. — Woodland.

Movements. — On May 3, 1952, a pair was seen and heard in a
thicket near the Reservation headquarters by Richard Fredrickson
and Mr. and Mrs. Bert Chewning. On May 15, 1953, I saw a male

280 Unin-ersity of Kansas Publs., Mus. Nat. Hist.

moving rapidly through the tops of large elms and it chased away
another small bird, perhaps of the same species. These are the only
definite records for the Reservation.

Passer doniesticus (Linnaeus)

English Sparrow

Status. — Frequent but irregular visitant; does not breed on the
Reservation.

Habitat. — This introduced species is characteristically associated
with man-made structures and altered habitats. In northeastern
Kansas, large flocks frequent nearly every farmyard. Roadside
hedges of osage orange and other thorny trees and shrubs that
border grain fields or are near to farm buildings often are the head-
quarters for such flocks. Where grain is grown or where poultry or
livestock are kept, the food supply encourages sparrows. Lone
sparrows, pairs or small groups often were attracted to the buildings
at the Reservation headquarters. Nesting sites, thorny thickets for
shelter, and grain for food ( that used as bait for trapping birds and
rodents ) were available, but the sparrows never became estabhshed
on this natural area.

Movements. — All but two of 25 recorded occurrences on the
Reservation were in late January, February, March, and April. The
visits to the area seemed to be motivated primarily by a search for
nesting sites. Individuals or pairs might visit the area daily for
several days or even several weeks. They might stay for only part
of the day. A barn and farmyard slightly less than a mile to the
west were the source of most of the individuals that came, or per-
haps of all. These pioneering sparrows were wary, and upon being
disturbed they would flush, circle over the area gaining altitude,
and, heading west, would pass out of sight in high, rapid flight.
Usually they would not return the same day. Those sparrows that
came alone were all males. On November 21, 1955, a pair was hov-
ering about the eaves of the garage, probably in search of a nest site.
On April 13, 1956, a pair was seen carrying nesting material to an
elm at the headquarters, but this nesting attempt was abortive.

In the spring of 1957 there were several dozen sparrows on the
Rockefeller Tract. They nested in the barn and several sheds, about
the house, and in nearby trees. By July most of these sparrows had
left. The buildings were no longer in use, and high weeds had
grown up in the adjacent areas that were formerly under cultiva-
tion, evidently creating conditions unfavorable for the sparrows.

Home Ranges of Vertebrates 281

Stumella magna ( Linnaeus )
Eastern Meadowlark

Status. — Common resident in general area but rarely present
on Reservation.

Habitat. — Meadowlarks in this area occur mainly in hay fields,
and prairie habitats where grass is fairly short. The habitat de-
ficiencies of the Reservation are not clearly apparent. As com-
pared with nearby occupied habitat on farmlands the habitat on
the Reservation differs in having a heavy accumulation of dead
vegetation on the ground, and in having denser grass. Neither the
extensive areas of brome grass fields, the original bluestem prairie
patch nor the re-established prairie have meadowlarks ordinarily.
In view of this bird's great abundance in the general area, its
sporadic appearance and brief sojourns on the Reservation are some-
what remarkable. In most instances the individuals heard or seen
on the area have not remained for more than a few minutes, but
nesting has been attempted on several occasions.

Movements. — In 1948, when the area was still grazed, the habitat
was unfavorable for meadowlarks. The first time one was seen on
the area was on June 1, 1949, in a hilltop field, where a crop of new
grass provided cover. None was known to have nested on the
Reservation that summer, but several were seen there in October
and November. In 1950, 1951 and 1952, meadowlarks appeared
briefly on the area in April and early May. Obviously these indi-
viduals were dispersing in search of territories for the nesting sea-
son. Some were in pairs. Singing and pursuit were common, but
in each instance the birds soon moved off the area. In 1953 on
March 20, a pair was found established in a hilltop field dominated
by brome grass. In the next two months these birds were seen
many times, and on several occasions were followed and caused
to flush several times in succession. The area covered was 2.8 acres.
In 1956 a male appeared on March 24, singing in the field near the
headquarters. Subsequently he appeared there regularly, some-
times accompanied by another. After arriving together, the two
would separate and sing from different trees, or from lower perches,
usually with no show of hostility. However, on April 12 three were
seen circling and dodging over the field in a high rapid pursuit. The
area which the male was seen to cover was 1.4 acres, and two trees
250 feet apart were his favorite singing stations. He did not ac-
quire a mate, and he disappeared from the area in mid-April.

282 Unr-ersity of Kansas Publs., Mus. Nat. Hist.

The failure of the meadowlarks that settled on the Reservation
almost every spring to consummate their nesting cycle there prob-
ably was due to the presence of grain fields nearby. As the grain
grew tall, these fields provided better nesting places than the fields
of brome grass on the Reservation, which were then deserted by
the meadowlarks.

In the spring of 1957 there were several pairs on the Rockefeller
Tract. Their activities were concentrated in the former pasture and
in the alfalfa field.

Stumella neglecta Audubon
Western Meadowlark

Status. — Uncommon transient.

Habitat. — Typically, short-grass prairie.

Movements.— On March 3, 1955, one was heard singing in a tall
grass prairie hilltop area of the Reservation near its northern edge.
Later in the day a flock of 19, probably of this species, were seen
flying northwest over the same field. On November 1, 1955, at
9:30 A. M., a small flock was perched on a wooded and brushy
hilltop near the west edge of the Reservation. They were resting,
and singing from time to time. One chased another, and both flew
out of sight to the west. Two days later a flock was seen foraging
at the edge of the county road % mile farther south, in an area of
big bluestem. After heavy snowstorms, in early January, 1956,
western meadowlarks were seen and heard frequently in a pasture
adjoining the Reservation on the west, and along the county road
just soutliwest of the Reservation section. One was singing in the
pasture on the Rockefeller Tract on October 9, 1957.

Xanthocephalus xanthocephalus (Bonaparte)
Yellow-headed Blackbird
Status. — Uncommon transient.
Habitat. — Marshland.

Movements.— On November 17, 1955, a flock of 50 or more was
seen flying south over the headquarters, strung out on a wide front,
as is characteristic in migrating blackbirds. On November 12, 1956,
a similar migrating flock, with a few starlings intermingled, was
seen at the same place.

Home Ranges of Vertebrates 283

Agelaius phoeniceus (Linnaeus)
Redwing

Status. — Several pairs usually are summer residents at the small
pond. Thousands migrate over the area without stopping in spring
and autumn, and many stop only for short periods.

Habitat. — The small pond and the marsh habitat about its edges
provide the only breeding habitat on the Reservation.

Movements. — Redwings were seen on the Reservation chiefly as
migrants in spring and autumn. Each spring the pond attracted
them, but because of the uncertain water supply, they sometimes
did not stay to nest. In the period of years 1948 through 1952, and
in 1957, with moderate to heavy precipitation, the pond retained
water at least through the early summer, and redwings nested in
the marsh at the edge of the water. In 1953 through 1956, with
more meager precipitation nesting was unsuccessful there. Singing
was first heard at the pond on March 20, 1951; February 29, 1952;
March 13, 1953; April 6, 1954; and March 15, 1955. Mass migra-
tions were noted each spring, sometimes beginning in early Febru-
ary, and sometimes lasting into late April. Autumn migration was
noted chiefly in October and November, In 1954 spring migration
reached its peak on approximately March 12, although it had been
in progress since early February. On March 12, weather was
unseasonably mild, with a high wind from the southwest, and red-
wings were passing over by the thousands. They were in sight or
hearing almost continually. Some were flying alone, others in twos
or threes or larger groups up to those numbering many hundreds.
Such large flocks strung out on a broad front, sometimes as much
as half a mile across.

In contrast to the loose formation of the flocks that were actually
migrating was the compactness of flocks that lingered in the vi-
cinity for periods of days, or even weeks, in spring. These flocks
came to the Reservation to roost in late afternoon or early evening,
usually not appearing until near sunset, and otherwise their daily
routine of activity was unknown. Presumably they foraged and
visited prospective nesting areas within a radius of a few miles.
Flying swiftly, in close formation, a flock of up to 200 birds would
maneuver over the area and then settle in the top of one of the
tallest trees available, where they would chatter and sing in chorus.

284 Unu-ersity of Kansas Publs., Mus. Nat. Hist.

In the hour or more that sometimes elapsed between its appearance
and its final settling to roost, the flock would normally shift several
times from one high tree to another, sometimes splitting into two
or more units, and sometimes joined by new contingents. When
finally it had become nearly dark, the redwings would fly into tall
grass to roost, each bird choosing its own moment for the descent,
and finding its own spot to settle. Usually only part of a flock
would settle in any one small area, and the remainder might shift
uneasily to another tree before finally going to roost. That such
roosting aggregations consisted largely of the same individuals over
periods of days or longer was strongly indicated by return of the
flock to the same trees in more or less the same sequence on suc-
cessive evenings. In 1954 the roosting aggregations were seen
daily over the period March 8 through April 5. Within this period
the largest aggregations were seen in late March and April.

In spring male redwings always appeared at the pond and sang,
well before any females were in evidence. In some seasons, when
habitat conditions were unfavorable, females probably did not come
there at all. The entire area about the pond that was used by red-
wings, was less than two acres in extent; the area actually available
for nesting was variable, but usually was considerably smaller.
The male redwings that came to the pond in the early part of the
nesting season spent only a part of each day there. Often two or
more males travelling in high rapid flight, would come to the pond
together, and perch separately in tall trees to sing and display. In
departing also, they usually kept together, in high rapid flight,
probably headed for distant roosting areas.

Nero (1956:130) found the average size of territory for 17 well
established males to be about 1/12 of an acre. He found that size
of territory varied inversely with the population density. Males
are polygamous, and one's territory may have several mutually
hostile females living within it. These females may be at different
stages of the breeding cycle.

Icterus spurius (Linnaeus)

Orchard Oriole
Status. — Rare visitor.

Habitat. — Parkland, and trees in relatively open situations.

Movements. — On May 16, 1951, a group of at least eight was in
a Kentucky coffee-tree (Gymnocladus dioica) at the Reservation
headquarters. Only two were males in breeding plumage. All

Home Ranges of Vertebrates 285

were feeding avidly on the flower buds of the tree. In 1958 orchard
orioles were seen or heard on the area almost daily in late May and
June, but in the intervening seven years none was recorded.

Icterus galbula (Linnaeus)

Baltimore Oriole

Status. — Moderately common summer resident in general ai'ea
but not known to breed on Reservation.

Habitat. — Groves of large deciduous trees.

Movements. — Earliest appearances over several years were:
April 28, 1951; May 16, 1952; May 6, 1953 (one seen three days
earlier at Ottawa, 30 miles south); May 4, 1954; April 27, 1955;
May 12, 1956; and May 1, 1957. In each instance the earliest record
was of a male, and males may have arrived a httle sooner than fe-
males. In no instance, so far as known, did Baltimore orioles re-
main on the area through the nesting season. Records were chiefly
from May (19 different dates) and from August (15 different
dates), with relatively few between (June 20 and July 16, 1954;
July 31, 1955). The species must have found conditions unsuitable
for nesting, for those individuals that had stopped on the area soon
shifted elsewhere. The reappearance in August probably coincided
with the end of the nesting cycle, at least for those individuals in-
volved, which shifted away from their nesting territories as con-
ditions changed there and became less favorable.

In 1953 the August visitation involved a pair and one or more
fledglings that were still dependent. These orioles appeared
abruptly on August 21; thereafter they were seen or heard daily
tlirough the remainder of the month, and from time to time in early
September (last record on the fourteenth). The records of tliis
group encompassed an area of 2.1 acres. In 1955, after the first
appearance on April 27, a male which probably was the same, was
present continuously until May 18. He was singing almost con-
tinually during the first few days, but on May 4 he abruptly became
much less vocal and remained so during the latter part of his stay.
The area encompassed by his movements was approximately 10.6
acres, in the shape of an elongate ellipse, with its long axis 1200
feet from end to end, extending along woodland edge.

In 1957, on the Rockefeller Tract, a cottonwood near the house
was much used by an oriole and nesting may have been attempted
there.

286 Untversity of Kansas Publs., Mus. Nat. Hist.

Euphagus carolinus (Miiller)
Rusty Blackbird

Status. — Regular transient in migration in spring and autumn;
occasional in winter.

Habitat. — Swampy areas of boreal forest in nesting season; in
winter range, which includes region of this study, requires such
areas as open fields or closely grazed pastures, because of habit of
foraging on ground.

Movements. — On October 21, 1948, several were seen and on the
morning of October 25 a flock of several thousand settled in the
grove at the edge of a hilltop field. These blackbirds may have
stopped to forage in the heavily grazed pastures which at that time
still had livestock in them. Soon afterward the livestock were
removed, and the blackbirds were never again seen in such numbers.

The two December records were of lone birds; on December 16,
1953, a male was seen foraging at the edge of a frozen pond, and
on December 2, 1955, a juvenile was foraging with a flock of star-
lings and robins beside the headquarters building. This was in
the yard where grass and weeds had been cut the preceding sum-
mer. On January 28, 1957, when the ground was snow-covered,
a flock of 35 was seen at the northwest corner of the Reservation,
and foraging in an adjacent field of milo stubble. On October 19,
1955, and November 12, 1954, southward migrating flocks passed
over. On November 3, 1948, and November 20, 1951, flocks were
seen resting in the headquarters area. In spring, March 11, 14, 22,
23, and 24, 1955; March 14 and 16, 1956; April 9, 1954; and April 3
and 5, 1955, rusty blackbirds were seen in compact flocks on the
Reservation. These flocks invariably came to the area in late
afternoon, attracted by the dense ground vegetation providing
favorable cover for roosting. A compact flock would often perch
for 10 minutes or more, chattering and singing, and would shift
from tree to tree several times before finally going to roost.

Quiscalus quiscula (Linnaeus)
Bronzed Crackle

Status. — Regular migrant in spring and autumn, but usually as
transient flying over area without stopping.

Habitat. — Short ground vegetation, permitting easy walking and
an unobstructed view, perhaps is the one requirement that is con-
sistently lacking on the Reservation, making it unattractive to the

Home Ranges of Vertebrates 287

grackle, which is a common breeding species locally on farmlands
and suburban areas.

Movements.— The only definite record of the grackle stopping
on the Reservation was obtained on April 4, 1951, when a flock of
more than 2000 were foraging in a hilltop field of brome grass. In
flushing they split into two groups that flew west across the area.
Within the next few hours a large flock was seen several times
maneuvering over different parts of the Reservation. These
grackles flew in a compact mass, and as they flushed or turned, their
wingbeats produced a roaring sound. While on the ground or
perched they kept up a chattering that was audible half a mile away.

In contrast, tliose that were passing over on migration were flying
parallel, strung out on a broad front. Often they were flying with
redwings. On October 31, 1952, at 9 A. M., a large flock passed
over headed south. On February 14 and 16, 1954, several were
seen flying high headed northeast. On March 2, 1954, a flock of
several hundred, on a front more than 100 yards wide, passed over
headed north.

Molothrus ater (Boddaert)

Cowbird

Status. — Common summer resident.

Habitat. — Groves along edges of bottomland fields are preferred
on Reservation in spring and summer, but probably every part of
area is used to some extent.

Movements. — Ordinarily cowbirds are present on the area only
during the nesting season. An unusually late record was November
7, 1949. A group of 30 or more was in a thicket at the edge of a
field, and many of them were feeding on the fruits of dogwood
{Cornus drummondii). These were probably transients from far-
ther north, that were passing on migration. Flocks of cowbirds
winter in the region of the Reservation, but they stay about farm-
yards and pastures, where there are cattle or other livestock, and
none has been seen on the Reservation. These wintering flocks
presumably are from farther north in the species' range.

In early spring each year, newly-arrived singing males appeared
abruptly on the Reservation. Within a few days their numbers
were much augmented by new arrivals and females began to
appear. As compared with other kinds of breeding birds on the
area, cowbirds were notable for the wide variation in time of ap-

8—3117

288 Unrtrsity of Kansas Publs., Mus. Nat. Hist.

pearance from year to year. Over a seven-year period earliest dates
of appearance were: April 14, 1951; March 24, 1952; March 13,
1953; March 28, 1954; March 31, 1955; March 22, 1956; and March
13, 1957. The earliest appearance, March 13, 1953, was somewhat
different from those of other years, as three males were seen perched
close together feeding on leaf buds, and not singing. On the fol-
lowuig day others were seen flying over. On March 16 two were
seen on tlie area, and from March 30 onward they were seen fre-
quently.

Cowbirds were most conspicuous on the area in April, May and
June. They were usually seen in small groups of two or three fe-
males and a somewhat larger number of males. These groups
moved about from tree to tree, usually over an area of several acres,
the males singing, displaying, and jostling each otlier for position.
From time to time one bird or several might separate from the
sexual aggregation and leave in strong direct fhght. For the groups
seen near the Reservation headquarters, the destination was obvi-
ously a pasture on the farm west of the Reservation, a distance of .5
mile to one mile. Only rarely were cowbirds ever seen on the ground
on the Reservation, and the rank vegetation seemed to create con-
ditions unfavorable for their foraging. Occasionally they were seen
feeding in trees, but probably they obtained most of their food in
pastureland beyond the limits of the Reservation, and came there
only for breeding. As the season advanced, throughout most of
the summer, the aggregations were seen much less frequently;
cowbirds seen were mostly females, which were silent and furtive
in behavior, lurking in thickets and crowns of trees. Although a
high proportion of all birds' nests on the Reservation are parasitized
by cowbirds, fledgling cowbirds have rarely been noticed on the
area.

Piranga olivacea (Gmclin)

Scarlet Tanager

Status. — Recorded on Reservation only as spring migrant.

Habitat. — Deciduous forest.

Movements. — My only record of this species was obtained on
June 13, 1951, when a pair was seen flying across a field to a large
elm near the Reservation headquarters, a few minutes before sunset.

Home Ranges of Vertebrates 289

Piranga rubra (Linnaeus)
Summer Tanager

Status. — Moderately common from late April to early September;
several nesting pairs each year.

Habitat. — Deciduous forest and edge.

In most instances pairs were observed in dense woods of pre-
dominantly oak-hickory type, but they also foraged frequently in
more open groves of elm and walnut adjacent to the denser woods.
The males tended to choose as singing stations large trees somewhat
apart from the main woodland. Several large honey locusts in
fields near woodland edges were the most used singing stations of
different males.

Movements. — Earliest recorded dates for several years were:
April 27, 1951; April 26, 1952; April 28, 1953; April 24, 1954; April
30, 1955; May 4, 1956; and April 30, 1957. In every instance the first
individual noted was a male, and possibly the males arrive a little
earlier than the females. Nest-building was observed in 1953
(Fitch and Fitch, 1955:48). Only the female carried material, but
the male stayed nearby. Some of the stems used were collected
on the ground beneath the nest, but nest-building for the first few
days proceeded at a leisurely pace. The female was first observed
at the nest site on May 11. By May 18 she was observed carrying
stems for the nest foundation. Building continued at an accelerated
rate through May 24. Much of the nesting material was gathered
along a ravine at the edge of woodland 200 feet from the nest.

When the young tanagers in the nest were just one week old, they
climbed out along the limb nearby, and one by one fell into tall
grass. At this stage they had only begun to feather out, and were
almost helpless. After spending three days in the grass near the
nest, they were sufiBciently developed to fly into nearby trees. The
whole family then soon shifted to the wooded ravine 200 feet or
more from the nest tree, and were seen there frequently in the
following three weeks.

The territory within which most of the activity of the pair of tan-
agers was hmited, was measured as 4.3 acres, in a broadly elliptical
area, which was mostly in an open grove of large American elms
at the edge of woodland. From time to time, however, one or both

290 University of Kansas Publs., Mus. Nat. Hist.

birds were observed well beyond the limits of this usual area. The
seven most outlying points encompassed an area totalling 20.8 acres.
Lack of territorial pressure probably encouraged these birds to
wander frequently beyond their home area. Each year at least
six pairs of summer tanagers nested on the Reservation, but their
territories were so well spaced that no more than one at a time
could be heard singing. Encounters between members of different
pairs were never observed, and must have occurred relatively sel-
dom. Each year singing ceased abruptly in the second week of
August. My latest records are on September 27 ( 1953 and 1957 ) .

Richmondena cardinalis (Linnaeus)

Cardinal

Status. — Common resident, using every part of Reservation ex-
cept extensive open fields; populations fluctuate from year to year
and from season to season; greatest numbers present in winter;
migrational shift of populations evidently occurs from summer to
winter.

Habitat. — Dense thickets in edge situations adjacent to woodland
and fields are preferred. Thick thorny shrubs such as osage orange,
crab apple and honey locust are favorite nesting sites. Dense
woodland edge, and thickets along roads and creeks provide shelter
and resting places, but cardinals forage in a wide variety of situa-
tions, and utilize many different food sources.

Movements. — In late autumn each year cardinals suddenly be-
came more abundant and conspicuous. Unusual concentrations,
probably made up mostly of new arrivals, were noted on: Novem-
ber 8, 1948; November 14, 1951; October 29, 1952; November 30,
1953; and October 17, 1955. Even larger concentrations were some-
times noted in late winter at times when weather was unusually
severe, and ground was snow-covered. The extent of population
shifts in response to extremes of weather was not determined.

Cardinals sing throughout the year, but singing is at low ebb in
November, December, and January. It occurs only on clear, still
mornings when weather is unusually mild. Singing tends to in-
crease throughout February, although it is temporarily stopped by
wind, low temperature, snow, or rain. By early March males are
beginning to establish territories.

In winter individuals are highly tolerant, and several may feed
or perch in proximity. However, aggregations are drawn together

Home Ranges of Vertebrates 291

by food supply, shelter, or other attractions, rather than by socia-
bility. Usually each individual is more closely associated with
other small passerines, such as Harris sparrows, tree sparrows, song
sparrows and juncos than with members of its own species.

On November 30, 1953, when a group of cardinals had gathered
in a spot exposed to the early morning sunshine, there was brief
singing, and a male chased a female persistently. Several other
males, excited by the pursuit, were trailing along, chiefly to harass
the first male, it seemed, rather than to catch the female.

For the cardinals live-trapped in winter, distances between suc-
cessive captures averaged 413 feet for 49 marked males, and 565 feet
for 31 records of females. Home ranges of 12.2 acres for males and
22.9 acres for females are indicated. Not included in this compu-
tation are three exceptionally long movements (one of a male and
two of females ) of nearly half a mile each. These are thought to
represent shifts in range. For those intervals including one or
more summers between successive captures, distances averaged no
more than for intervals within the same winter.

Approximately the first week of March the males, having estab-
lished territorial singing stations, become increasingly intolerant.
A typical encounter was observed on the morning of March 8, 1957,
at the edge of woodland. Two males were hopping about briskly,
with crests erect and tails jerking, within a few feet of each other,
occasionally giving metallic chirps. When one would fly, the other
would follow close behind. There was no actual contact. Usually
the chase led back and forth along the edge of the woods in an old
diversion ditch, but occasionally the birds crossed a narrow field
to edge of woodland on the opposite side, 150 feet or more away.

For a period of weeks stress increases as males defend their newly
established territories against the encroachment of neighbors, and
against the wintering cardinals still wandering over home ranges
in the general area. However, territoriality develops gradually.
Singing and hostile behavior is most noticeable in early morning.
Later in the day the males may relax their vigilance and mingle with
others with little indication of intolerance. In periods of stormy
weather, territorial behavior may be suppressed entirely. Forma-
tion of pairs occurs only after the males are thoroughly ensconced
on their territories. Females still ranging freely over their large
winter ranges, are followed and courted by the males whose terri-
tories they cross. In competition for mates actual fights are fre-
quent, along with intimidating displays and pursuits. On March 9,

292 University of Kansas Publs., Mus. Nat. Hist.

1954, two males that were chasing the same female stopped to fight
violently. They fluttered near the ground, sparring, their wings
ratthng together. In the period March 16 to 19, 1952, two males
having newly established territories along the edge of the woods
near the Reservation headquarters were observed frequently. Their
territorial centers seemed to be about 150 feet apart. Each often
strayed into the other's territory and was chased back, the pursuer
usually stopping abruptly near the same place, where a mutually
accepted boundary seemed to be situated. After such a pursuit
the males would stop a few yards apart, each on his own side of
the boundary. They would watch each other, flicking their tails
and uttering clicking notes, then would fly to high perches and
sing. A female seemed to be staying in the general area, ranging
over both territories, and she was the object of frequent pursuits,
sometimes by both males simultaneously. She would fly rapidly
from tree to tree and bush to bush, along the ravine in the edge
of the woods avoiding the males with frequent quick turns. In
these pursuits the males became so preoccupied as to forget defense
of their territories, but occasionally one would dart at another.
Once a second female, which had been watching with excited
chirping, interposed herself, so that all four birds were strung out
in a line, the two males bringing up the rear.

On April 16, 1952, an encounter was observed between four
cardinals which seemed to consist of a male chasing a prospective
mate, and an established pair whose territory was violated in the
course of the pursuit. The paired female was not chased but she
followed the pursuit with hostile behavior directed largely toward
the unattached female. The paired male followed the otlier male
and from time to time intercepted him and engaged him in short
fights. The wrangling continued for approximately half an hour
that the birds were watched. On April 22, 1955, when persistent
scolding attracted my attention, I observed a male trying to head
off a female and drive her back into the area presumed to be his
territory. A second male was interfering. Occasionally the two
males closed in and struck at each other. Much of the month of
March is spent in such maneuvering.

By early April most cardinals are paired. An unusual feature
of territoriality in this species is the singing of the female. On
occasion, persistently singing birds assumed to have been males
turned out to be females. Paired females have been heard to give
a burst of song in reply to each burst of the male, as he sang from
a nearby territorial perch.

Home Ranges of Vertebrates 293

No territories of cardinals were actually measured. The some-
what secretive habits of the birds in the season of nesting made it
difficult to follow their movements. Nine intervals between occu-
pied nests ranged from 220 feet to 560 feet and averaged 391. That
this distance was somewhat representative of the diameter of a
territory was borne out by the distances measured between adja-
cent singing males in 169 instances. This distance between the
singing males averaged 416 feet indicating an average territory of
3.1 acres if it be assumed that tlie territories are circular and have
common boundaries.

Brewer (1955:141) recorded an average area of .37 acre (.31
to .45) covered by five cardinals in a swamp thicket habitat in
southern Illinois. The small size of the "minimum home range"
obtained by Brewer, in this and other instances probably result from
inadequately small series of records.

Pheucticus ludovicianus (Linnaeus)

Rose-breasted Grosbeak

Status. — Regular migrant or transient in spring, some stopping
on the Reservation for periods of days or weeks, but not knov^oi to
nest.

Habitat. — Deciduous forest and parkland situations are occu-
pied in the breeding season. On the Reservation it has been seen
and heard chiefly on a northwest slope in mesic mixed forest, with
ash, hickory, elm, chestnut oak and black oak, and in a willow grove
at the pond.

Movements. — Seen or heard on the following dates: 1951, April
30; 1952, April 29 and 30, May 17 and 22; 1953, May 19; 1954,
May 15; 1955, May 8, 9, 13, 14 and 16; 1956, April 26, 28, 29, May 2,
4, 5, 7 and 26. It is noteworthy that each year all had disappeared
by June 1, although they seemed to be settling down and estab-
lishing territories in May. Often those seen were feeding and mov-
ing about in pairs. On April 30, 1951, two males were eating leaf
buds of ash in the same tree. Similarly on May 7, 1956, two males
were feeding together in the same tree. On other occasions, pur-
suits were seen, but on May 5, 1956, three pairs were feeding to-
gether in a willow grove, all within a 50-foot radius, with no sign
of hostility. Abundant larvae of chrysomelid beetles on the willow
leaves made this an attractive feeding place. Probably all those
seen on the Reservation were recent arrivals, which had not yet
settled down permanently or established territory.

294 University of Kansas Publs., Mus. Nat. Hist.

Guiraca caerulea (Linnaeus)

Blue Grosbeak

Status. — Summer resident on areas immediately adjoining Reser-
vation, but rarely crosses boundary.

Habitat. — Brush.

A wooded slope of north and west exposure adjoining the west
edge of the Reservation, was bulldozed in late winter of 1949. By
1952 the area had grown up into a thicket, with numerous stump
sprouts of hickory, oak and elm, high weeds, and with the par-
tially burned debris of uprooted trees piled at the bottom of the
slope. Grosbeaks were first definitely recorded in 1952, and were
present regularly thereafter.

Movements. — Two pairs were present on the brushy strip west
of the Reservation in 1955 and 1956. The male of one pair was
frequently seen along a 250-foot stretch of road, perched on tele-
phone wires and occasionally his mate was with him. The territory
was at least 200 feet wide, and thus included a minimum area of
IM acres. Farther south along the road a second pair was often
seen, but a stretch of some 350 feet intervened where none was re-
corded.

Passerina cyanea (Linnaeus)

Indigo Bunting

Status. — Common summer resident, late April through Septem-
ber.

Habitat. — Low thickets in or near fields with grass, weeds or
cultivated crops provide the preferred habitat of the indigo bunt-
ing. From 1948 through 1958 conditions have steadily improved
for the bunting, because thickets have encroached into formerly
pastured areas. These thickets, of crab apple, honey locust, osage
orange, elm, and a few other trees, have created abundant shelter
and nesting sites for the buntings. The most dense population of
buntings noted was on a north slope just west of tlie Reservation,
wooded until 1948, when it was cleared by bulldozing. The piles
of broken tree trunks, limbs and roots were partly burned. In the
next few years saplings of oaks, hickories and other trees sprang
up in large numbers. Also, there were many vines and thick weeds,
so that by June, 1955, the area was an almost impenetrable tangle.

Movements. — Earliest recorded occurrences of the indigo bunting
for the period 1952 through 1957 on the Reservation were in early
May each year, the third, fourteenth, ninth, second, fourth and

Home Ranges of Vertebrates 295

fourth. Singing was chiefly in May and June, but continued into
early August. My latest record is for September 15, 1953.

Despite the abundance of indigo buntings on the area, it proved
difficult to trace out the territory of any one individual because of
their shyness and frequent flight, and the difficulty of recognizing
individuals. One territory, measured on the basis of 15 recorded
movements within it, was 2.7 acres. In 32 instances the distances
between adjacent singing males were recorded, and they varied
from 1300 feet to 110 feet, averaging 504 feet. A territory of this
latter diameter would cover approximately 4.6 acres. Brewer
(1955:141) recorded an average area of only .26 acre (.15 to .52)
for five indigo buntings in swamp thicket habitat in southern Illinois.

Passerina ciris (Linnaeus)

Painted Bunting

Status. — Rare, probably occurs on Reservation only as summer
transient.

Habitat. — Brush and woodland edges.

Movements. — The only recorded occurrence of this species was
obtained on May 14, 1950, by Mr. and Mrs. Bert Chewning. The
bunting was heard ( but not seen ) singing at the edge of a brushy,
south-facing slope where the woodland adjoined a meadow.

' Spiza americana (Gmelin)

Dickcissel

Status. — The dickcissel is one of the most abundant of breeding
birds in the general area. Since 1948, when the Reservation con-
sisted of woodland, closely grazed pasture and cornfields, plant
succession has proceeded. Certain areas have grown up to tall
grass and weeds and have become favorable habitat for dickcissels.

Habitat. — Grassland.

Under modern conditions in northeastern Kansas most dickcis-
sels nest in fields of wheat, barley, oats, or alfalfa. Singing stations
are provided mainly by fence posts and by telephone wires at the
fields' margins, and the singing males tend to be rather regularly
spaced. On the Reservation dickcissels have become abundant in an
old field that was sown to seeds of native grasses in 1949 and has
taken on the aspect of a prairie since then, with tall grass clumps
(big bluestem, little bluestem, switch grass, Indian grass, side-oats
grama) in varying density interspersed with goldenrod and other

9—3117

296 Uxi\'ERsi'ri' of Kansas Publs., Mus. Nat. Hist.

weeds in patches, with numerous saphngs of ehn, locust and other
trees.

It is noteworthy that dickcissels are more numerous on parts of
this field where a mixture of several kinds of grass had been planted
than they were on parts where the several kinds were growing in
separate strips.

In 1953 a single pair of dickcissels nested in the field of brome
grass and bluegrass with numerous small trees, near the Reserva-
tion headquarters. In the years 1953 through 1955 another pair
had a territory which included part of a field of bluegrass and
brome, and part of a formerly cultivated field with stunted giant
ragweed, and large patches of foxtail grass. In 1954 and 1955 sev-
eral pairs were located in the southwestern part of the Reservation
in a field of brome and bluegrass with thickets of saplings, adjoin-
ing an oat field.

Movements. — Arrival dates for tlie years 1951 through 1957 were
April 30, May 3, May 4, April 29, May 1, May 2, and May 6. Within
a few days after the first arrival males were abundant and conspicu-
ous, spending much of their time singing from territorial perches.
No dickcissels have been seen on the area after the end of July. If
they remain on the area at all, they must become extremely silent
and secretive.

Because of their tameness, and the open nature of their habitat,
dickcissels could be studied, with respect to territory, more readily
than any other species of bird present on the Reservation, While
singing, they would often allow approach to within 25 feet, and
when flushed, tliey would usually show no alarm but might move
only a few yards, to a nearby perch. By following such an indi-
vidual from perch to perch for a short time, it was easy to determine
the approximate extent of its territory. Nine territories investigated
in this way ranged from 1.4 to .5 acres and averaged .9 acre. In
14 instances the distances between adjacent singing males, where
the habitat appeared to be continuous, varied from 450 to 60 feet
and averaged 296. Assuming that this distance represents the
diameter of an average territory, an area of 1.6 acres would be
represented. Presumably the discrepancy between this figure and
the .9 acre of territories actually measured results in part from the
fact that some territories were incompletely recorded, and in part
from the fact that there are some unoccupied spaces between ter-
ritories. Therefore, an intermediate figure of approximately IJ*
acres may be most typical.

Home Ranges of Vertebrates

297

The largest territory measured, one of 1.4 acres, was in the
meadow of brome grass near the headquarters, a habitat different
from the tall-grass prairie where most of the dickcissels were ob-
served. In the absence of competition this pair tended to cover a
larger area than they would have otherwise. Records are far more
complete for this pair, than for any of the other pairs observed. The
territory of 1.4 acres was plotted from 30 locations, many of which
were used many times by the birds during the course of observa-
tions. Occasionally the male was noted to sing from perches well
beyond the limits of the area usually occupied, in each direction.
The total area encompassed by these outlying points totalled 5.6
acres, but obviously most of this area fell outside the regular ter-
ritory (Fig. 23).

Fig. 23. Territory of a pair of dickcissels in June and July, 1953. So

far as known the female kept to the relatively small central area, but the

singing male, in the absence of competitors, often wandered to outlying

stations. Lines show observed movements.

The pair first appeared on this territory on June 10, 1953, prob-
ably after an unsuccessful nesting attempt elsewhere, because more
than a month had elapsed since arrival of dickcissels and establish-
ment of territories in the general area. Each time the male sang,
the female answered with a low buzzing sound. The female was
seen carrying nesting material on the morning of July 9, and in the

298 University of Kansas Publs., Mus. Nat. Hist.

afternoon of the same day the nest was found completed with one
egg in it. By July 14 the clutch of four eggs had been completed,
and hatching occurred on July 23-24. The nest was robbed on July
31, and on the following day the adults had left the territory.

On a few occasions, singing males that were followed too per-
sistently and caused to take alarm, moved somewhat beyond the
limits of their territories, with accompanying change in behavior.
Ceasing to sing, they would skulk through low vegetation, instead
of perching on the highest and most exposed perches available, as is
usual with singing males.

Elsewhere in the general region, telephone wires at the edges
of grain fields provide the most favored territorial perches. In the
prairie area where most of the pairs were studied on the Reserva-
tion, there were no trees other than saplings. A telephone line
along the north edge of the Reservation, on the edge of the prairie
tract, therefore provided the best perches for singing males and it
was used regularly by all those having territories along the north
edge of the field. It was used also by several others whose territories
did not adjoin the edge, unless a narrow corridor of travel to and
from the wire be considered part of the territory.

In this area where several pairs lived in adjacent territories, pur-
suits were frequent, but usually they were brief and did not involve
actual fighting. So far as observed, these territorial pursuits in-
volved only the male.

Carpodacus purpureus (Gmelin)
Purple Finch
Status. — Occasional transient in winter.

Habitat. — On the Reservation purple finches were most often seen
near the pond, which was the focal point of their activity. Stands
of sunflower and other weeds attracted them, the seeds supplying
an abundant food source. At other times, in fall, they were seen
foraging in treetops, probably for dried fruits and berries.

Movements. — Purple finches recorded on the Reservation were
usually travelling in small groups in high, rapid flight and they
were among the most vagile of all birds in tlie area. Also, they
were more wary tlian any except raptors and water birds, and gen-
erally would not permit approach closer than 200 feet. Twenty-
two specific records were distributed as follows: October — 8, No-
vember— 6, December — 3, January — 1, February — none, March — 2,

Home Ranges of Vertebrates 299

April— 2. Earliest fall records were October 8, 1956; October 15,
1955; October 24, 1954; and October 15, 1957. Latest spring rec-
ords were April 2, 1955, and April 12, 1956.

Spinus pinus (Wilson)

Pine Siskin

Status. — Occasional transient.

Habitat. — In breeding season, northern forests, usually associated
with coniferous trees.

In its more southern winter range the species may prefer conifers
also. No conifers are present on the Reservation, and in any case
the area is, in general, unattractive to siskins, which have been ob-
served there on only a few occasions.

Movements. — In the first week of November, 1954, siskins were
seen many times at the pond where they were attracted by sun-
flower patches, and were feeding on the seeds. Again in late
October and November, 1955, siskins were abundant at this same
place and at a fallow field on the east side of the Reservation, where
there were patches of sunflower.

Spinus tristis (Linnaeus)
Eastern Goldfinch

Status. — Common resident, using every part of Reservation;
moves about freely, and numbers vary markedly according to season
and food supply.

Habitat. — Activities are concentrated about weedy fields. Seeds
of sunflower seem to be the one most important food source, and
are used throughout the fall, winter, and early spring. Seeds of
various other composites are used to a lesser extent. Since 1948
when formerly cultivated fields produced their first crop of pioneer
annual weedy vegetation, the quantity of sunflower has steadily
decreased, because this plant is most characteristic of the early
stages of secondary succession. As a result, the area is used less
by goldfinches. The dwindling sunflower patches continue to be
focal points of their activity. In late March and April, when seeds
of elm are maturing and are still on the trees, they are preferred
food, and large restless flocks of goldfinches are much in evidence,
moving about in the treetops or maneuvering over them.

Movements. — Goldfinches have not been recorded on the area
in the months of February or June. While they doubtless visit

300 University of Kansas Publs., Mus. Nat. Hist.

the area occasionally at all seasons, they are relatively scarce in
winter and summer, and are much more frequent in September
and October, and again in late March and early April. Those pres-
ent in fall are seen singly or in pairs, while those present in spring
are in large roving flocks attracted to the area by the abundant
supply of elm seeds for food. Occasionally at this time of year
they may be among the most abundant of birds present. Nesting
activity has been noted chiefly in August and September. In Au-
gust, 1954, several were noted in different places circling and singing
at heights between 100 and 200 feet over hilltop grassland areas.
In September, 1953, a pair nested in the headquarters area. Fre-
quently they flew over the territory, in circles of perhaps 200 yards
diameter, keeping a httle above the level of the treetops. To for-
age, these goldfinches almost always left the Reservation, and on
many occasions v.'ere seen to make flights of at least half a mile
west or north to pastures and cultivated fields. Sometimes both
members of the pair travelled together on these foraging expedi-
tions, but more often they travelled alone. On September 14, a
family group of five was seen maneuvering over the territory, and
on September 15 a group of seven was noted flying toward the same
area. In mid-October, 1954, goldfinches seen foraging in sunflower
patches were in pairs.

Stokes (1950:114) studying the goldfinch's nesting habits in a
dry marsh near Madison, Wisconsin, found territories were only 95
feet in diameter in an area of favorable habitat. He found that the
territory, including the nest site, did not necessarily provide food,
water, or nesting material. Territorial defense was strongest at
the beginning of the nesting cycle, and was much reduced by the
time the young had hatched. Nickell (1951:450) found three nests
within a 50-yard radius in a dry swamp habitat, and concluded that
the species is highly tolerant. His findings contrasted with those of
Drum ( 1939:71-72) in a different type of habitat. The latter author
found that territories were vigorously defended by the male, and
sometimes also by the female, and that feeding areas were some-
times a mile or more from the nest.

Pipilo erythrophthalmus (Linnaeus)

Red-eyed Towhee

Status. — The towhee is moderately common throughout most of
the year on the Reservation. However, the breeding population
present in summer is of a subspecies different from the one present
in winter. It is not definitely known whether there is seasonal over-

Home Ranges of Vertebrates 301

lapping between the summer and winter populations. Representa-
tives of both populations may be present for a short time in April.
Towhees have not been definitely recorded in September; there
may be a period in autumn when neither population is represented
on the area.

Habitat. — Blackberry thickets at or near the edge of woodland
seem to provide the essential habitat feature for the wintering popu-
lation. The birds seem to do much of their foraging in the ground
litter beneath these thickets, and in their daily routine they shift
from one thicket to another, spending relatively little time in other
situations. They are seen most often away from thickets at times
when the ground is snow covered and food is scarce. In summer
the same blackberry thickets are used by the breeding population,
but these towhees also use a variety of other situations. Thickets
of wild plum, crab apple, fragrant sumac, osage orange, and locust
along roadsides, streamsides or woodland edge provide preferred
habitat. Territories may also include parts of the woodland where
the crown canopy is not continuous and there is underbrush.

Movements. — The earliest spring records of the recently arrived
singing males for the years 1951 through 1957 were: March 19,
March 28, April 19, April 4, April 14, April 14, and April 6. Late
dates of singing for the breeding population are: August 21, 1953;
August 13, 1954; and August 10, 1956. Each year territories were
established in the area of the Reservation headquarters. Most com-
plete information was obtained in 1953, 1954, and 1956. In each
of those years sizes of territories covered by singing males were
measured. In 1956 there were two contiguous territories of 6.7
and 4.4 acres, one consisting of two woodland edge areas separated
by a field, and the other consisting of three such units (Fig. 24).
The towhees crossed the fields from one to another of the thicket
areas — a maximum distance of approximately 500 feet, but they
did none of their foraging there and ordinarily they stayed in the
edge of the woods. In 1954 the same area was divided between
three territories, of 4.7, 2.4, and 1.9 acres, of which only the largest
had separate units. In 1953 there were two territories with 6.2 and
4.2 acres, the larger divided into separate units. Pursuits were
observed from time to time. Only males were involved. In at
least one instance the same tree was used by two different males
for singing, on different occasions. On 14 occasions the distances
between males, in adjoining territories, singing and answering each
other, were recorded. These distances ranged from 1000 feet to
100 feet and averaged 238. If this distance represents the average

302

University of Kansas Publs., Mus. Nat. Hist.

Fig. 24. Territories of spotted towhees, as revealed by singing stations of
males (dots) in 1954 and 1956. Activity centered in blackberry thickets
about margins of a field. In 1954 territories were relatively large; one en-
compassed the entire east end of the field and adjoining woodland. In 1956
most of this area was divided between three smaller territories. Although
the towhees usually kept to woodland edge tangles, they occasionally
crossed open fields. Routes followed in two such crossings in 1954 are

shown by arrows in upper figure.

Home Ranges of Vertebrates 303

diameter of a territory, the latter would cover approximately 4.1
acres. This figure agrees fairly well with the 4.4-acre average of
the seven territories actually measured. In the nesting season of
1955 territories near the Reservation headquarters were not con-
tinuously occupied. Towhees arrived on April 13 and singing and
chasing were noted frequently in the following three weeks. Then
they disappeared from the area and were not recorded again until
June 30.

On several occasions a surprising degree of tolerance was noted
between cardinals and towhees, which might even sing from the
same tree simultaneously with no indication of hostility.

Towhees wintering on the area were much more vagile than
those present in summer. A series of 50 distances between succes-
sive captures in males averaged 492 feet (2080 to 40) indicating
an average home range of approximately 17.4 acres. Females
probably wander less widely. Of 16 towhees that were banded
and recaptured, 11 were males, three were females and sex was
unrecorded in two. One female was caught four times in succes-
sion at the same place, and nowhere else. Another female made a
movement of 220 feet and then was caught twice in the same place,
and the third female made a movement of 380 feet.

Barbour (1941:593) studied wintering populations of the red-
eyed towhee in eastern Kentucky. He found them to be associated
in loosely organized flocks. Only 23.7 per cent of the towhees in
these flocks were females, and Barbour surmised that the females
tend to migrate farther south than the males. One flock covered
an area of approximately 31 acres, another 9.4 acres.

Ammodramus savannarum (Gmelin)

Grasshopper Sparrow

Status. — Occasional resident or transient.
Habitat. — Grassland.

Movements. — In the first week of June, 1958, one was seen and
heard several times, singing in fields north of the buildings on the
Rockefeller Tract. When followed, it moved about in an area ap-
proximately 500 feet in diameter, including parts of two fields that
were formerly cultivated and the northwest part of the adjacent

304 Unr'ersity of Kansas Publs., Mus. Nat. Hist.

pasture. In early July, two were singing in the northern part of
the pasture. No grasshopper sparrows have been recorded on the
Reservation.

Passerherbulus caudacutus (Latham)

Leconte Sparrow

Status. — Transient and temporary resident in spring and autumn,
and occasionally in winter.

Habitat. — Dense grass.

Movements. — Specific dates recorded are: October 23, 1953;
November 2 and 6, 1954; March 11 and 14 and October 21 and 23,
1955; several times from March 22 to April 14, 1956; and on March
12 and October 17, and October 24, 1957.

Pooecetes gramineus (Gmelin)

Vesper Sparrow

Status. — Occasional transient.

Habitat. — Meadows and fields; patches of bare ground seem to
be essential.

Movements. — On October 14, 1949, two were seen in a hilltop
field of brome grass near the quarry, in the wheel tracks where there
was little vegetation. On April 10, 1957, one was watched as it
foraged in an almost barren field at the south edge of the Rocke-
feller Tract.

Chondestes grammacus (Say)

Lark Sparrow

Status. — Occasional summer resident.

Habitat. — Meadows, pastures and short-grass prairies where
there are occasional trees.

In 1948 and earlier, when livestock grazed the area keeping her-
baceous vegetation cropped short, lark sparrows were of much more
general occurrence.

Movements. — Earliest records are: April 7, 1952, and April 11,
1953. An old eroded upland cornfield in the northeastern part of
the Reservation had lost most of its topsoil and become dissected
with deep gullies in the nineteen thirties. By 1952 tlie vegetation
was still sparse, consisting chiefly of three-awn grass, lespedeza,
and bindweed; patches of bare ground remained. Because of its
barrenness this area was still used by lark sparrows after they had
disappeared from other parts of the Reservation. On June 11, 1952,

Home Ranges of Vertebrates 305

a pair seen in this field behaved as if they had a nest nearby. On
June 11, 1954, in another part of the field, a bob-tailed fledgling
was flushed, and the parents were nearby. On April 11, 1953, and
May 29, 1955, lark sparrows were seen in the meadow near the
headquarters. On the latter date a male and female were moving
about together, and the male repeatedly chased away another male
in different directions from an area of approximately 300 feet by
500 feet along a gully beside which there were several perch trees.
On April 5, 1956, a group was seen foraging at the northwest comer
of the Reservation where bluestem prairie had been burned off the
previous month, and new grass was beginning to appear.

In 1957 lark sparrows were present in abundance on the Rocke-
feller Tract, in formerly cultivated fields and a pasture. Most ar-
rived in the first week of May. One pair was observed and fol-
lowed frequently in the second week of May, this pair moved about
in an area that was 1050 feet across in one direction and 650 feet
across in another. The entire area encompassed by the birds*
movements was more than 15 acres, but foraging was concentrated
in relatively small parts of it, usually within 100 feet of woodland
edge, or of isolated trees. An adjoining pair had a territory 700
feet across, and they were seen to defend both boundaries against
their neighbors. In foraging, the lark sparrows of a pair tended to
keep within a few yards of each other as they wandered on open
ground. When one flew the other usually would follow within
a few seconds. Many territorial encounters were seen. The males
were most active in these encounters. They would approach each
other with wings drooping and tails spread, and such meetings
usually led to fighting or pursuit. In one encounter between two
pairs a male seemed sexually excited by the presence of the second
pair. After each short flight he would momentarily mount the fe-
male.

Junco hyemalis (Linnaeus)

Slate-colored Junco

Status. — Regular winter visitor, usually most abundant of all
birds during time it is present on area — early October to late April.

Habitat. — Chiefly open situations; fields, meadows and wood-
land edge.

There is some habitat shift according to weather conditions. The
first juncos to arrive in autumn may be present while weather is
still warm, and they tend to keep in deep shade in cool and shel-
tered situations. When ground is snow-covered they forage mostly

306 UNiNTRSiTi' OF Kansas Publs., Mus. Nat. Hist.

in open fields. To roost, the juncos of a flock usually settle in
thick grass — most often they have been flushed from awnless brome
grass which is the dominant grass on the Reservation. When fields
are snow-covered many roosting juncos have been flushed (usually
one at a time) from the protected niches beneath overhanging
banks along ditches and gulUes.

Movements. — From 1949 through 1957 the earhest records of
juncos on the area in fall were: October 22, October 17, October 20,
October 4, September 29, October 17, October 24 and October 11.
Late records in spring include: April 23, 1950; April 8, 1953; May
6, 1954; April 22, 1955; April 20, 1956; April 25, 1957. A few strag-
glers remain in spring, weeks after most juncos of the wintering
population have left. Likewise in early fall while weather is still
mild, weeks before the main influx of wintering juncos, small groups
appear.

In several instances the same banded individuals were caught on
the area after an intervening breeding season. One male was re-
corded in each of four consecutive winters, but each winter most
of the juncos caught were individuals not recorded in previous
years. Of the juncos banded, many were never recaptured. A
total of 76 were recaptured at new locations providing data con-
cerning movements. For a total of 122 recorded movements (dis-
tances between successive points of capture), the average was 485
feet. Not included in this average are movements of five individuals
recaptured after an intervening summer. Distances for these aver-
aged 858 feet, probably indicating that returned migrants altered
their home ranges somewhat, according to changed availability of
food and shelter or other essentials. For 57 movements of males the
average was 604 feet, indicating an average home range of approxi-
mately 26 acres, while for 47 movements of females the average was
434 feet, indicating an average home range of approximately 13
acres. These figures apply to the areas where individuals foraged,
but do not necessarily include the places where these juncos roosted.
Observations on flocks of juncos going to roost or those flushed
after dark have suggested that they may travel beyond the usual
daytime range to find a suitable roosting place. Loose flocks have
been seen streaming across fields and hillsides and gathering in
bushes or low trees before going to roost on the ground in thick
brome grass. Occasionally in walking across fields of brome grass
at night I have flushed juncos in large numbers in areas where none
was to be found by day. These may have travelled considerable
distances, even from beyond the limits of the Reservation, to find
suitable roosting cover.

Home Ranges of Vertebrates 307

Spizella arborea (Wilson)
Tree Sparrow

Status. — Regular winter visitor, late October to April, exceeded
in numbers only by junco.

Habitat. — Various types of open land and edge situations, usually
where there are brush piles or thickets for shelter.

Old fields with weedy mixtures of herbaceous plants provide a
type of foraging ground favored by the tree sparrow. Thickets
whether at woodland edges or in open fields, are preferred escape
shelters. Flocks of tree sparrows mingle with flocks of juncos, but
the latter are somewhat more arboreal.

Movements. — EarHest fall records, 1951 through 1957, are No-
vember 3, November 12, November 13, October 16, October 29,
October 26, and October 23. The latest definite spring record is
April 10, 1957. On that date a large wintering flock was still pres-
ent. Wintering tree sparrows are highly gregarious. Flocks of
several hundred may move about and forage together. Often these
flocks are mixed with juncos. The two species are sufficiently
similar in their habits that they prefer the same type of food and
cover, and they seem completely tolerant. Song sparrows also
frequently intermingle in the flocks, although they prefer denser
low cover and more moist situations. Less regularly, Harris spar-
rows, white-throated sparrows, cardinals, and towhees associate
with the flocks.

A large flock of several hundred tree sparrows stayed in an area
that included weedy fallow fields on the Reservation near its west-
em edge and adjoining it, a milo field bordered on one side by an
intermittent creek with weeds and brush, and on the other by a
county road, and by thickets and debris from a bulldozed hillside.
The milo field and bordering thickets comprised approximately five
acres, and this was the headquarters of the flock, but their entire
range covered a little more than 20 acres, along the creek, and
adjacent areas of fields and thickets. The range was some 2200 feet
long and 550 feet wide. The flock tended to keep together, con-
centrated in one small part of the range, and moved about slowly,
usually following the line of brush along the edges of the field.
TravelUng from one part of the range to another the sparrows
would move in small straggling groups.

In recaptured banded tree sparrows, distances between succes-
sive captures averaged 378 feet, indicating home ranges of approxi-
mately ten acres in winter.

308 University of Kansas Publs., Mus. Nat. Hist.

Spizella passerina (Bechstein)
Chipping Sparrow

Status. — Migrant and occasional temporary resident in spring.

Habitat. — In the breeding season chipping sparrows choose for
territories areas of parkland and groves, with short ground vegeta-
tion. Lack of such areas with suitably short ground vegetation
seems to prevent establishment of a permanent population on the
Reservation.

Movements. — In 1951, in the last week of April and the first two
days of May, a chipping sparrow was heard and seen frequently,
singing from a large elm beside the house at the Reservation head-
quarters. It soon disappeared from this area. In early May, 1957,
one briefly established a territory on the Rockefeller Tract. This
sparrow was nearly always heard singing from the same large elm,
in the edge of woodland adjoining a cultivated field. It disappeared
within a week of the time it was first noticed.

Spizella pallida (Swainson)

Clay-colored Sparrow

Status. — Occasional transient.

Habitat. — Thickets and woodland edges.

Movements. — The only record is of an individual foraging in
short grass and weeds beside tlie Reservation residence at mid-day
on April 29, 1951.

Spizella pusilla (Wilson)

Field Sparrow

Status. — Regular summer resident; one of most abundant birds
from April till October.

Habitat. — Border between woodland and grassland, and espe-
cially former pastures and cultivated fields that are in process of
growing up into thickets.

Since cultivation and grazing were discontinued on the Reserva-
tion in 1948, the habitat has steadily improved, as brush and sap-
lings have encroached into the fields. Occasionally, field sparrows
have been found established in woodland, where canopy was open
and understory vegetation was not dense. So far as observed, they
occupy open areas only where thickets are available.

Movements. — Field sparrows appear on the area earlier than any

Home Ranges of Vertebrates

309

other summer residents that do not spend the winter there. Table 3
illustrates the seasonal schedule.

Each year singing was first heard on mild sunny days of late
winter. Such early season singing was of a pecuHarly subdued
quality. With the return of stormy or cold weather, singing stopped
altogether for periods of days, or even weeks. In February and

Table 3. — Year by Year Record of Events in the Annual Cycle of
THE Field Sparrow on the University of Kansas Natitoal History

Reservation.

Year

Earliest
record

Earliest
singing

Earliest
pairing

Last
record

IQJiO

October 26

1951

19i2

1953

1954

1955

1956

1957

March 5
February 10
February 7
March 25
March 3
March 17
March 11

March 16
March 16
February 7
March 25
March 3
March 17
March 11

March 25
March 16
March 13
April 2

October 30
October 5
November 12

April 1

early March the sparrows were present in small numbers, and new-
comers might arrive gradually over a period of weeks. By mid-
April most or all had arrived and established themselves in terri-
tories.

At this time of year males sing persistently from elevated perches
wherever tliese are available in the territory. Males usually sing
from perches five to twenty feet above the ground. In singing the
bird is usually perched well inside the crown of the tree, and almost
never perches on the top, unless it is a small sapling. Arrival of
additional field sparrows makes necessary readjustments of terri-
tories as those estabhshed early are compressed. On March 26, 1954,
only one field sparrow was present in the field near the Reservation
headquarters. As it moved from tree to tree singing along wood-
land edge it was followed for a distance of 820 feet, indicating an
unusually large territory. At times when the sparrows were pres-
ent in normal numbers 500 feet seemed to be about the maximum
diameter of any territory.

Even in April, territoriality may be temporarily suppressed in
inclement weather. This was strikingly demonstrated when as
many as six field sparrows were caught simultaneously in the same
funnel trap in rainy weather. In rain singing stopped. The spar-

310 Unhtrsity of Kansas Publs., Mus. Nat. Hist.

rows were reluctant to forage in their usual place, in high grass
where their plumage might soon become soaked; instead they con-
gregated in open places chiefly along the road. Several pairs
might forage within a few square yards, with no evidence of hos-
tility, although the area involved was within the territory of one
pair and beyond the usual boundaries of the others. With the
return of clear weather the sparrows withdrew to their established
territories with resumption of singing, and intolerant behavior
toward intruders. However, even in mild weather singing and ter-
ritorial behavior was much more prominent in the morning and was
partly suppressed later in the day, with some intermingling between
different pairs.

The approximate extent of a territory could soon be determined
by flushing a singing male from perch to perch. Within a few
minutes he would make the circuit of his territory. Seven terri-
tories that were traced in some detail, in 1952, 1953, 1954, and 1956,
averaged 1.6 acres (2.5, 2.2, 1.8, 1.6, 1.4, 1.0, .8). The largest were
those that extended over open areas. Those that were relatively
small were partly beneath trees and in brushy places.

In 1950, 58 pairs were located in an area of approximately 300
acres, the northwestern half of the Reservation. They tended to
be linearly arranged in edges of fallow fields and former pastures
bordering woodland. In a few instances there were intervals of
hundreds of yards between pairs, but in most instances territories
were contiguous and actual distances between the birds of adja-
cent pairs were 100 to 600 feet. For 46 such intervals the average
was 320 feet. A territory of this diameter would cover 1.8 acres.

Territorial fighting and pursuits were seen frequently, especially
in April. In 1954 a remarkably prolonged territorial quarrel was
observed. On April 6, when the population was increasing rapidly
through an influx of new migrants, two males were noticed perching
and hopping about nervously within a few inches of each other,
each trying to round up the other and force it back. There was no
actual fighting, and neither bird would give ground. On April 13
two males, probably the same, were observed in the same area, and
there were frequent long chases. Both males had been banded. The
pursued sparrow, always the same, fled consistently but would not
leave the territory. Several times the chase led to a female, who
did not participate in the quarrel. After several hours the chase
was still in progress. It was continued on the following day. Sel-
dom more than a few seconds elapsed until the aggressor renewed

Home Ranges of Vertebrates 311

the pursuit, and the birds might be in motion 30 seconds or more
before the next pause. As before, the pursued persistently remained
within the disputed area, and the pursuer, so far as observed, did
not press to the point of actual attack. Continuations of the pursuit
were observed on the 16th, 19th, 20th, 23rd, 24th, and 26th. When-
ever the pursuer flew back to the female he was followed by the
other male, whose presence distracted him and disrupted the nor-
mal course of courtship and nesting. Eventually the pursuits be-
came less spirited, with longer intervening pauses, during which
the pursued, the pursuer and the female perched or foraged in
close proximity. On the morning of April 26, the pursuing male
was bedraggled in appearance, with feathers disarranged and some
missing. Obviously the continuous tension was telling on him. The
pursued showed no such effects. However, the pursuer may have
become involved in fights with other outsiders and could not have
maintained his boundaries against adjoining pairs while distracted
by the rival within the area. Later the same day he was located
singing in a more open and less favorable area beyond the limits
of his former territory. It seemed that he had abandoned both
territory and mate to the interloper whom he had failed to dislodge
in two weeks of almost unremitting effort.

In 1953 the same territory contested by these two males was
occupied by different individuals. The bob-tailed male was color-
banded on April 6, probably soon after his arrival. On April 7 he
was seen associating briefly with each of two different females, one
of which had been color-banded. On April 8 he had obtained a
mate, and the pair moved together about the territory. As they
foraged near together on the ground, movements involved a steady
advance of the male, and retreat of the female to avoid actual con-
tact. When the male moved toward her for a few inches, the female
would quickly hop away, maintaining her distance, and then would
pivot around facing the male or standing broadside to him. How-
ever, whenever the male flew, she promptly followed. Later in
the day the male became increasingly aggressive, and frequently
would cause the female to flush and would chase her back and forth
across the territory several times without pause. Finally he caught
her, and both fell into the wet grass, fluttering and pecking. They
lay sprawled and interlocked, fluttering feebly and pecking at each
other for more than two minutes. The female broke away but im-
mediately the male gave chase and caught her, and again they
fell into the grass, interlocked, and remained there almost as long

312 University of Kansas Publs., Mus. Nat. Hist.

as they had the first time. After disengaging, they flew to a for-
aging place preening and shaking, the hostilities ended. On the
following day the female was no longer present. The male's overly
aggressive and hostile behavior may have caused her to desert the
territory, and for several weeks thereafter the male remained un-
paired.

Singing reaches a peak in April and tapers off later in the season.
In April song may be heard at any time of day, or even at night.
In June and July singing occurs chiefly at daybreak. Occasionally
singing is heard in August and early September.

On several occasions the development and dispersal of broods
were observed. On October 2, 1952, three or four fledglings of an
unusually late brood, probably just out of the nest, were chirping
within an area of a few square yards. On the following day they
had shifted about 70 feet farther east and were more scattered,
still concealed in high grass. The adults were delivering food fre-
quently. On the following day they had moved still farther in the
same direction. Two were flushed 200 feet and 270 feet from the
original location. They were still bob-tailed ( tails less than one inch
long ) and their flights were short and weak. On October 7 one was
observed near the same place it was seen October 4. It had
emerged from the grass, and was fluttering about in a bush as the
adults came to feed it at frequent intervals. On October 8, one
was near the same place perched ten feet above ground in a small
tree, chirping steadily. The two other young were chirping in
bushes 30 feet farther southeast. One hopped and fluttered along
the ground to the base of the tree where the first young bird was
perched, and an adult came and fed it there. Soon afterward the
young that was perched ten feet high flew strongly for 100 feet
west and within a few minutes made several more flights. Late in
the afternoon all three young were assembled in the same tree.
Their tails were about VA inches long. Later one young was seen
following an adult as the latter moved about slowly, foraging on
the ground in high grass. The fledgling's peeping changed to more
metallic chirps as the adult approached to feed it.

In the following two weeks the family group remained intact and
the young and adults were often seen foraging together in or about
a dry pond bed some 450 feet east of the nesting area, outside the
original territory. By October 21 the fledglings' tails had grown to
full length, and the young birds were able to fly rapidly and strongly.
They were moving about freely ^vith no attempt at concealment.
On October 24 the fledglings were foraging independently most of

Home Ranges of Vertebrates 313

the time though occasionally one would give a juvenal chirp. The
young field sparrows were mingling freely with recently arrived
juncos and song sparrows, and with field sparrows of other families.

Zonotrichia querula (Nuttall)
Harris Sparrow

Status. — Common winter resident, also migrant in spring and
autumn.

Habitat. — Thick brush for shelter adjoining open land, with
weeds or cultivated crops providing a food supply, are the chief
requirements on the winter range in northeastern Kansas.

Movements. — In the winter of 1956-1957, a flock of these spar-
rows made their headquarters in several large brush piles where a
grove of osage orange trees had been cut and bulldozed, on the
southern edge of the Rockefeller Tract adjoining the Reservation.
They foraged in an adjacent field of milo stubble, and ranged across
this field to tlie brushy woodland edge, 900 feet north of the favored
brush piles. Also they ranged 800 feet east of the brush piles, in
thickets bordering the county road.

Of 71 Harris sparrows banded, only 11 have been recaptured. In
eight recaptured within a few weeks of banding, distances (in feet)
were as follows: 0, 0, 40, 50, 70, 180, 360, 1030. One that was
caught at a brush pile on the Rockefeller Tract was brought to the
Reservation headquarters for banding, and released there. Several
hours later it was recaptured only 100 yards from the place of re-
lease, having remained in the vicinity instead of returning half a
mile to tlie flock and home range. Two of those banded were re-
captured in later seasons. One of these, banded March 17, 1955,
was recaptured 600 feet farther along the same fence row thicket
on February 3, 1956. The other, banded on January 22, 1955,
was retrapped on January 28, 1957, 1800 feet away ( near the brush
pile on the Rockefeller Tract). Harris sparrows have been seen as
early as October 5 (1953) in fall and as late as May 10 (1957), in
spring.

Zonotrichia leucophrys (Forster)

White-crowned Sparrow

Status. — Occasional migrant.

Habitat. — Thickets, especially those at margins of weedy fields.

Movements. — On October 30, 1952, one was seen at the brushy
edge of a ravine near the Reservation headquarters. On November
8, 1954, one was seen one hundred yards farther east at the pond.

314 University of Kansas Publs., Mus. Nat. Hist.

One was trapped with a group of Harris sparrows on February 5,
1958. These are the only definite records for the area, but on Octo-
ber 21, 1951, several were seen in an osage orange thicket beside
the county road near the southeast corner of the section.

Zonotrichia albicollis (Gmelin)

White-throated Sparrow

Status. — Regular migrant in spring and fall, and occasional winter
resident.

Habitat. — On Reservation chiefly in woodland edge thickets.

Movements. — Autumn appearances in several years were: Octo-
ber 14, 1949; October 13, 1950; October 17, 1951; October 30, 1952;
October 15, 1953. For a week or two after the first arrival these
sparrows were abundant and conspicuous in such situations as
brush piles, woodland edge thickets, and weedy margins of the
pond, but they soon moved on. Spring influxes were noted on April
26, 1951; April 28, 1952; April 21, 1953; April 17, 1954; and April
25, 1956. These migrants were gregarious, and flocks of a dozen
or more settled temporarily in thickets providing shelter and food,
often remaining two or three weeks. Latest spring records are May
8, 1954, and May 4, 1956. The woodland edge thickets between the
pond and the Reservation headquarters had a flock almost every
spring. At times their song was one of the most prominent in the
spring chorus. They were usually associated with Harris sparrows.
Occasional individuals remain all winter in the same situations
where flocks are seen in spring and fall. One banded on December
9, 1954, was recaptured regularly ( 14 times in all ) up to April 10,
1955. The area encompassed by its records was 700 feet long and
150 feet wide. The flocks present in spring have often been seen
feeding in trees, on leaf buds and seed clusters of elm, while in
autumn seeds of weeds seem to provide most of the food.

Borror (1948:428) found that migrating white-throated sparrows
settled temporarily in "stopover ranges" up to an acre or two in size
(average of greatest distances between captures for 74 repeaters
was 58 yards). Borror found that in fall the stay was somewhat
longer and the range somewhat larger than in spring. Fischer and
Gill ( 1946:407) in a report based on analysis of the data from 43,000
banded white-throated sparrows, found that returns from the birds
banded while on migration were rare.

Home Ranges of Vertebrates 315

Passerella iliaca (Merrem)
Fox Sparrow
Status. — Occasional transient.
Habitat. — Woodland thickets.

Movements. — Recorded from the Reservation on only a few
occasions. On November 5, 1954, several were seen in weed thick-
ets near the edge of the pond. These birds must have settled tem-
porarily in the vicinity, as they were seen frequently through No-
vember 17. R. W. Fredrickson reported seeing one on October 23,
1955. On March 1, 1956, one was seen at a weed patch beside the
pond. It had perhaps stopped for the day at this place, as it was
seen twice, with several hours intervening.

Melospiza lincolnii ( Audubon )

Lincoln Sparrow

Status. — Regular transient and temporary resident.

Habitat. — Low, dense thickets in swampy places; on Reservation
thick grass is usual habitat.

Movements. — Arrival dates recorded over several years were
April 28, 1951; April 25, 1952; April 25, 1953; April 10, 1954; April
26, 1956. Curiously, none was seen in the spring of 1955. In
1952 one was banded and was caught a total of six times within
the period April 27 to May 7. All captures were made with live-
traps set for mice, and all were well within the area covered with
traps in a 50-foot grid. The most remote points of capture were
500 feet apart; however all but the first capture were within a 30-
foot radius. Although only one other banded Lincoln sparrow was
recaptured ( 50 feet from the original site ) , the impression is borne
out that individuals stopping on their northward migration settle
for periods of days or weeks in small areas. Patches of foxtail
grass (Setaria viridis) near the edge of woodland at the Reservation
headquarters were much used by these sparrows in late April, 1956.
When disturbed, they would fly back to brush a few yards away,
and were seen at these small grass patches many times daily. In
1956 the sparrows were observed in much greater numbers than
they had been in any previous year. One was seen on October 13,
1955, and in the last week of September, 1956, and on October 1,

316 University of Kansas Publs., Mus. Nat. Hist.

many were seen at the fo.xtail patches near the headquarters. An-
other was caught many times at the same brush pile in late Decem-
ber, 1955, indicating that occasional individuals winter in the area.

Meiospiza georgiana ( Latham )

Swamp Sparrow

Status. — Rare migrant.

Habitat. — Swamps and marshes.

Movements. — R. W. Fredrickson reported one on April 14 and
another on April 27, 1952. On several other occasions in late
autumn swamp sparrows have been flushed from a patch of smart-
weed beside the dry pond bed.

Meiospiza melodia (Wilson)

Song Sparrow

Status. — Regular winter resident, present in abundance from Oc-
tober into April.

Habitat. — Fields, meadows, and marshy places with dense ground
vegetation satisfy the habitat requirements of the song sparrow in
winter. Fence row thickets of dogwood, various saplings and vines
are frequented, and brush piles provide preferred escape covert.
Greatest concentrations were noted in marshy areas about the edge
of the small pond, where there were tangles of smartweed, rice
cutgrass, cat-tail and other marsh vegetation. In old-field areas
with weedy mixtures, patches of foxtail grass ( Setaria sp. ) provided
preferred types of food and shelter.

Movements. — From 1952 to 1957, inclusive, October appearances
of immigrating song sparrows were on the 19th, 11th, 5th, 17th, 21st
and 16th. Last dates of record, in April each year from 1953
through 1956, were the 24th, 10th (except for a transient heard
singing on the evening of June 3), 20th, and 26th. Though requir-
ing dense cover, a song sparrow having settled in its winter quar-
ters requires only a small area. For 29 banded and recaptured song
sparrows, distances between successive capture points, representing
a total of 35 movements, ranged from 1040 feet to 40 feet and aver-
aged 300 feet. Assuming this distance to represent an average home
range radius, home ranges of approximately 6.8 acres are indicated.
The records of one individual caught eleven times in the winter of
1953-54, encompassed 8.9 acres. Only one individual was recap-
tured in the same winter range after an intervening summer.

Home Ranges of Vertebrates 317

SUMMARY AND CONCLUSIONS

In this study of spatial relationships, 3 species of fish, 10 of
amphibians, 24 of reptiles, 29 of mammals, and 167 of birds are
discussed. In an area under observation so regularly and inten-
sively as the University of Kansas Natural History Reservation, the
negative evidence obtained through the lack of records of species
that might be expected to occur is significant. For example,
among the species recorded in Douglas County, or on all sides of it,
but never recorded from the Reservation itself, there are five kinds
of amphibians, 22 kinds of reptiles, 20 kinds of mammals, and 131
kinds of birds. Many of these are rare species which might be ex-
pected to appear only at widely spaced intervals, and several of
them perhaps ultimately will appear on the Reservation. Another
large group of species are those characteristic of extensive fluvia-
tile or marshland habitats that are not represented on the Reser-
vation. A total of 81 species belong to this category. They include
river turtles, water snakes, beaver, otter, mink, loons, grebes, herons,
ducks, shorebirds, terns, and others. Among the mammals are eight
species of bats which might be expected but have not yet been re-
corded because of inadequate sampling. Approximately 26 species
are characteristic of prairie habitat, and presumably do not occur
on the Reservation because of the hmited extent, isolation, or un-
suitabiHty of the prairie relict there. About the same number of
kinds are characteristic of deciduous forest habitats of the eastern
United States (notably the warblers, among birds).

Among those species that have been recorded from the Reserva-
tion approximately half the total reach the area only as wanderers,
and have no permanent population on the area. These include the
tiger salamander, spadefoot toad, painted turtle, woodchuck, house
cat, American rough-legged hawk, goshawk, bald eagle, killdeer,
Franklin's gull, barn owl, belted kingfisher, and a number of other
kinds of birds that merely fly over the area, without stopping on it.
Also included in this category are the great majority of individuals
of Woodhouse's toad, muskrat, starling, house sparrow, marsh
hawk and perhaps a few others, although individuals of each of
these kinds has been known to live on the Reservation.

There are few species, if any, which wander completely at ran-
dom. However, there seems to be no clear-cut line of demarcation
between those that habitually wander and those that occupy definite
home ranges. All degrees of intermediacy have been found be-
tween the two categories. The snapping turtle may be considered

318 University of Kansas Publs., Mus. Nat. Hist.

a typical representative of the species with strongly developed
nomadic tendencies. At times individuals of various sizes and of
both sexes wander for long distances up and down the intermittent
stream courses. They also may frequent certain pools or ponds
or sections of a stream, habitually using an available shelter or food
supply. That a home range having definite boundaries exists is
not demonstrable. In the bullfrog, leopard frog, narrow-mouthed
toad, cricket frog and chorus frog, wandering tendencies are
marked, and the animals' habits of staying at the same pool, or be-
neath the same rock, for periods of days or weeks may be forced
upon them by drought limiting their movements, rather than by
any aflBnity for the location where the animal happens to be. In
the American toad and Woodhouse's toad, an individual returns
regularly to the same diurnal shelter, over periods of days or weeks,
at least, and emerging to forage in the evening, it may cover more
or less the same area on successive occasions. However, even in
these animals, the home range seems to have no definite limits, ex-
cept where natural barriers prevent free movement, and wandering
tendencies may at almost any time cause the toad to shift to a
new area more or less remote from its former range.

In most kinds of hzards, snakes, small mammals, and birds for
which sufficient data were obtained, the trend was somewhat simi-
lar, with varying degrees of attachment to a familiar area, but with,
at best, vaguely defined boundaries that are subject to frequent al-
teration, and with occasional wandering. Dice and Clark (1953:10)
have emphasized that one respect in which home ranges differ from
territories is the poorly defined boundaries of the former, and they
have suggested defining the home range in terms of the average
radius, rather than as a definite area.

In a few species, notably the collared lizard, the larger snakes
{Coluber, EJaphe, Agkistrodon) , the blue jay, and (in winter)
chickadee, titmouse, and cardinal, attachment to a home area is
strong, and the same adult may remain for periods of years in ap-
proximately the same range. In these same species other indi-
viduals may wander, both frequently and extensively, and the
average occupancy of an area is short.

Territoriality exists in most of the species of birds that breed on
the area, and is less developed in the collared hzard, the woodrat
and perhaps a few other kinds of small mammals. In its most
typical form a territory constitutes a defended home range, and
possession of a well defined home range is a prerequisite to ter-

Home Ranges of Vertebrates 319

ritoriality. All stages in the development of territoriality were seen
in the numerous species investigated. In some, like the skinks
(Eumeces sp.), the possession of a regular home range is associated
with hostility (in the adult male, especially in the breeding season).
Fighting is frequent, and the establishment of a true territory is per-
haps prevented only by the retiring and secretive habits, which
normally prevent detection of a rival until it has come within close
range.

Among the birds there are relatively few kinds having permanent
territories that live on the area; the red-tailed hawk, homed owl,
barred owl, hairy woodpecker and Carolina wren are the most
typical of those kinds that do have permanent territories. Even
in these species territorial boundaries are flexible, and are subject to
seasonal alteration. In other species, some of which may remain
permanently mated, territorial boundaries are largely dissolved at
times of year other than the breeding season. A pair may continue
to use its former territory, but at the same time may mingle v^dth
other members of a social group, and range over a much more
extensive area than the territory had encompassed. The blue jay,
crow, tufted titmouse, black-capped chickadee, downy woodpecker
and red-belhed woodpecker are among the best examples of this
group. In contrast, only temporary and seasonal attachment to
territory and mate is charactertistic of the bob-white, mourning
dove. Cooper's hawk, bluebird and goldfinch among resident spe-
cies. Most species of birds that breed on the area make annual mi-
grations, abandoning both territory and mate. The territory seems
especially subject to alteration, and may change continually in size,
shape and position, according to immediate needs at the particular
stage of the nesting cycle in these species that have only temporary
attachment to it.

Size of territory is somewhat characteristic of each species.
Among the migratory species, males that are early arrivals in spring
may occupy unusually large territories; an instance is cited in the
species account of the field sparrow. In this sparrow the breeding
population increases over periods of weeks and the new arrivals
must usurp parts of areas that are already occupied in order to
establish their territories. The males or pairs that are established
early are then subjected to increasing territorial pressure, with en-
forced retrenchment. In many other species, whose populations
are sparse, including the barred owl, Carolina wren, bluebird, wood

320 University of Kansas Publs., Mus. Nat. Hist.

thrush, Kentucky warbler, and summer tanager, a pair establishing
a territory was usually subject to but little territorial pressure or to
none at all, because territories seldom occupied adjoining areas.
Even under these conditions, the pair usually tended to confine their
activities to a relatively small central area, which perhaps should
be considered the whole territory, but the daily routine was varied
from time to time by visits to any of several outlying points.

For territories and home ranges alike, one of the chief factors
determining size was the distribution of suitable habitat. Where
the essential features of the habitat were scarce and scattered, the
individual was forced to cover a relatively large area. This is well
illustrated by the red-tailed hawk territories shown in Fig. 11; the
one including a high proportion of cultivated land, unproductive of
prey was consistently larger than the other. A more extreme ex-
ample is provided by the territories of towhees shown in Fig. 24.
Blackberry thickets at the edges of woodland were the essential
features and the territory might include several of the thickets and
also intervening areas of less favorable habitat — brush, groves, or
even open fields. Towhees crossed the fields regularly in moving
from one blackberry patch to another. Although they did not for-
age in the fields, tliey occasionally did stop in isolated trees to sing
there. Therefore, part of a field might be considered part of a
towhee's territory, even though it furnished neither food nor shelter,
and was significant to the bird only as an intervening space, to be
crossed at some hazard. In the mourning dove and goldfinch, ac-
tivities are still more extended; in some instances foraging areas are
so remote from the nest that they could not be construed as part of
the territory, and intermediate stages of all degrees exist.

Odum and Kuenzler (1955) have emphasized the ephemeral
nature of territories in many kinds of birds, and have described
fairly rapid alteration in size and shape. They imply that territories
become progressively smaller later in the season, with the transi-
tion from nest building to incubation, to feeding of young. Most
territories are established early in the breeding season, and the
pair, having successfully defended their chosen area against inter-
lopers during this stage, may not be required to withstand much
territorial pressure in later stages. Rather, they may be free to
wander through the original territory, and to concentrate on those
parts of it that best satisfy their seasonal requirements.

In certain kinds of birds, exemplified by the raptors, phoebe,
chickadee, titmouse and others, the members of a pair share more

Home Ranges of Vertebrates 321

or less equally in defense and utilization of their territory, and
characteristically travel together. In other kinds exemplified by
the horned lark, the redwing, the indigo bunting and the dickcissel,
the sexes di£Fer greatly in habits, and territorial defense devolves
chiefly upon the male. He moves about more frequently and more
conspicuously than the female, and usually covers a somewhat
larger area. In mammals, although most are not clearly territorial,
the same tendency toward a larger range in the male was manifest.
In the opossum, fox squirrel, gray squirrel, deer mouse, and prairie
vole, the male's range is markedly larger than the female's. In
the cotton rat, harvest mouse, white-footed mouse and cottontail,
the difference between the sexes, in size of area covered, is slight.
In the larger snakes (garter snake, yellow-bellied racer, black rat
snake, copperhead) the adult male's range is at least twice the
size of the female's, and in the common lizards including collared
lizard, Great Plains skink, and glass lizard, a comparable disparity
exists. The six-lined racerunner seems to be a notable exception
to the general trend, as the female's range averaged 40 per cent
larger than the male's, in my sample.

In general, young animals ranged less widely than adults of the
same species. As the young animal grows and becomes familiar
with its surroundings, it gradually increases its range. Although
individuals of some kinds of animals remain permanently near the
place where they were born, dispersive wandering is characteristic
of the young of most kinds. In amphibians dispersal occurs chiefly
at the time of metamorphosis, and in reptiles it may occur after
birth or hatching. In mammals the dispersal takes place after
weaning, and in birds usually just after the fledglings have learned
to forage for themselves.

Among the factors that determine size of home range, potential
mobility of the animal is, of course, important, but the correlation
is not close. The smallest home ranges, less than half an acre in
area, were found in lizards, toads, voles, the woodrat, and the white-
footed mouse — animals with limited capacity for rapid travel.
Home ranges or territories in the size range of half an acre to two
acres were found in the remaining small mammals, the short-tailed
shrew, jumping mouse, cotton rat, harvest mouse, house mouse,
and in several species of birds, mostly the smaller kinds, and espe-
cially those that live in thick woods or in dense ground vegetation
— pewee, Bell vireo, gnatcatcher, long-tailed chat, blue grosbeak,
and bob-white. Areas from two to ten acres in extent are occupied

322 Untversity of Kansas Publs., Mus. Nat. Hist.

by individuals of most kinds of small birds, by the cottontail, and
the ornate box turtle. Relatively large areas, ten to fifty acres in
extent, are utilized by the larger snakes, including the copperhead,
black rat snake, yellow-bellied racer, and common garter snake, by
certain birds, including those that are persistent fliers — the homed
lark, bam swallow, whip-poor-will, chuck-will's-widow and hairy
woodpecker — and by various small passerines on their winter ranges
when they are not territorial — junco, tree sparrow, towhee, chick-
adee, and titmouse. The opossum, fox squirrel, and gray squirrel
also have home ranges within this size range. Territories and home
ranges larger than 50 acres were found to be maintained by only
a few kinds of vertebrates, nearly all of them predators and all rela-
tively large. They include most, if not all, of the hawks and owls
living on the area (the screech owl is a probable exception), the
turkey vulture, the coyote, red fox, spotted skunk, and white-tailed
deer.

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1956. Colonies of great blue heron (Ardea herodias) in Kansas. Trans.
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Barbour, R. W,

1941. Winter habits of the red-eyed towhee in eastern Kentucky. Amer.
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1939. Territory and population in the great homed owl. Auk, 56:274-
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1940. Life histories of North American cuckoos, goatsuckers, humming-
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1940. Home ranges and populations of the jumping mouse. Amer. Midi.
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1942. Size of home range and notes on the life history of the woodland
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1948. Analysis of repeat records of banded white-throated sparrows.
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Home Ranges of Vertebrates 323

Breckenrtoge, W. J.

1943. The life history of the black-banded skink, Eumeces septentrionalis

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1955. Size of home range in eight bird species in a southern Illinois
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1951. An ecological survey of the Fort Leavenworth MiUtary Reservation.
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1940. Territorial behavior and populations of some small mammals in

southern Michigan. Misc. Publ. Univ. Michigan 45, 58 pp., 2 pis.,

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Cade, T. J.

1955. Experiments on winter territoriality of the American kestrel, Falco
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Carpenter, C. C.

1952. Comparative ecology of the common garter snake (Thamnophis
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1953. A study of hibernacula and hibernating associations of snakes and
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Chapman, H. H.

1947. Some observations on the CaroHna wren in La Salle Parish, Lou-
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Crabb, W. D.

1948. The ecology and management of the prairie spotted skunk in Iowa.
Ecol. Monogr., 18:201-232.

Dice, L. R.

1952. Natural Communities. Univ. Michigan Press, Ann Arbor, xii +
547 pp., 52 figs.

Dice, L. R., and Clark, P. J.

1953. The statistical concept of home range as applied to the recapture
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Dice, L. R., and Howard, W. E.

1951. Distance of dispersal by prairie deermice from birthplaces to

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1 map,
DiLGER, W. C.

1956. Hostile behavior and reproductive isolating mechanisms in the
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DRxmi, M.

1939. Territorial studies on the eastern goldfinch. Wilson Bull., 51:
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324 UNi\'ERsrri' of Kansas Publs., Mus. Nat. Hist.

Engels, W. L.

1949. Crested flycatchers nesting some distance from their foraging area.
Auk, 66:287-288.

Fischer, R. B., and Gill, G.

1946. A cooperative study of the white-throated sparrow. Auk, 63:
402-418.
Fitch, F. W., Jr.

1950. Life history and ecology of the scissor-tailed flycatcher, Musdvora
forflcata. Auk, 67:145-168.

Fitch, H. S.

1949. Study of snake populations in central California. Amer. Midi.
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1952. The University of Kansas Natural History Reservation. Umv.
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1954. Life histor>' and ecology of the five-lined skink, Eumeces fasciatus.
Univ. Kansas Publ. Mus. Nat, Hist., 8:1-156.

1955. Habits and adaptations of the Great Plains skink (Eumeces
obsoletus). Ecol. Monogr., 25:59-83.

FrrcH, H. S., and Fitch, V. R.

1955. Observations on the summer tanager in northeastern Kansas. Wil-
son Bull., 67:45-54.

Fitch, H. S., and Lokke, D. H.

1956. The molluscan record of succession on the University of Kansas
Natural History Reservation. Trans. Kansas Acad. Sci., 59:442-454.

Fitch, H. S., and McGregor, R. L.

1956. The forest habitat of the University of Kansas Natural History
Reservation. Univ. Kansas Publ. Mus. Nat. Hist, 10:77-127.
Fitch, H. S., and Packard, R. L.

1955. The coyote on a natural area in northeastern Kansas. Trans. Kan-
sas Acad. Sci., 58:211-221.

Fitch, H. S., and Rainey, D. G.

1956. Ecological observations on the woodrat, Neotoma floridana. Univ.
Kansas Publ. Mus. Nat. Hist., 8:499-533.

Fitch, H. S., and Sandidge, L. L.

1953. Ecology of the opossum on a natural area in northeastern Kansas.
Univ. Kansas Publ. Mus. Nat. Hist., 7:305-338.

Fitch, H. S., Swenson, F., and Tillotson, D. F.

1946. Behavior and food habits of the red-tailed hawk. Condor, 48:
205-237.
Freiburg, R. E.

1951. An ecological study of the narrow-mouthed toad (Microhyla oliva-
cea) in northeastern Kansas. Trans. Kansas Acad. Sci., 54:374-386.

Gloyd, H. K.

1934. Studies on the breeding habits and young of the copperhead,

Agkistrodon mokasen Beauvois. Papers Michigan Acad. Sci. Arts

and Letters, 19:587-604, pis. 77-79.
GoiN, C. J., and Coin, O. B.

1954. Nest-building behavior of the Carohna wren. Wilson Bull., 66:59.

Home Ranges of Vertebrates 325

Hall, E. R.

1956. Handbook of mammals of Kansas. Univ. Kansas Mus. Nat. Hist.
Misc. Publ. 7, 303 pp., 100 figs.
Haugen, a. O.

1942. Home range of the cottontail rabbit. Ecology, 23:354-367.
Hayne, D. W.

1949. Calculation of size of home range. Joum. Mamm., 30:1-18.
Hensley, M, M.

1950. Notes on the breeding behavior of the Bell's vireo. Auk, 67:243-
244.

How^ard, H. E.

1948. Territory in bird life. Collins, London. 224 pp.
Imler, R. H.

1945. Bullsnakes and their control on a Nebraska wildlife refuge. Joum.
Wildlife Mgt., 9:265-273.
Janes, D. W.

MS The home range and movements of the eastern cottontail in Kan-
sas. Unpubhshed thesis, Univ. Kansas Library.
Laskey, a. R.

1940. The 1939 nesting season of bluebirds at Nashville, Tennessee.
Wilson Bull., 52:183-189.

Leonard, A. B., and Goble, C. R.

1952. Mollusca of the University of Kansas Natural History Reservation.
Univ. Kansas Sci. Bull., 34:1013-1053, 2 pis.
MacQueen, p. M.

1950. Territory and song of the least fl>'catcher. Wilson Bull., 62:
194-204.

Manville, R. H.

1949. A study of small mammal populations in northern Michigan. Misc.
Publ. Mus. Zool. Univ. Michigan, 73:83 pp., 4 pis., 6 figs., 1 map.

Martin, E. P.

1956. A population study of the prairie vole (Microtus ochrogaster) .
Univ. Kansas Publ. Mus. Nat. Hist., 8:361-416.
Nero, R. W.

1956. A behavior study of the red-winged blackbird. IL Territoriality.
Wilson Bull., 68:129-150.
Nickell, W. p.

1951. Studies of habitats, territory and nests of the eastern goldfinch.
Auk, 68:447-470.

Odum, E. p.

1941. Annual cycle of the black-capped chickadee. Auk, 58:314-333;
518-535.

1942. Annual cycle of the black-capped chickadee. Auk, 59-499-531.
Odum, E. P., and Kuenzler, E. J.

1955. Measurements of territory and home range size in birds. Auk,
72:128-137.

Packard, R. L.

1956. The tree squirrels of Kansas: ecology and economic importance.
Univ. Kansas Mus. Nat. Hist. Misc. Publ., 11:1-67, 2 pis., 11 figs.

326 University of Kansas Publs., Mus. Nat. Hist.

Putnam, L. S.

1949. The life histor>' of the cedar waxwing. Wilson Bull., 61:141-182.
QUIMBY, D. C.

1951. The life history and ecology of the jumping mouse, Zapus hud-
sonius. Ecol. Monogr., 21:61-95.

Rainey, D. G.

1956. Eastern wood rat, Neotoma fioridana: life history and ecology.
Univ. Kansas Publ. Mus. Nat. Hist., 8:535-646, 12 pis., 13 figs.
S\UTH, H. B.

1956. Handbook of amphibians and reptiles of Kansas. Univ. Kansas
Mus. Nat. Hist. Misc. Publ., 9:357 pp., 232 figs.
Smith, W. P.

1942. Nesting habits of the eastern phoebe. Auk, 59:410-417.
Stew.\rt, R. E.

1949. Ecology of a nesting red-shouldered hawk population. Wilson
Bull., 61:26-35.

1953. A life history study of the yellow-tliroat. Wilson Bull, 65:99-115.
Stickel, L. F.

1950. Populations and home range relationships of the box turtle, Terra-
pene c. Carolina (Linnaeus) Ecol. Monogr., 20:351-378.

Stickel, W. H., and Cope, J. B.

1947. The home ranges and wandering of snakes. Copeia, 1947:127-136.
Stokes, A. W.

1950. Breeding behavior of the goldfinch. Wilson Bull., 62:107-127.
Stone, R. H.

1950. Roosting brown creepers, Certhia familiaris. Auk, 67:391.
Stuew^er, F. W.

1943. Raccoons, their habits and management in Michigan. Ecol.
Monogr., 13:203-258, 16 tables, 55 figs, in text.

TORDOFF, H. B.

1956. Check-list of the birds of Kansas. Univ. Kansas Publ. Mus. Nat.
Hist., 8:307-359.
Trautman, M. B.

1940. The birds of Buckeye Lake, Ohio. Misc. Publ. Mus. Zool. Univ.
Michigan, 44:466 pp., 15 pis.
Whitney, L. F., and Underwood, A. B.

1952. The raccoon. Practical Science Publ. Co., Orange, Connecticut.
177 pp.

Wing, L.

1946. Species association in winter groups. Auk, 63:508-511.
Young, S. P., and Jackson, H. H. T.

1951. The clever coyote. Stackpole, 411 pp.

Transmitted March 11, 1958 with minor additions on September 22, 1958.

D

27-3117

(Continued from inside of front cover)

15. The pigmy woodrat, Neotoma goldmani, its distribution and systematic posi-
tion. By Dennis G. Rainey and Rollin H. Baker. Pp. 619-624, 2 figs, in
text. June 10, 1955.

Index. Pp. 625-651,
Vol. 8. 1. Life history and ecology of the five-lined skink, Eumeces fasciatus. By Henry
S. Fitch. Pp. 1-156, 26 figs, in text. September 1, 1954.

2. Myology and serology of the Avian Family FringiUidae, a taxonomic study.
By William B. Stallcup. Pp. 157-211, 23 figures in text, 4 tables. November
15, 1954.

3. An ecological study of the collared lizard (Crotaphytus coUaris). By Henry
S. Fitch. Pp. 213-274, 10 figures in text. February 10, 1956.

4. A field study of the Kansas ant-eating frog, Gastrophryne olivacea. By Henry
S. Fitch. Pp. 275-306, 9 figures in text. February 10, 1956.

5. Check-list of the birds of Kansas. By Harrison B. Tordoff. Pp. 307-359, 1
figure in text. March 10, 1956.

6. A population studv of the prairie vole (Microtus ochrogaster) in northeastern
Kansas. By Edwin P. Martin. Pp. 361-416, 19 figures in text. AprU 2,
1956.

7. Temperature responses in free-living amphibians and reptiles of northeastern
Kansas. By Henry S. Fitch. Pp. 417-476, 10 figmres in text, 6 tables. June
1, 1956.

8. Food of the crow, Corvus brachyrhynchos Brehm, in south-central Kansas. By
Dwight Piatt. Pp. 477-498, 4 tables. June 8, 1956.

9 Ecological observations on the woodrat, Neotoma floridana. By Henry S.
Fitch and Dennis G. Rainey. Pp. 499-533, 3 figures in text. June 12, 1956.
10. Eastern woodrat, Neotoma floridana: Life history and ecology. By Dennis G.
Rainey. Pp. 535-646, 12 plates, 13 figures in text. August 15, 1956.

Index. Pp. 647-675.
Vol. 9. 1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 fig-
ures in text. December 10, 1955.

2. Additional records and extensions of ranges of mammals from Utah. By
Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80.
December 10, 1955.

3. A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rol-
lin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.

4. Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming.
By Sydney Anderson. Pp. 85-104, 2 figiures in text. May 10, 1956.

5. The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6
figures in text. May 19, 1956.

6. Additional remains of the multituberculate genus Eucosmodon. By Robert W.
Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.

7. Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures
in text. Jime 15, 1956.

8. Comments on the taxonomic status of Apodemus peninsulae, with description
of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 337-346, 1
figure in text, 1 table. August 15, 1956.

9. Extensions of known ranges of Mexican bats. By Sydney Anderson, Pp. 347-
351. August 15, 1956.

10. A new bat (Genus Leptonycteris ) from Coahuila. By Howard J. Stains.
Pp. 353-356. January 21, 1957.

11. A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico.
By Robert J. RusseU. Pp. 357-361. January 21, 1957.

12. Geographic variation in the pocket gopher, Thomomys bottae, in Colorado.
By Phillip M. Youngman. Pp. 363-384, 7 figs, in text, 1 table. February
21, 1958.

13. A new bog lemming (genus Synaptomys) from Nebraska. By J. Knox
Jones, Jr. Pp. 385-388.

More numbers will appear in volume 9.
VoL 10. 1. Studies of birds killed in nocturnal migration. By Harrison B. Tordoff and
Robert M. Mengel. Pp. 1-44, 6 figvues in text, 2 tables. September 12, 1956.

2. Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. WooHenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.

3. The forest habitat of the University of Kansas Natural History Reservation.
By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures
in text, 4 tables. December 31, 1956.

(Continued on outside of back cover)

(Continued from inside of back cover)

4. Aspects of reproduction and development in the prairie vole (Microtua ochro-
gaster). By Henry S. Fitch. Pp. 129-161, 8 figures in text. 4 t.ibles. Decem-
ber 19. 1957.

5. Birds found on the Arctic Slope of northern Alaska. By James W. Bee. Pp.
163-211. 2 plates. 1 figure in text. March 12. 1958.

6. The wood rats of Colorado: distribution and ecology. By Robert B. Finley,
Jr. Pp. 213-552, 34 plates, 8 figiu-es in text, 35 tables. November 7, 1958.
More numbers will appear in volume 10.

Vol. 11. 1. TTie systematic status of the colubrid snake, Leptodeira discolor Giinther. By
William E. Ducllman. Pp. 1-9, 4 figures. July 14, 1958.

2. Natural history of the six-lined racerunner. ( Cnemidophorus sexlincatus). By
Henry S. Fitch. Pp. 11-62, 9 figures, 9 tables. September 19. 1958.

3. Home ranges, territories, and seasonal movements of vertebrates of the Natural
History Reservation. By Henry S. Fitch. Pp. 63-326, 6 pis., 24 figures in
text, 3 tables. December 12, 1958.

More numbers wUl appear in volume 11.

, • ; •" U C =^ w «^ €^ irx c e.3 ■

IVlUd.

1.^^

FEB 19i;J5l

\mtMn

University of Kansas PublicationI
Museum of Natural History

Volume 11, No. 4, pp. 327-334, 2 figs.
January 28, 1959

A New Snake of the Genus Geophis
From Chihuahua, Mexico

BY

JOHN M. LEGLER

University of Kansas

Lawrence

1959

University of Kansas Publications, Musexjm of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 11, No. 4, pp. 327-334
Published January 28, 1959

University of Kansas
Lawrence, Kansas

FEB 1 9 1059

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA. KANSAS

I 9S9

27-6682

A New Snake of the Genus Geophis
From Chihuahua, Mexico

BY
JOHN M. LEGLER

In July, 1957, members of a field party from the University of
Kansas Museum of Natural History, under the direction of Mr.
Sydney Anderson, spent 12 days collecting vertebrates in the vi-
cinity of Creel in southwestern Chihuahua. Among the specimens
are two snakes representing an undescribed species of the genus
Geophis. A description and illustrations of these two specimens
were prepared and submitted for publication in the spring of 1958.
At that time it came to my attention that Dr. Norman Hartweg,
Museum of Zoology, University of Michigan, was also preparing
a report on four specimens of the same species from two additional
localities in southwestern Chihuahua. Upon learning of my work
on the species, Dr. Hartweg generously loaned me his specimens
and notes and allowed me to incorporate them in the present report.
The snakes may be known and described as:

Geophis aquilonaris new species

Holotype. — Adult female, KU 44265, alcoholic; 23 mi. S and VA mi. E
Creel, Chihuahua, Mexico; 23 July 1957; original number 198, Ronald H. Pine.

Paratypes. — (Total of five alcoholic specimens, all from Chihuahua) Male
(probably subadult), KU 44266, same data as holotype; male, UMMZ 117770,
Mojarachic, 23 July 1957, Irving W. Knobloch; females, UMMZ 111501-2,
Maguarachic, August 1954, and UMMZ 117771, Maguarachic, August 1957,
Irving W. Knobloch.

Diagnosis. — Size small; 15 rows of smooth scales; a high number of ventral
(173 to 183) and subcaudal (55 to 64) scales; alternating dark and pale rings
on body and tail; dark rings, and often pale rings, complete.

Description of Holotype. — Snout-vent length 327 mm.; length of tail 93
mm.; anal scale entire; ventral scales 181 + anal; subcaudal scales 63 -4- tip;
dorsal scales in 15 rows on all parts of body; six supralabials (fifth and sixth
fused on left side), third and fourth entering orbit; fifth supralabial largest
and in broad contact with parietal, posterior temporal, and postocular; six
infralabials on each side, first pair in contact behind mental; enlarged chin
shields in two pairs, anterior pair longer than posterior pair; anterior chin
shields in contact for half their length with fourth infralabials; rostral nearly
as high as broad; intemasal and prefrontal scales paired and distinct; anterior
and posterior segments of nasals distinct and nearly equal in size; loreal twice
as long as high, in contact with eye; preocular lacking (represented by minute
scale on left side); vertical diameter of eye equal to distance from lower rim

(329)

330

University of Kansas Publs., Mus. Nat. Hist.

of orbit to free edge of upper lip; temporal formula 0+1, the single temporal
scale separating sixth supralabial and parietal; one postocular and one supra-

FiG. 1. Geophis aquilonaris new species, KU 44265, Holotype; lateral, dorsal,
and ventral views of head and neck (approximately X 6).

ocular on each side; all scales perfectly smooth; no scale-pits evident; dentary
bone bearing eight teeth; maxillary bone bearing seven teeth; posterior tooth
longest, thirmest, and separated from other teeth by slight diastema (maxillary

A New Snake of the Genus Geophis

331

teeth in UMMZ 111502, 6/7, dentary teeth 8/8, no diastema in maxillary
series).

Head slate-black above, having dim pale mark on anterior part of each
prefrontal and another on interparietal seam; an indistinct pale gray crescent
on posterior border of nostril; narrow cream band covering posterior edges
of parietal and half of first dorsal scale row, widening laterally to include

rvc

Fig. 2. Geophis aquilonaris new species, KU 44265, HoIot3^e; scalation
and coloration at mid-body showing 19th and 20th white rings (approx-
imately X 7).

temporal and posterior two or three supralabials; throat cream (except for
dark markings on mental, on first three infralabials, and on anterior chin
shields), its pale area continuous with pale band on head; body and tail
marked with alternating white and black rings; white rings (excluding band
on head) 38 on body, 17 on tail; each white ring alternately one and two
scales wide dorsally (producing wavy or zigzag e£Fect), widened laterally, and
three to four scales wide on belly; black rings three to four and one half

332

Unr'ersity of Kansas Publs., Mus. Nat. Hist.

scales wide on middorsal line, and two to three scales wide on belly; black and
white rings (excepting first black ring behind head) continuous around body
and tail. Colors described above nearly same as in living specimens.

Range. — The species is known only from three locahties on the Pacific
drainage of southwestern Chihuahua; the geographic range probably includes
parts of southern Sonora and northern Sinaloa. The discovery of Geophis in
southern Chihuahua increases to 21 the number of species of the genus known
to occur in Mexico and extends the known range of the genus approximately
560 miles northwestward from the type locality of G. latifrontalis, a point 50
miles south (in Guanajuato?) of San Luis Potosi, or, a slightly lesser number
of miles north-northwestward from an indefinite locality for G. bicolor in
western Jahsco ( La Cumbre de los Arrastrados ) ( Boulenger, Catalogue of
the snakes in the British Museum, Vol. 2, 1894, p. 298).

Variation. — Standard counts of scales are given for the paratypes as well
as the holotype in table 1. The fifth and sixth supralabial scales are fused
on both sides of one specimen (UMMZ 117771) as is the case on the left
side of the holotype. Except for one specimen (UMMZ 117770) that has a
small anterior temporal separating the posterior two-thirds of the fifth supra-
labial from the parietal on each side, the temporal formula in the type series
is uniformly 0 -f 1.

Table 1. — Counts of Scales, Measurements, and Other Data Pertaining
TO Holotype and Paratypes of Geophis aquilonaris New Species.

Sex

Dorsal
scale
rows

Ven-
trals

Sub-
caudals

Pale

rings

Snout-
vent
length

Length

of

tail

Body

Tail

UMMZ

111501

9

15

173

55

29

12

237

60

UMMZ

111502

9

15

181

58

34

14

355

88

UMMZ

117771

9

15

182

55

39

16

371

90

KU

44265

9

15

183

63

38

17

327

93

KU

442G6

cT

15

175

60

30

13

160

43

UMMZ

117770

cf

15

174

64

34

15

245

74

Considerably more variation occurs in color and in the arrangement of
markings than in squamation. The ground color of the two specimens from
Creel is black with little or no trace of brown, and the rings are white. Ground
color in the remaining paratypes ranges from grayish black, with some brown-
ishness on the belly, to dark brown, the colors in one specimen approximating
the range from Mummy brown to Dresden brown, becoming paler posteriorly
and ventrally. The head is slate gray to blackish brown in all the specimens.
Those having a suggestion of brown on the head tend also to have more brown
on the body.

The dark band on the neck is complete in four of the paratypes and in-

A New Snake of the Genus Geophis 333

complete (as in holotype) in one. Pale marks on the prefrontals are lacking
in three of the paratypes and the pale mark between the parietals is lacking
in two specimens (fused with white band on neck of one specimen). Pale
postnarial crescents are evident in three paratypes.

As stated above, the rings on the holotype are mostly complete. Excep-
tions occur between the 13th and 15th white rings where two black rings are
fused on the left side, rendering one black and one white ring (the 14th)
incomplete. Also, where the ninth and tenth white rings fuse on the left
side, they enclose a black ring and render it incomplete. The markings of the
three smaller paratypes are substantially the same as those of the holotype —
complete rings with a small number of variations in each specimen. In the
two largest paratypes nearly all the white rings are incomplete ventrally, ap-
pearing to have been encroached upon by the darker ground color. In the
larger speicmens there is a tendency also for the white rings to be one scale
wide (rather than alternately one and two scales wide) and to lack a zigzag
appearance; this appears to be due to the darkening of entire scales and to
the darkening of the edges of other scales.

Relationships. — Geophis aquilonaris is distinct from all other
Mexican representatives of the genus in having, on the body and
tail, numerous, alternating pale and dark bands. Both sets of bands
are in the form of complete rings or the dark bands are joined ven-
trally rendering the belly dark.

Of the seven other Mexican Geophis having 15 rows of scales,
four species (cancellatus, dugesii, chalyheus, and semidoliatus)
have alternating pale and dark transverse markings and therefore
superficially resemble aquilonaris. Of the latter two species, the
poorly known G. chalyheus (Veracruz) has a much lower (137 to
142) number of ventrals than aquilonaris, and G. semidoliatus
(southeastern Mexico — Veracruz, Hidalgo, and Oaxaca) has a
narrower head, fewer supralabials (four to five with only the third
entering the orbit), and fewer ventrals (136 to 169) than aquilo-
naris. Geophis aquilonaris seems to be most closely allied to G.
cancellatus (Chicharras, Chiapas) and G, dugesii (known from
two localities in northern Michoacan); all three species resemble
one another in the number and arrangement of the scales of the
head, in general coloration, and in having relatively high numbers
of ventral scales (171 in cancellatus, 150 to 164 in dugesii). G.
cancellatus differs from the other two species in lacking internasal
scales. Geophis aquilonaris differs from both species in having a
higher number of ventral and subcaudal scales, a longer tail (tail
contained in snout-vent length three to four times in aquilonaris,
four and one half to six times in dugesii, 11 to 12 times in cancel-
latus), and in having more bands on the body (28 to 32 in cancel-

334 University of Kansas Publs., Mus. Nat. Hist.

latus, two to seven in dugem ) . The belly in dugesii and cancellatus
is pale but in aquilonaris it is ringed or of a solid dark color.

As more specimens of Ceophis become available from intermedi-
ate localities in Mexico, it will perhaps be demonstrated that many
of the kinds now thought of as full species (including those dis-
cussed above) are subspecies of a few wide-ranging species.

Remarks. — The type locality of G. aquilonaris is the small village
of Barranca at the bottom of the valley of the Rio Urique, several
miles south and west of the continental divide. The Urique Valley,
known as the Barranca del Cobre in the region south of Creel, is
a deep canyon, the walls of which slope abruptly from approxi-
mately 7300 to 3000 feet and are dissected by deep side-canyons.
Coniferous forest on the upper rim of the canyon is replaced by
scrub vegetation on the rocky walls and by an arid tropical flora
on the bottom.

Maguarachic (elevation approximately 5400 feet, longitude and
latitude respectively, 108 degrees, 03 minutes W and 27 degrees,
50 minutes N ) and Mojarachic ( elevation approximately 7000 feet,
longitude and latitude respectively, 108 degrees W and 27 degrees,
52 minutes N) are situated approximately three miles from each
other and approximately 27 miles northwest of Creel. Maguarachic
is given as "Mafuarachic" on the American Geographical Society
map (NG 12, Baja California-Mexico, Prov. Ed., 1924). Mojara-
chic is not on any map of Chihuahua that I have examined.

The type and topotypic paratype were given to a member of the
K. U. field party by a Mexican youth who had obtained them the
previous night on the lower rocky slopes of the canyon. Both
specimens were damaged by the collector piercing their heads with
thorns, presumably to kill them. The type contained three oviducal
eggs, each about four millimeters long. The stomachs of both speci-
mens from Creel contained earthworms.

The presence of Geophis in this area suggests that the distribu-
tion of the genus is more or less continuous, on tlie western slope
of the Sierra Madre Occidental, from Jalisco to southern Sonora.

I am grateful to Mr. Sydney Anderson and Mr. Ronald Pine for permission
to use their field notes, to Dr. Hobart M. Smith for his examination of the
specimens from Creel, to Mrs. Loma Cordonnier for the drawings of the type,
to Dr. Norman Hartweg for permitting me to study materials in his care and
upon which he was making an independent study, and to Mr. Thomas M.
Uzzell for locaUty data pertaining to the UMMZ paratypes.

Transmitted November 10, 1958.

D

27-6682

-DJI^-L

MUS. COM?. ZOOL
JUN181959

University of Kansas PublicaticIjms.

Museum of Natural History

yiliVERSlTY

Volume 11, No. 5, pp. 335-343, pis. 7 and 8
April 24, 1959

A New Tortoise, Genus Gopherus,
From North-central Mexico

BY

JOHN M. LEGLER

University of Kansas
Lawrence
"=^1959

University of Kansas Publications, Museum of Natural History

Editors: E. Ra>aiiond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 11, No. 5, pp. 335-343
Published April 24, 1959

University of Kansas
Lawrence, Kansas

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA. KANSAS

t9B9

27-6858

A New Tortoise, Genus Gopheruf,
From North-central Mexico

'ilil COMP. ZOOL
JUN181959

BY
JOHN M. LEGLER

In the course of taxonomic studies of the genus Gopherus it was
found that specimens from north-central Mexico belong to an
hitherto unrecognized species, which is named and described be-
low.

Gopherus flavomarginatus new species

Holotype. — U. S. National Museum 61253, adult; stufiFed specimen with dis-
associated skull; 30 to 40 miles from Lerdo, Durango, Mexico; obtained by Dr.
Elswood Chaffee, 1918.

Paratypes. — USNM 61254, adult, stuffed specimen with skull in place,
other data the same as those for holotype; USNM 60976, adult, stuffed speci-
men with disassociated skull, Lerdo, Durango, Dr. Elswood Chaffee, May
1918; University of Illinois 42953-4, adults, two carapaces, Carrillo, Chihuahua,
Pete S. Chrapliwy and Kermeth L. WiUiams, 1 August 1958; University of
Kansas 39415, adult, carapace only, 8 mi. E and 2 mi. S Americanos, Coahuila,
Robert W. Dickerman, 19 May 1954.

Diagnosis. — A large tortoise of the genus Gopherus, having a flat-topped,
posteriorly-flared shell, carapacal laminae that are pale with contrasting dark
centers (at least on the lateral margins of the carapace), intergular seam
longer than interhmneral, obtuse gular projections, and a relatively wider skull
than other Mexican representatives of the genus.

Description of species. — Ground color of carapace pale yellow to straw in
smaller specimens, pale brown in larger specimens; in some specimens a slight
indication of darker radial markings on carapace; epidermal areolae of cara-
pace (if present) and first two or three zones of growth (around areolae) dark
brown to black, their contrast vdth the ground color greater in smaller speci-
mens; ground color of lateral marginals yellowish, much paler than remainder
of carapace; black areolae of marginals contrasting sharply with ground color
even in old individuals; plastron pale, approximately the same shade of yellow
as lateral marginals, marked with sharply contrasting blotches of black or dark
brown in younger specimens, becoming immaculate with age.

Carapace low, evenly arched or shghtly flat-topped in cross section, its
height shghtly more than 50 per cent of its length; top of carapace, between
highest parts of first and fourth central laminae, more or less flat in profile;
carapace flared, wider posteriorly than anteriorly; central laminae all broader
than long, the first not (or but shghtly) narrower posteriorly than anteriorly;
plastron deeply notched behind, the inner margins of the notch straight,
neither convex nor concave; gular projections truncate (not prongUke) having
no notch or but a shallow notch between them; greatest combined width of gular
laminae much greater than greatest length; interlaminal length of gular greater

(337)

338

University of Kansas Publs., Mus. Nat. Hist.

than that of humeral; one large axillary scute on each side, approximately rec-
tangular but slightly wider below than above; one or two inguinal scutes on
each side, the anterior roughly triangular and much larger than posterior.

Anterior surface of antebrachium having enlarged, juxtaposed (or shghtly
imbricated), osteodermal scales arranged in seven to eight longitudinal rows,
each scale yellowish with a black or dark bro^^^a center; scales on lateral edge
of antebrachium, from fifth claw to elbow, ten in number and darker than other
antebrachial scales; two enlarged black-tipped, yellow spurs on posterior sur-
face of thigh, inner spur approximately half the size of outer. Head relatively
wide, greatest vi^dth of skull shghtly more than 80 per cent of condylobasilar
length; width of head 1.3 to 1.7 times diameter of hind foot; scales on top of
head large but irregularly arranged (see table 1 for measurements of type and
paratyTpes ) .

Table 1. Measurements, in Millimeters, of the Holotype and Paratypes
OF Gopherus Flavomarginatus New Species

4-1

o

?,

(U

a

^

Catalogue Number

1

i

1

J

13

q
3

13

AND Collection

u

Qi

o

M

•*4

Mh

o

^

O

^

•t

a

1

a

H

13

^

h-1

0)

Q

^

USNM 60976

246

212

252

108

31

41

USNM 61253

222

166

212

96

21

36

USNM 61254

371

292

358

157

43

55

lU 42953

281

220

lU 42954

278

214

KU 39415

303

232

Duges (1888:146-147, and 1896:479) twice reported specimens of Gopherus
polyphemus from Chihuahua ( see discussion of distribution ) ; his description of
two living examples (1888) agrees generally with the above description of G.
■flavomarginatus. He stated that the iris of his smaller specimen was yellow
and the iris of a larger specimen brown.

It will be most interesting to leam the color of juveniles of G. flavomar-
giriatus. Considering the dark color of the epidermal areolae of adults, I
would expect the color of hatchhngs to be dark brown to nearly black, with
each lamina acquiring paler borders as it grew, rather than acquiring darker
borders as is usually the case in G. berlandieri (personal observation), agassizii
(MiUer, 1932:194, pi. 2; Woodbury and Hardy, 1948:165, figs. 9 and 10), and
polyphemus (personal observation).

PLATE 7

Plate 7. Gopherus fiavumarginatus new species: Top — Dorsal and ventral
views of liolotype (x /»); Middle — Lateral view of holotype (X/s); Bot-
tom— Head of paratype (USNM 61254) (XJO and antebrachium of
liolotype (X %)• Dark outline on plastron of holotype indicates where
portion of shell was removed (and then replaced) in course of preparation.

PLATE S

Pi.AiK 8. Coiihcms fi(Hnin(ii^i)uitiis ii(V\ specie's: l)i)isal xicws of

paratypes ( Jroiii top to bottom, W 42953, 11' 42954, and KU 39415),

approximately i,-, natural si/e.

A New Tortoise, Genus Gopherus

339

Fig. 1. Gopherus flavamarginatus new species: la and lb — Lateral
and ventral views of skull of holotype ( x 1 ) ; Ic — Cross section of
paratype (USNM 60976) taken through middle of third central

lamina ( X %) .

Anomalies. — There is a high incidence of anomaly in the type series. Three
of the specimens (USNM 60976, lU 42954, and KU 39415) have a super-
mmierary central lamina. All of the specimens from Durango are anomalous
in regard to the marginal laminae. In the holotype the first marginals are
small, approximately as wide as the small precentral. In the largest para-
type (USNM 61254) the first and second marginals are fused to form a single
large scute on each side. In the remaining Durangan paratype (USNM 60976)
there are eleven normal marginals on each side but a supernumerary lamina
occurs between the first and second marginal on the left and between the
precentral and first marginal on the right side. The remaining three paratypes
have the number of marginals that is normal for the genus.

Relationships. — Gopherus flavomarginatus differs from all other species of
the genus by its larger size and by having an intergular seam that is longer
than the interhumeral seam. The possession of a single, rectangular axillary
scale, wider below than above, also tends to distinguish favomarginatus from
other living Gopherus; in the other species this scute (frequently paired in
herlandieri) is generally triangular or at least pointed below. Femoral spurs
are developed to varying degrees in the other species of Gopherus; they are

340

Untversity of Kansas Publs., Mus. Nat. E[ist.

usually lacking in indi\iduals of small to medium size. In large specimens of
agassizii and pohjphemus that I have examined, there is but one enlarged spur,
although several pointed osteodermal scales may be present on the back of the
thigh.

Gopherus herlandieri, the closest relative of G. jlavomarginatus in a geo-
graphic sense, has a highly arched shell and pronglike gular projections, be-
tween which there is nearly always a deep notch. Gopherus agassizii, morpho-
logically the closest Mexican relative of jlavomarginatus, differs from it in hav-
ing a narrower head and in lacking the contrasting pale and dark coloration of
the plastron and lateral marginal area. Gopherus polyphemus seems to be the
closest relative of jlavomarginatus and differs from it in having an unflared
shell (anterior and posterior wddths of carapace subequal or carapace narrower
behind) and in the characteristics of color mentioned above. Certain propor-
tional differences between the living species of Gopherus are summarized in
table 2.

Distribution. — The discovery of a new Gopherus from north-central Mexico
increases to four the recognized forms of the genus and extends the known
range of the genus onto the Mexican highlands, approximately 410 miles east-
ward from Alamos, Sonora (G. agassizii), approximately 100 miles westward
from a point 2 mi. W of Monclova, Coahuila (G. herlandieri), and approxi-
mately 200 miles westward from Monterrey, Nuevo Le6n (G. herlandieri).

Table 2. Bodily Pboportions in the Knowtsi Species of Gopherus. Two
Specimens of G. polyphemus from Harmon, Mississippi (USNM
53166-7), Are Considered Separately Because of Their Notably Nar-
rower Heads. The Size of Each Sample Is Given in Parentheses Above
the Average; Extremes Abe in Parentheses Followtng the Averages.

Species

and

General Locality

Greatest
width of
skull as a
percentage
of condy-
lobasilar
length

Height

of

shell as a

percentage

of width

of
carapace

Height

of

shell as a

percentage

of length

of
carapace

Diameter

of
hind foot

as a

percentage

of width

of head

G. polyphemus
(Florida)

(6)

.94 (.92-.97)

(13)

.53 (.48-.58)

(13)

.39 (.36-.42)

G. polyphemus
(Mississippi)

(2)

.87 (.84-.89)

(2)

.53 (.52-.54)

(2)

.43 (.40-.45)

G. herlandieri

(Texas and NE Mexico)

(21)

.78 (.70-.83)

(22)

.56 (.48-.62)

(21)

.46 (.43-.51)

G. agassizii

(U.S. and N. Sonora)

(6)

.77 (.70-.83)

(8)

.53 (.49-.58)

(8)

.41 (.38-.44)

G. agassizii

(Alamos, Sonora)

(4)

.56 (.52-.59)

(4)

.39 (.37-.41)

(4)
.92(.87-1.00)

G. Jlavomarginatus
(North-central Mexico)

(2)

.83 (.82-.83)

(3)

.54 (.61-.68)

(3)

.43 (.42-.44)

(3)

.71 (.58-.78)

A New Tortoise, Genus Gopherus 341

Thus far, G. flavomarginatus is known only from the three localities repre-
sented in the type series and from the indefinite locality, "Bolson de Mapimi",
Chihuahua, given by Duges ( 1888:146-7, and 1896:479). These long-doubted
references of Duges to Gopherus polyphemus seem clearly to represent G. jlavo-
marginatus, which probably occurs in several or all of the internally drained
basins in northeastern Chihuahua, western Coahuila, and northern Durango.
This region is within the Basin and Range physiographic province of Fenne-
man (1931:326-8) and the Mapimi biotic province of Smith (1949:231).
Duges ( supra cit. ) seems to have used "Bolson de Mapimi" in a restricted sense,
as it is shown on some recent maps ( Nat. Geog. Soc, Map of Mexico and Cen-
tral America, 1953) (American Geog. Soc, Cuhcan Map, NG 13, 1935).
Other maps show this bolson to include internally drained portions of the
Mexican highlands from northeastern Chihuahua to the region near Lerdo and
Torreon. Thayer (1916:73) pointed out that the Bolson de Mapimi, in its
larger sense, consists of a series of basins separated by mountains of con-
siderable elevation.

The range of G. flavomarginatus appears to be limited ultimately by the
higher elevations of the Sierra Madre Occidental and the Sierra Madre Oriental
to the west and east, respectively, and to the south by the Mexican plateau.
The northern limits of the range are less clear; possibly the range extends as far
as the portions of the United States adjacent to northeastern Chihuahua, but
this is doubtful. The range seems not to overlap that of any other species of
Gopherus, although the ranges of G. flavomarginatus and G. berlandieri closely
approximate each other in central Coahuila.

In September, 1958, when I was collecting turtles near Cuatro Cienegas,
Coahuila, I took the opportunity to query natives, as well as an American
rancher, about the possible occurrence of tortoises in the area. Most persons
had seen no tortoises in the area or said they had seen them only rarely. Sev-
eral older men who had herded goats in the area all their lives said that tor-
toises (referred to as "Tortuga del Monte") were common on the other side
of the Sierra de La Madera and Sierra de La Fragua ranges west of Cuatro
Cienegas. These men referred probably to G. -flavomarginatus. Americanos
lies approximately 75 miles west-northwest of Cuatro Cienegas.

It is indeed remarkable that a population of large tortoises in northern
Mexico has so long escaped the notice of naturalists. Also remarkable is the
fact that the late Dr. Leonhard Stejneger, in view of his intense interest in North
American chelonians and his familiarity with the genus Gopherus, did not re-
mark on the specimens from Durango when he saw them; a thorough search of
Stejneger 's notes revealed no clue that he had ever studied the specimens.
Perhaps his first reaction to the specimens, like mine when I first examined the
two disassociated skulls, was to consider "Lerdo" a misspelling of "Laredo."
However, a check of old correspondence and accession records at the National
Museimi confirmed, beyond doubt, the validity of the locahty data for these
specimens. Dr. ChaflFee, the collector, frequently sent specimens of plants
and animals to the Museum from Diurango and Zacatecas; his home was in
Lerdo.

Size. — The largest specimen of Gopherus flavomarginatus (USNM 61254)
exceeds by more than 50 millimeters the length of the largest knovra speci-
mens of G. agassizii, and slightly exceeds the length of the largest specimen

342 Unr'ersity of Kansas Publs., Mus. Nat. Hist.

of G. polyphemus that I have examined (USNM 51357, length of carapace
360 millimeters, a specimen formerly kept at the National Zoological Park). G.
polyphemus is the largest tortoise inhabiting the United States; the maximum
length of 343 millimeters given for the species by Carr (1952:334) probably
more closely approaches the true maximum in free-living populations. G.
agassizii rarely attains a length greater than 300 millimeters (Woodbury and
Hardy, 1948:152-5). The fact that representatives of the genus do not attain
a larger size in the United States may be due to the decimation of natural popu-
lations by man for food and souvenirs.

There have been several indications that a large species of tortoise existed
in north-central Mexico. The Chihuahuan specimens reported by Duges (248
and 202 miUimeters long, respectively) rank in size with the smaller paratypes
of G. flavomarginatus; Duges stated, however, that the species was said to attain
a length of one meter but that he thought such large size surely to apply to
another species.

Mr. Charles M. Bogert recently related to me two stories that are here
worthy of note. One of them, which Bogert learned from the late Dr. Karl
P. Schmidt, concerns a collector who was sent by the Chicago Natural History
Museum (then Field Museum) to Coahuila in the early part of the century to
obtain materials for a desert exhibit. When the exhibit had been completed
several decades later, the collector, visiting the museum, inquired as to where
the large tortoises were that he had collected in Coahuila. Dr. Schmidt could
find no record of such tortoises and suggested that possibly they had been
confused with Galapagos tortoises (Geochelone). Dr. Robert F. Inger is unable
to locate the specimens or add anything to the story. The collector stated that
the large tortoises had been found in only one bolson in Coahuila.

Mr. Bogert further related that, in 1946 when he was camped at Las
Delicias, Coahuila, a Mexican mule herder told him of seeing a large tortoise
"three bolsons north" of the bolson in which Las Dehcias is located (the gen-
eral area referred to would be near Americanos ) . The man awoke one morn-
ing to find his saddle missing; following tracks from the place where the saddle
had been left, the man eventually found it on the back of a large tortoise.
The tortoise had apparently sought shelter beneath the saddle and, finding
it unsatisfactory, had walked away with the saddle on his back.

Either of the incidents related above, or the reference of Duges (loc. cit.)
to large size, since they are based on second or third-hand accounts, would
seem to be unreahstic and of Uttle scientific value when considered alone and
at face value. However, in the fight of the discovery of Gopherus fiavonwr-
ginatus, these accounts have new meaning and tend to support my proposal
that the species is the largest tortoise in North America.

Remarks. — The paratypes at the University of Illinois were found in a trash-
dump in Carrillo. According to what the collectors learned from local inhabit-
ants, the species is used for food and the shells are sometimes used for poultry
dishes or even for eating utensils. Although the species is found on the flat,
sandy desert near Carrillo, it is more common on the nearby mountain slopes
and is seen there most frequently after rains. Dickerman (field notes, 1954)
likewise noted that the species was eaten near Americanos; the K. U. para-
type represents the best (and only specimen saved) of several broken shells
found in that area. Judging by the habits of other members of the genus and
by the notes of Duges (1888:147), G. ilavomarginatus is probably herbivorous.

A New Tortoise, Genus Gopherus 343

Acknowledgments. — A portion of this study was completed in the summer
of 1958, while I was an internee at the United States National Museum. I am
grateful to Dr. Doris M. Cochran of that institution for helpful information re-
garding the origin of the type and for permission to study other specimens
in her care. Thanks are due also to Messrs. Pete S. ChrapUwy and Kenneth
L. Williams who collected two of the paratypes and who generously provided
their notes on the specimens when learning of my study, to Dr. Hobart M.
Smith and Mr. Charles M. Bogert for offering helpful suggestions and for the
loan of specimens, and to Mr. Jebb Taylor for assistance with taking data.
The drawings are the work of Mrs. Connie Spitz; the photographs are by the
author.

LITERATURE CITED
Cabr, a.

1952. Handbook of tiuiles; the turtles of the United States, Canada, and
Baja California. Cornell Univ. Press, xv -f- 542 pp., 37 figs., 82 pis.,
15 tables, 23 maps.
DuGES, A.

1888. La Tortuga Polifemo. La Naturaleza, l(ser. 2): 146-147.
1896. Reptiles y Batracios de los E. U. Mexicanos. La Natxiraleza,
2(ser. 2):479-485.
Fenneman, N. M.

1931. Physiography of western United States. McGraw-Hill, New York,
1st. ed„ v-xii -f 534 pp., 173 figs.

Miller, L.

1932. Notes on the desert tortoise {Testudo agassizii). Trans. San Diego
Soc. Nat. Hist., 7:187-208.

Smith, H. M.

1949. Herpetogeny in Mexico and Guatemala. Ann. Assn. American
Geographers, 34(3):219-238, 1 fig.
Thayer, W. N.

1916. The physiography of Mexico. Journ. Geol., 24:61-94, 2 figs.
Woodbury, A. M., and Hardy, R.

1948. Studies of the desert tortoise, Gopherus agassizii. Ecol. Monogr,,
18:145-200, 25 figs., 4 tables.

Transmitted November 18, 1958.

n

27-6856

in J^ L-

H!!S. COM?. ZCOL

JUN181959

University of Kansas Publication*.

Museum of Natural History

Volume 11, No. 6, pp. 345-400, 2 plates, 2 figs, in text, 10 tables
May 6, 1959 -—.

Fishes of

Chautauqua, Cowley and

Elk Counties, Kansas

BY

ARTIE L. METCALF

University of Kansas

Lawrence

T959

UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY

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ming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text.
July 23, 1954.

8. A new subspecies of bat (Myotis velifer) from southeastern California and
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9. Mammals of the San Gabriel mountains of California. By Terry A. Vaughan.
Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954.

10. A new bat (Genus Pipistrellus) from northeastern Mexico. By Rollin H.
Baker. Pp. 583-586. November 15, 1954.

11. A new subspecies of pocket mouse from Kansas. By E. Ra>Tnond Hall. Pp.
587-590. November 15, 1954.

12. Geographic variation in the pocket gopher, Cratogcomvs castanops, in Coa-
huila, Mexico. By Robert J. Russell and Rollin H. Baker. Pp. 591-608.
March 15, 1955.

13. A new cottontail (Sylvilagus florfdanus) from northeastern Mexico. By Rollin
H. Baker. Pp. 609-612. April 8, 1955.

14. Taxonomy and distribution of some American shrews. By James S. Findley.
Pp. 613-618. June 10, 1955.

15. The pigmy woodrat, Neotoma goldmani, its distribution and systematic posi-
tion. By Dnnnis G. Rainey and Rollin H. Baker. Pp. 619-624, 2 figures in
text. June 10, 1955.

Index. Pp. 625-651.

(Continued on inside of back cover)

University of Kansas Publications
Museum of Natural History

Volume 11, No. 6, pp. 345-400, 2 plates, 2 figs, in text, 10 tables
May 6, 1959

Fishes of

Chautauqua, Cowley and

Elk Counties, Kansas

BY

ARTIE L. METGALF

University of Kansas

Lawrence

1959

Unfv'ersity of Kansas Publications, Musexim of Natural History

Editors: E. Raymond Hall, Chainnan, Henry S. Fitch,
Robert W. Wilson

Volume 11, No. 6, pp. 345-400, 2 plates, 2 figs, in text, 10 tables
Published May 6, 1959

MUS. COiJ?. ZCOl

JUN181959
KARPRfl

University of Kansas
Lawrence, Kansas

A Contribution From
The State Biological Survey of Kansas

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA. KANSAS

1959

27-7079

Fishes of

Chautauqua, Cowley and

Elk Counties, Kansas

BY

ARTIE L. METCALF

CONTENTS

PAGE

Introduction 347

Physical characteristics of the streams 351

Chmate 351

Present flora 353

History 354

Conservation 357

Previous ichthyological collections 357

Acknowledgments 358

Materials and methods 358

Collecting stations 359

Annotated list of species 362

Fishes of doubtful or possible occurrence 383

Faunal comparisons of different streams 384

Distributional variations within the same stream 387

Faunas of intermittent streams 390

East-west distribution 392

Summary 394

Literature cited 397

INTRODUCTION

Aims of the distributional study here reported on concerning
the fishes of a part of the Arkansas River Basin of south-central
Kansas were as follows:

( 1 ) Ascertain what species occur in streams of the three counties.

(2) Ascertain habitat preferences for the species found.

(3) Distinguish faunal associations existing in different parts
of the same stream.

(4) Describe differences and similarities among the fish faunas
of the several streams in the area.

(5) Relate the findings to the over-all picture of east- west dis-
tribution of fishes in Kansas.

(347)

348 UNi\'ERsmL' OF Kansas Publs., Mus. Nat. Hist.

(6) List any demonstrable effects of intermittency of streams on
fish distribution within the area.

Cowley and Chautauqua counties form part of the southern
border of Kansas, and Elk County lies directly north of Chau-
tauqua. The following report concerns data only from tliose three
counties unless otherwise noted. They make up an area of 2,430
square miles having a population of 50,960 persons in 1950 ( 55,552
in 1940, and 60,375 in 1930). The most populous portion of the
area is western Cowley County where Arkansas City with 12,903
inhabitants and Winfield with 10,264 inhabitants are located. Each
of the other towns has less than 2,000 inhabitants. In the Flint
Hills, which cross the central portion of tlie area surveyed, popu-
lation is sparse and chiefly in the valleys.

Topographically, the area is divisible into three general sections:
the extensive Wellington formation and the floodplain of the
Arkansas River in western Cowley County; the Flint Hills in the
central part of the area; and the "Chautauqua Hills" in the eastern
part. The drainage pattern is shown in Figure 1.

The Wellington formation, which is devoid of sharp relief,
borders the floodplain of the Arkansas River through most of its
course in Cowley County. A short distance south of Arkansas City,
however, the Arkansas is joined by the Walnut River and enters a
narrow valley walled by steep, wooded slopes. Frye and Leonard
(1952:198) suggest that this valley was originally carved by the
Walnut River, when the Arkansas River flowed southward west
of its present course. They further suggest that during Nebraskan
glacial time the Arkansas probably was diverted to the rapidly
downcutting Walnut. The Arkansas River has a gradient of 3.0
ft. per mile in Cowley County. This gradient and others cited were
computed, by use of a cartometer, from maps made by the State
Geological Survey of Kansas and the United States Geological
Survey.

Northward along the Walnut, steep bluffs and eroded gulleys
characterize both sides of the river, especially in southern Cowley
County. Two massive limestones, the Fort Riley and the Winfield,
form the bluffs in most places. The well-defined Winfield lime-
stone is persistent on the west bank of the river across the entire
county. The Walnut has only a few small tributaries in the south-
ern half of Cowley County (Fig. 1). In the northern half, how-
ever, it is joined from the east by Timber Creek and Rock Creek.
Timber Creek drains a large level area, formed by the eroded upper

Fishes of Arkansas River Basin

349

portion of the Fort Riley limestone, in the north-central portion of
the county. The gradient of Timber Creek is 12.9 feet per mile.
The gradient of the Walnut River is only 2.3 ft. per mile from its
point of entrance into the county to its mouth.

Fig. 1. Map of Cowley, Chautauqua and Elk counties, Kansas, showing the

streams mentioned in tlie text.

Grouse Creek, like the Walnut, has formed a valley of one to
three miles in width, rimmed by prominent wooded bluflFs. Those
on the west side are capped by the Fort Riley limestone with the
resistant Wreford and Crouse limestones forming lower escarp-
ments. On the east side the Wreford and Crouse limestones pro-
vide the only escarpments along the stream above the Vinton
community, except for occasional lower outcrops of Morrill lime-
stone. Below Vinton the Fort Riley limestone again appears,
capping the hills above the Wreford limestone. The headwaters
of the western tributaries of Grouse Creek are generally in the
Doyle shale formation; the eastern tributaries are in the Wreford
Hmestone, Matfield shale, and Barnestone limestone formations.
The gradient of Grouse Creek is 9 ft. per mile, of Silver Creek 14.6
ft. per mile, and of Crab Creek 14.4 ft. per mile.

The Big Caney River ( Fig. 1 ) , having a gradient of 15.4 ft. per
mile in the area studied, drains an area with considerable geological
and topographic variation. The main stream and its western trib-

350 Unix-ersity of Kansas Publs., Mus. Nat. Hist.

utaries originate in Permian formations, whereas the eastern trib-
utaries originate in Pennsylvanian formations. Cedar Creek is
exemplary of western tributaries of Big Caney. This creek arises
in the Wreford limestone, as do several nearby tributaries of Grouse
Creek. Although the Grouse tributaries descend through only
part of the Council Grove group. Cedar Creek flows downward
through the entire Grove, Admire, and Wabaunsee groups and part
of the Shawnee Group (Moore, 1951). In only 15 miles. Cedar
Creek traverses formations comprising more than 60 per cent of
the entire exposed stratigraphic section in Cowley County. Bass
(1929:16) states that reliefs of 350 feet within a mile are present
in parts of this area.

Large terraces of limestone characterize the eastern flank of the
Flint Hills, which the western tributaries of Big Caney drain. Most
striking is the Foraker limestone. It characteristically consists of
three massive members in Cowley County, the uppermost of which
forms the prominent first crest of the Flint Hills. As the rapid-
flowing western tributaries of Big Caney descend over these suc-
cessive limestone members, large quantities of chert and limestone
rubble are transported and deposited in stream beds of the system.
In many places the streams of the Big Caney system flow over re-
sistant limestone members, which form a bedrock bottom. The
eastern tributaries of Big Caney drain, for the most part, forma-
tions of the Wabaunsee group of the Pennsylvanian. Most of these
streams have lower gradients than those entering Big Caney from
the west. The tributaries of Big Caney, along with length in miles
and gradient in feet per mile, are as follows: Spring Creek, 7.1, 54.5;
Union Creek, 6.3, 42.9; Otter Creek, 14.6, 27.4; Cedar Creek, 11.6,
31.0; Rock Creek, 15.9, 26.5; Wolf Creek, 9.3, 17.2; Turkey Creek,
8.5, 26.4; Grant Creek, 13.9, 23.4; and Sycamore Creek, 8.9, 27.0.

Spring Creek and Union Creek are short and have formed no
extensive floodplain. The high gradients of these creeks are char-
acteristic also of the upper portions of several other tributaries
such as Cedar Creek and Otter Creek.

Middle Caney Creek (Fig. 1) has its source in the Wabaunsee
and Shawnee groups of the Pennsylvanian but its watershed is
dominated by the "Chautauqua Hills" of the Douglas Group. This
area is described by Moore (1949:127) as "an upland formed by
hard sandstone layers." The rough rounded hills supporting thick
growths of oaks differ in appearance from both the Big Caney
watershed on the west and the Verdigris River watershed on the
east. The gradient of Middle Caney in Chautauqua County is

Fishes of Arkansas River Basin 351

10.8 feet per mile. Its largest tributary, North Caney Creek, has
a gradient of 15.5 feet per mile.

The Elk River Basin resembles the Big Caney River Basin topo-
graphically. Elk River has a gradient of 14.4 feet per mile.

PHYSICAL CHARACTERISTICS OF STREAMS

The stream channels derive their physical characteristics from the geological
make-up of the area and from land-use. The Arkansas River typically has low
banks; however, in a few places, as in the NE /4 of Section 21, T. 33 S, R. 3 E,
it cuts into limestone members to form steep rocky banks. The bottom is pre-
dominantly sand. In years of heavy rainfall the river is turbid, but during
1956, when it occupied only a small portion of its channel, it was clear each
time observed. All streams surveyed were clear except after short periods
of flooding in June, and except in some isolated pools where cattle had
access to the water.

In the Walnut River, sand bottoms occur in the lower part of the stream
but the sand is coarser than that of the Arkansas River. Upstream, gravel
and rubble bottoms become more common. Steep rocky banks border
most of the course of the Walnut. During 1956, stream-flow was confined
to the center of the channel, remote from these rocky banks.

The rubble and bedrock bottoms found in most streams of the Flint Hills
have been described. In the alluvial valleys of their lower courses mud
bottoms are found. Gravel is present in some places but sand is absent.
Banks are variable but often steep and wooded. Along east- or west-flowing
streams the north bank characteristically is low and sloping whereas the
south bank is high, rises abruptly, and in many places is continuous with
wooded hills. The lower sections of Otter Creek, Cedar Creek, and Rock
Creek fit this description (Bass, 1929:19) especially well, as does Elk River
near Howard.

Streams in the Chautauqua Hills resemble those of the Flint Hills in physi-
cal characteristics, except that a larger admixture of sandstone occurs in the

rubble.

CLIMATE

The climate of the area is characterized by those fluctuations of tempera-
lure, wind, and rainfall typical of the Great Plains. The mean annual temper-
ature is 58 degrees; the mean July temperature is 81 degrees; the mean Janu-
ary temperature is approximately 34 degrees. The mean annual precipitation
is 32.9 in Cowley County, 38.5 in Chautauqua County, and 35.1 in Elk County.
Wind movement is great; Flora (1948:6) states that south-central Kansas
ranks close to some of the windiest inland areas in the United States.

The area has been periodically subjected to droughts and floods. Such
phenomena are of special interest to ichthyological workers in the area. At
the time of this study drought conditions, which began in 1952, prevailed.
Even in this period of drought, however, flooding occurred on Grouse Creek
and water was high in Big Caney River after heavy local rains on the head-
waters of these streams on June 22, 1956. Some of the lower tributaries of
these same streams (such as Crab Creek and Cedar Creek) did not flow while
the mainstreams were flooding. This illustrates the local nature of many of
the summer rains in the area.

352

Unr-ersity of Kansas Publs., Mus. Nat. Hist.

Table 1 indicates maximum, minimum, and average discharges in cubic
feet per second at several stations in the area and on nearby streams. These
figures were pro\'ided by the U. S. Geological Survey.

Table 1. — Cubic Feet Per Second of Water Discharged at Gauging
Stations in Chautauqua, Elk, Montgomery, and Cowley CotrNTiES for
Years Prior to 1931.

Gauging
station

Drain-
age
area

(sq. mi.)

Aver-
age
dis-
charge

Maxi-
mum
dis-
charge

Date

Mini-
mum
dis-
charge

Date

Arkansas River
at Arkansas City

43,713

1,630

103,000

June 10,
1923

1

October 9,
1921

Walnut River at
Winfield

1,840

738

105,000

April 23,
1944

0

1928, 1936

Big Caney River
at Elgin

445

264

35,500

April 10,
1944

0

1939, 1940,
1946, 1947

Elk River near
Elk City

575

393

39,200

April 16,
1945

0

1939, 1940,
1946

Fall River near
Fall River

591

359

45,600

April 16,
1945

0

1939, 1940,
1946

Verdigris River
at Independence

2,892

1,649

117,000

April 17,
1945

0

1932, 1934,
1936, 1939,
1940

Something of the effect that drought and flash-flood have had on Big Caney
River is shown by the monthly means of daily discharge from October, 1954, to
September, 1956, at the stream-gauging station near Elgin, Kansas (Table 2).
Within these monthly variations there are also pronounced daily fluctuations;
on Big Caney River approximately 34 mile south of Elgin, Kansas, discharge
in cubic feet per second for May, 1944, ranged from .7 to 9,270.0 and for May,
1956, from .03 to 20.0.

Table 2. — Monthly Means of Daily Discharge in Cubic Feet per Second
FOR Big Caney River at Elgin, Kansas

Month
October . .
November
December
January . .
February .
March . . .

1954-55 1955-56
103.00 69.60

.31

.18

.78

4.76

3.37

.78
1.92
1.65
2.08
1.27

Month 1954-55 1955-56

April 4.91 47

May 624.00 7.37

June 51.30 35.20

July 1.20 1.85

August 0.00 0.00

September . . .04 0.00

Fishes of Arkansas River Basin 353

PRESENT FLORA

The flora of the region varies greatly at the present time. Land-use has
altered the original floral communities, especially in the intensively cultivated
area of western Cowley County and in the river valleys.

The sandy Arkansas River floodplain exhibits several stages ranging from
sparsely vegetated sandy mounds near the river through stages of Johnson
grass, willow, and cottonwood, to an elm-hackberry fringe-forest. The Well-
ington formation bordering the floodplain supports a prairie flora where not
disturbed by cultivation; Gates (1936:15) designates this as a part of the
mixed bluestem and short-grass region. Andropogon gerardi Vitman., Andro-
pogon scoparius Michx., Sorgliastrum nutans (L. ), and Panicum virgatum L.
are important grasses in the hilly pasture-lands. Although much of this land
is virgin prairie, the tall, lush condition of the grasses described by early
writers such as Mooso (1888:304), and by local residents, is not seen today.
These residents speak of slough grasses (probably Tripsacum dactyloides L.
and Spartina pectinate Link.) that originally formed rank growths. These
no doubt helped conserve water and stabilize flow in small headwater creeks.
Remnants of some of these sloughs can still be found. The streams in the
Flint Hills have fringe-forests of elm, hackberry, walnut, ash, and willow.

Eastward from the Flint Hills these fringe-forests become thicker with a
greater admixture of hickories and oaks. The north slopes of hills also be-
come more wooded. However, grassland remains predominant over wood-
land in western Chautauqua and Elk counties, whereas in the eastern one-half
of Chautauqua County and the eastern one-third of Elk County the wooded
Chautauqua Hills prevail. This is one of the most extensive wooded upland
areas in Kansas. Hale (1955:167) describes this woodland as part of an
ecotonal scrub-oak forest bordering the Great Plains south through Texas.
He found stand dominants in these wooded areas to be Quercus marilandica
Muenchh., Quercus stellata Wang., and Quercus velutina Lam.

Few true aquatic plants were observed in the Arkansas River although
mats of duckweed were found in shallow backwater pools at station A-3
(Fig. 2) on December 22, 1956. In the Walnut River Najas guadalupensis
Spreng. was common at station W-2. Stones were usually covered with algae
in both the Arkansas and Walnut rivers. A red bloom, possibly attributable
to Euglena rubra (Johnson), was observed on a tributary of the Walnut
River on July 9, 1956, at station W-4.

Green algae were abundant at all stations in the Caney, Elk, and Grouse
systems during May and June, 1956, and reappeared late in September.
Chara sp. was common in these streams in April and May.

The most characteristic rooted aquatic of streams in the Flint Hills was
Justicia americana L. At station G-7 on Grouse Creek and Station C-8 on
Big Caney River (Fig. 3), Nelumbo lutea (Willd. ) was found. Myriophyllum
heterophyllum Michx. formed dense floating mats at a number of stations.
Other aquatic plants observed in the Caney, Elk, and Grouse systems in-
cluded Potamogeton gramineus L., Potamogeton nodosus Poir., Potamogeton
foliosus Raf., Sagittaria latifolia WUld., Typha latifolia L., and Jussiaea diffusa
Forsk.

354 University of Kansas Publs., Mus. Nat. Hist.

HISTORY

In 1857, a survey was made of the southern boundary of Kansas. Several
diaries (Miller, 1932; Caldwell, 1937; Bieber, 1932) were kept by members
of the surveying party, which traveled from east to west. These accounts
contain complaints of difficulty in traversing a country of broken ridges and
guUeys as the part\' approached the area now comprising Chautauqua County.
One account by Hugh Campbell, astronomical computer for the party (Cald-
well, 1937) mentions rocky ridges covered with dense growth of "black jack,"
while another by Col. Joseph Johnson, Commander (Miller, 1932) speaks
of "a good deal of oakes in the heights" — indicating that the upland oak
forest of the Chautauqua Hills was in existence at that time. On reaching
Big Caney River near Elgin, Campbell wrote of a stream with very high
banks and of a valley timbered with oak and black walnut. While the party
was encamped on Big Caney River some fishing was done. Campbell (Cald-
well, 1937:353) described the fish taken as "Cat, Trout or Bass, Buffalo and
Garr." Eugene Bandel (Bieber, 1932:152) wrote, "This forenoon we did
not expect to leave camp, and therefore we went fishing. In about two hours
we caught more fish than the whole company could eat. There were some
forty fish caught, some of them weighing over ten pounds." It was noted that
the waters of Big Caney and its tributaries were "very clear." Progressing
up Rock Creek, Johnson wrote of entering a high rolling plain covered with
fine grass, and crossed occasionally by clear wooded streams (probably Big
and Little Beaver Creeks and Grouse Creek). The diary of Hugh Campbell
(Caldwell, 1937:354) contains a description of the Arkansas River Valley
near the Oklahoma border. "The Arkansas River at this point is about 300
yards wide, its waters are muddy, not quite so much so, as those of the
Mississippi or Rio Bravo. Its valley is wooded and about two miles in width,
the main bottom here, being on the east side. On the west it is a rolling
prairie as far as the eye can see, affording excellent grass." Some seining
was done while encamped on the Arkansas River and "buffalo, catfish, stur-
geons, and gars" were taken (Bieber, 1932:156).

An editorial in the Winfield Courier of November 16, 1899, vigorously
registers concern about a direct effect of settlement on fish populations in
rivers of the area:

"The fish in the streams of Cowley Coimty are being slaughtered by the
thousands, by the unlawful use of the seine and the deadly hoop net. Fish
are sold on the market every day, sometimes a tubful at a time, which never
swallowed a hook.

"The fish law says it is imlawful to seine, snare, or trap fish but some of
the smaller streams in the county, it is said are so full of hoop and trammel
nets that a minnow cannot get up or down stream. These nets not only
destroy what fish there are in the streams but they keep other fish from
coming in, they are not operated as a rule by farmers to supply their own
tables but by fellows who catch the fish to sell with no thought or care for
the welfare of others who like to catch and eat fish.

"If there is a fishwarden in Cowley County so far as his utility goes the
county would be as well off without him and his inactivity has caused many
of those interested to get together for the purpose of seeing that the law is
enforced.

Fishes of Arkansas River Basin 355

"Depredations like this work injury in more ways than one. They not only
deplete the streams of fish large enough to eat and destroy the source of supply
but if the U. S. Fish Commission discovers that the law is not enforced and
the fish not protected, there will be no free government fish placed in Cowley
County streams. It is useless for the Government to spend thousands of
dollars to keep the streams well supplied if a few outlaws are allowed to
nithlessly destroy them. The new organization has its eye on certain parties
now and something is liable to drop unexpectedly soon."

Graham (1885:78) hsted 13 species of fish that had already been intro-
duced into Kansas waters prior to 1885 by the State Fish Commission.

These early references indicate that direct effects of settlement on the
native flora and fauna were recognized early. Concern such as that expressed
in the editorial above persists today; however, it is not clear whether the fish
fauna of the streams of the area has been essentially changed by man's
predation. The indirect effects tlirough human modifications of the environ-
ment seem to be of much importance. Three modifications which have
especially affected streams have been agricultural use, urbanization, and in-
dustrialization.

The effect of land-use on streams is closely related to its effect on the flora
of the watershed. Turbidity, sedimentation, and the rate, periodicity, and
manner of flow all bear some relationship to the land-use of the watershed.
Stream-flow in the area has been discussed in the section on climate.

The effects of urbanization are more tangible and better recognized than
those of agricultural land-use. Streams that flow through cities and other
populous areas undergo some modification, especially of the streamside flora.
Another effect of urbanization has been increased loads of sewage dis-
charged into the streams. The combined populations of Arkansas City and
Winfield rose from 3,986 in 1880 to 23,167 in 1950. Arkansas City found
it necessary to construct a sewage system in 1889; Winfield in 1907.

There are, at the present time, nine towns within the area that have
municipal sewage systems. The State Training Home at Winfield also has
a sewage system. The Kansas State Board of Health, Division of Sanitation,
has provided information concerning adequacy of these systems and certain
others in nearby counties as of February 5, 1957. This infonnation is shown
in Table 3.

Representatives of the Division of Sanitation, Kansas State Board of Health,
expressed the belief that pollution by both domestic sewage and industrial
wastes would be largely eliminated in the "lower Arkansas" and in the
Walnut watershed by 1959.

Important oil and gas resources have been discovered in each of the three
counties. The first producing wells were drilled between 1900 and 1902
(Jewett and Abernathy, 1945:24). The Arkansas River flows through sev-
eral oilfields in its course across Cowley County (Jewett and Abernathy,
1945:97). A number of producing wells have been drilled in the Grouse
Creek watershed since 1939 and many of these wells are near the banks of
the creek. In the Big Caney watershed of Cowley and Chautauqua counties
there has been little oil production in recent years; however, a few small
pools are presently producing in southwestern Elk County.

Clapp (1920:33) stated that "Many of the finest streams of our state are
now destitute of fish on account of oil and salt pollution. The Walnut River,

356

Unu'ersity of Kansas Publs., Mus. Nat. Hist.

Table 3. — Sewage Dispos.al Facilities in Some South-central Kansas

Communities.

Community

Status on February 5, 1957

Remarks

Cowley County:

Arkansas City

Geuda Springs

Winfield

State training school
Udall

Discharging raw sewage ....

Discharging raw sewage

Inadequate

Adequate

Adequate

Adequate plant in
design stage.

Chautauqua Count}':
Cedar Vale

Inadequate

Adequate

Sedan

In operation 30 days.

Elgin

Adequate

Elk County:

Moline

Howard

Inadequate
Adequate

Sumner County:

Belle Plaine

Mulvane

Oxford

Discharging raw sewage ....
Discharging raw sewage ....
Discharging raw sewage ....

Adequate plant under

construction.
Adequate plant under

construction.
Construction on

adequate plant

to start soon.

Butler County:

Augusta

El Dorado

Douglass

Adequate

Discharging raw sewage ....

Discharging raw sewage ....

Adequate plant under
construction.

Adequate plant to go
into operation
within 30 days.

once as fine a bass stream as could be found anywhere, and a beautiful
stream, too, is now a murky oil run, and does not contain a single fish so far
as I know. The Fall and Verdigris rivers are practically mined. Both the
Caney rivers are affected, and may soon be ruined for fishing." Doze ( 1924:31 )
noted "Some of the finest streams in the state have been ruined as habitat for
wild life, the Walnut River is probably the most flagrant example."

Pollution by petroleum wastes from refineries has also affected tlie streams
studied. The only refinery within the area is at Arkansas City. In Butler
County there are four refineries on the Walnut watershed upstream from the
area surveyed. Metzler (1952) noted that "fish-kills" occurred from the
mid-1940's until 1952 in connection with wastes periodically discharged from
these refineries. However, the largest kill, in 1944, was attributed to exces-
sive brine pollution.

In Arkansas City a meat-packing plant, a large railroad workshop, two flour
mills, two milk plants, and several small manufacturing plants contribute
wastes which may figure in industrial pollution. There arc milk plants and
small poultry processing plants at Winfield. In Chautauqua and Elk Counties
there is little industrial activity.

Fishes of Arkansas River Basin 357

CONSERVATION

In recent years several measures have been implemented or
proposed to conserve the water and land resources of the Arkansas
River Basin. Droughts and floods have focused public attention
on such conservation. Less spectacular, but nevertheless important,
problems confronting conservationists include streambank erosion,
channel deterioration, silting, recreational demands for water, and
irrigation needs.

Congress has authorized the U. S. Corps of Engineers (by the
Flood Control Act of 1941) to construct six dam and reservoir
projects in the Verdigris watershed. Two of these — Hulah Res-
ervoir in Osage County, Oklahoma, on Big Caney River, and Fall
River Reservoir in Greenwood County, Kansas — have been com-
pleted. Other reservoirs authorized in the Verdigris watershed
include Toronto, Neodesha, and Elk City (Table Mound) in Kansas
and Oologah in Oklahoma. Construction is underway on the To-
ronto Reservoir and some planning has been accomplished on the
Neodesha and Elk City projects.

The possibilities of irrigation projects in the Verdigris and Wal-
nut River basins are under investigation by the United States
Bureau of Reclamation (Foley, et ah, 1955 :F18).

An area of 11 square miles in Chautauqua and Montgomery
Counties is included in the Aiken Creek "Pilot Watershed Project,"
a co-operative effort by federal, state, and local agencies to obtain
information as to the effects of an integrated watershed protection
program (Foley, et al, 1955:131).

PREVIOUS ICHTHYOLOGICAL COLLECTIONS

Few accounts of fishes in the area here reported on have been pubhshed.
Evermann and Fordice (1886:184) made a collection from Timber Creek at
Winfield in 1884.

The State Biological Survey collected actively from 1910 to 1912, but
localities visited in the Arkansas River System were limited to the Neosho
and Verdigris River basins (Breukelman, 1940:377). The only collection
made in the area considered here was on tlie Elk River in Elk County on July
11, 1912. The total species list of this collection is not known.

In the years 1924-1929 Minna E. Jewell collected at various places in
central Kansas. On June 30, 1925, Jewell and Frank Jobes made collections
on Timber Creek and Silver Creek in Cowley County.

Hoyle (1936:285) mentions collections made by himself and Dr. Charles
E. Burt, who was then Professor of Biology at Southwestern College, Winfield,
Kansas. Records in the Department of Biology, Kansas State Teachers Col-
lege at Emporia, indicate that Dr. Burt and others made collections in the
area which have not been published on.

358

University of Kansas Publs., Mus. Nat. Hist.

Table 4. — Collections Made by Dr. Frank B. Cross of the State

Biological Survey in 1955.

Collection number

Date

River

Location

C-131

April 5, 1955

Elk

Sec. 3, T31S, RUE

C-132

April 5, 1955

Sj'camore

Sec. 5, T34S, RIOE

C-133

April 5, 1955

Big Caney

Sec. 12, T34S, R8E

C-136

April 6, 1955

Walnut

Sec. 29 or 32, T32S, R4E

Claire Schelske ( 1 957 ) studied fishes of the Fall and Verdigris Rivers in
Wilson and Montgomery counties from March, 1954, to February, 1955.

In the annotated list of species that follows, records other than mine are
designated by the following symbols:

E&F — Evermann and Fordice

SBS— State Biological Survey (1910-1912)

J&J — Jewell and Jobes (collection on Silver Creek)

C — Collection number — Cross (State Biological Survey, 1955)

UMMZ — University of Michigan Museum of Zoology

OAM — Oklahoma A&M College Museum of Zoology

ACKNOWLEDGMENTS

I am grateful to Professor Frank B. Cross for his interest in my investigation,
for his counsel, and for his penetrating criticism of this paper. This study
would have been impossible without the assistance of several persons who
helped in the field. Mr. Artie C. Metcalf and Mr. Delbert Metcalf deserve
special thanks for their enthusiastic and untiring co-operation in collecting
and preserving of specimens. Mrs. Artie C. Metcalf, Miss Patricia Metcalf,
Mr. Chester Metcalf, and Mr. Forrest W. Metcalf gave help which is much
appreciated. I am indebted to the following persons for numerous valuable
suggestions: Dr. John Breukelman, Kansas State Teachers College, Emporia,
Kansas; Dr. George Moore, Oklahoma A&M College, and Mr. W. L. Minckley,
Lawrence, Kansas.

MATERIALS AND METHODS

Collections were made by means of: ( 1 ) a four-foot net of nylon screen;
(2) a 10x4-foot "common-sense" woven seine with /^-inch mesh; (3) a 15x4-
foot knotted mesh seine; (4) a 20x5-foot ;4-inch mesh seine; (5) pole and hne
(natural and artificial baits). At most stations the four-foot, ten-foot, and
twenty-foot seines were used; however, the equipment that was used varied
according to the size of pool, number of obstructions, nature of bottom, amount
of flow, and type of strcambank. Usually several hours were spent at each
station and several stations were revisited from time to time. Percentages
noted in the List of Species represent the relative number taken in the first
five seine-hauls at each station.

Fishes of Arkansas River Basin

359

COLLECTING STATIONS

Collecting was done at stations listed below and shown in Fig. 2. Each
station was assigned a letter, designating the stream system on which the
station was located, and a number which indicates the position of the station
on the stream. This number increases progressively upstream from mouth
to source. Code letters used are as follows: A — Arkansas River; W — Walnut
River System; B — Beaver Creek System; C — Big Caney River System; G —
Grouse Creek System; M — Middle Caney Creek System; E — Elk River System.
All dates are in the year 1956.

Fig. 2. Map of Cowley, Chautauqua and Elk counties, Kansas, showing
stations at which collecting was done.

35 S, R. 4 E. June 14 and August
Water slightly turbid, with layer

Sec. 22, T. 34 S, R. 3 E. August 25. Flowing
Average depth 12 inches. Bottom sand. (Plate 9,

A-1. Arkansas River. Sec. 2 and 3, T
20. Braided channel with sand bottom,
of oil sludge on bottom.

A-2. Arkansas River,
tlirough diverse channels,
fig. 1.)

A-3. Arkansas River. Sec. 21, T. 33 S, R. 3 E. August 27 and December
22. Flowing over fine sand. Average depth 11 inches. Some areas of back-
water with oil sludge on bottom.

W-1. Walnut River. Sec. 20, T. 34 S, R. 4 E. July 7. Flowing rapidly,
with large volume, because of recent rains. Average width 300 feet. Bottom
gravel. Water turbid.

W-2. Walnut River. Sec. 11, T. 34 S, R. 4 E. July 20. Rubble riffles
and large shallow pools with gravel bottoms. Average width, 100 feet. Water
clear.

W-3. Walnut River. Sec. 29, T. 32 S, R. 4 E. July 17. Pools and
riffles below Tunnel Mill Dam at Winfield. Water clear.

W-4. Badger Creek. Sec. 6, T. 33 S, R. 5 E. July 17. Small pools.
Average width 7 feet, average length 40 feet, average depth 8 inches. Water
turbid and malodorous. Bottoms and banks mud. Much detritus present.

360 University of Kansas Publs., Mus. Nat. Hist.

W-5. Timber Creek. Sec. 35, T. 31 S, R. 4 E. June 6. Intermittent
pools, widely separated. Average width 9 feet, average depth 8 inches. Bot-
tom mud and gravel.

B-1. Big Beaver Creek. Sec. 8, T. 35 S, R. 7 E. May 28. Isolated
pools. Average width 10 feet, average depth one foot. Water turbid. Bot-
tom rubble.

B-2. Little Beaver Creek. Sec. 18, T. 35 S, R. 6 E. July 21. Inter-
mittent pools. Average width 10 feet, average length 35 feet, average depth
10 inches. Bottoms rubble, mud, and bedrock.

B-3. Big Beaver Creek. Sec. 28, T. 34 S, R. 7 E. July 22. Series of
small turbid pools.

G-1. Grouse Creek. Sec. 5, T. 35 S, R. 5 E. May 30, September 5, and
September 24. Intermittent pools in close succession. Average width 22
feet, average depth 16 inches. Water turbid on May 30 but clear in Sep-
tember. Bottom rubble. Steep banks. Little shade for pools.

G-2. Grouse Creek. Sec. 23, T. 34 S, R. 5 E. August 29. Series of
shallow intermittent pools. Average width 42 feet, average length 120 feet,
average depth 15 inches. Bottom bedrock and mud. (Plate 9, fig. 2.)

G-3. Grouse Creek. Sec. 6, T. 34 S, R. 6 E. July 12. Intermittent
pools. Average width 20 feet, average length 65 feet, average depth 14
inches. Bottom bedrock and gravel. Justicia americana L. abundant.

G-4. Grouse Creek. Sec. 12, T. 33 S, R. 6 E. June 1 and September 7.
Intermittent pools. Average width 15 feet, average length 100 feet, average
depth 18 inches. Water turbid in June, clear in September. Najas guadalu-
pensis Spreng., and Myriophyllum heterophyllum Michx. common.

G-5. Grouse Creek. Sec. 19, T. 32 S, R. 7 E. July 2. Succession of
riffles and pools. Water clear. Volume of flow approximately one cubic foot
per second, but creek bankful after heavy rains on June 22. Average width 20
feet, average depth 18 inches.

G-6. Grouse Creek. Sec. 32, T. 31 S, R. 7 E. July 8. Small intermittent
pools to which cattle had access. Water turbid, bottom mud and rubble.
Average width 10 feet, average depth 8 inches. Stream-bed covered with
tangled growths of Sorghum halepense ( L. ) .

G-7. Grouse Creek. Sec. 34, T. 30 S, R. 7 E. July 8. Stream flowing
slightly. Water clear. Average width of pools 30 feet; average depth 20
inches. Bottom bedrock and gravel. Myriophyllum heterophyllum Michx.,
Nelumho lutea (Willd. ), and Justicia americana L. common in shallow water.

G-8. Silver Creek. Sec. 1, T. 33 S, R. 5 E. July 17. Intermittent pools.
Average width 30 feet, average length 120 feet, average depth 12 inches.
Water clear.

G-9. Silver Creek. Sec. 4, T. 32 S, R. 6 E. July 17. Small upland
brook with volume less than one-half cfs. Average width 12 feet, average
depth 10 inches. Water clear, bottom mostly rubble.

G-10. Crab Creek. Sec. 33, T. 33 S, R. 6 E. June 24. Intermittent pools,
showing evidence of having flowed after rains on June 22. Average width 15
feet, average depth 16 inches.

G-11. Crab Creek. Sec. 35, T. 33 S, R. 6 E. July 16. Small intermittent
pools. Average width 13 feet, average length 55 feet, average depth 11
inches. Water clear. Bottom rubble and mud.

G-12. Crab Creek. Sec. 28, T. 33 S, R. 7 E. June 2 and July 20. Iso-
lated pools. Average width 18 feet, average depth one foot. Water turbid.
Bottom bedrock and rubble. Myriophyllum heterophyllum and Justicia
americana abundant.

G-13. Crab Creek. Sec. 21, T. 33 S, R. 7 E. July 29. Isolated pools
300 feet by 24 feet. Average depth 12 inches. Water turbid.

G-14. Unnamed creek (hereafter called Grand Summit Creek). Sec. 26,
T. 31 S, R. 7 E. August 30. Intermittent pools. Average width 15 feet,
average length 45 feet, average depth 11 inches. Water clear. Bottom rubble.

PLATE 9

1. Station A-2. Arkansas River. (Cowley County, Section 22, T. 34 S, R. 3 E.)

2. Station G-2. Grouse Creek. (Cowley County, Section 23, T. 34 S, R. 5 E.)

PLATE 10

1. Slatiun C-12. Cedar Creek. (Cowley County, Seetion 17, T. 34 S, R. 8 K.)

2. Station C-lfi. Spring Creek. (Elk County, Seetion 20. '!'. .n S, H. « E.)
N'ohnne ol flow ol tliis small creek is iiidicrited l)\ ridlc in foreiironnd.

Fishes of Arkansas River Basin 361

G-15. Unnamed creek (same as above). Sec. 17, T. 31 S, R. 8 E. July
27. Small upland creek bordered by bluestem pastures. Pools with average
width of 10 feet, average length 30 feet, average depth 9 inches. Water
slightly turbid. Bottom rubble and mud.

G-16. Crab Creek. Sec. 22, T. 33 S, R. 7 E. July 25. Small isolated
pools. Average width 17 feet, average length 58 feet, average depth 9 inches.
Water turbid.

G-17. Crab Creek. Sec. 23, T. 33 S, R. 7 E. July 25. Upland brook
bordered by bluestem pastures. Unshaded intermittent pools. Average width
7 feet, average length 40 feet, average depth 9 inches. Water turbid.

C-1. Big Caney River. Sec. 16, T. 33 S, R. 10 E. July 19. Intermittent
pools. Average width 47 feet, average length 90 feet, average depth 13 inches.
Bottom rubble and bedrock. Water clear to slightly turbid.

C-2. Big Caney River. Sec. 1, T. 35 S, R. 9 E. September 5. Series
of intermittent pools. Bottom rubble and large stones.

C-3. Big Caney River. Sec. 29, T. 34 S, R. 9 E. ^^ June 17. Large
shallow pool below ledge 3 feet high forming "Osro Falls." Bottom bedrock.

C-4. Big Caney River. Sec. 32, T. 34 S, R. 9 E. June 3. Three large
pools (50 feet by 300 feet) with connecting riflBes. Water turbid. Bottom
bedrock and rubble.

C-5. Big Caney River. Sec. 11 and 12, T. 34 S, R. 8 E. May 27, May
29, June 11, June 18, June 19, and June 27. From a low-water dam, 6 feet
high, downstream for /4 mile. Pools alternating with rubble and bedrock
riffles. Collecting was done at different times of day and night, and when
stream was flowing and intermittent.

C-6. Big Caney River. Sec. 26, T. 33 S, R. 8 E. June 16. Intermittent
pools with bedrock bottom. Water slightly turbid. Average width 16 feet,
average depth 10 inches.

C-7. Otter Creek. Sec. 26, T. 33 S, R. 8 E. June 16. Pools and riffles.
Water clear. Algae abundant. Average width 10 feet, average depth 10
inches.

C-8. Big Caney River. Sec. 1, T. 33 S, R. 8 E. June 10. Intermittent
pools. Average width 10 feet, average depth 14 inches. Water clear. Bottom
rubble and gravel. Aquatic plants included Chara sp., Sagittaria latifolia
Willd., Jussiaea diffusa Forsk., and Nelumbo lutea (Willd. ).

C-9. Big Caney River. Sec. 6 and 7, T. 32 S, R. 9 E. June 27. Clear,
flowing stream, 20 feet wide, volume estimated at 5 cfs. Bottom gravel and
rubble. Extensive gravel riffles.

C-10. Big Caney River. Sec. 29 and 32, T. 31 S, R. 9 E. June 27.
Water clear and flowing rapidly, volume estimated at 5-6 cfs. Bottom rubble
with a few muddy backwater areas.

C-11. Big Caney River. Sec. 7, T. 31 S, R. 9 E. July 26. Flowing, vdth
less than 1 cfs. Average width 20 feet, average depth 22 inches. Water
extremely clear. Bottom gravel and rubble. Myriophyllum heterophyllum,
Potamogeton foliosiis, and Justicia americana common.

C-12. Cedar Creek. Sec. 17, T. 34 S, R. 8 E. March 10, April 2, June
1, June 6, and August 24. Pools and riffles along M mile of stream were
seined in the early collections. In August only small isolated pools remained.
Bottom bedrock and rubble. Much detritus along streambanks. (Plate 10,

fig. 1.)

C-13. Otter Creek. Sec. 16, T. 33 S, R. 8 E. June 15. Flowing, less than
1 cfs. Pools interspersed with rubble riffles. Water clear.

C-14. Otter Creek. Sec. 30, T. 32 S, R. 8 E. May 31, and September
3. Series of small pools. Average width 10 feet, average depth 15 inches.
Shallow rubble riffles. Water extremely clear. Temperature 68° at 6:30
p. m. on May 31; 78° at 2:00 p. m. on September 3.

C-15. Spring Creek. Sec. 35, T. 31 S, R. 8 E. June 28. Small, clear,

2—7079

362 University of Kansas Publs., Mus. Nat. Hist.

upland brook with nibble bottom. Pools 10 feet in average width and 11
inches in average depth. Numerous shallow rubble riflBes.

C-16. Spring Creek. Sec. 26, T. 31 S, R. 8 E. July 9. Small intermittent
pools. Average width 10 feet; average depth 8 inches. Bottom gravel.
(Plate 10, fig. 2.)

C-17. West Fork Big Caney River. Sec. 36, T. 30 S, R. 8 E. July 27.
Small pool below low-water dam. Pool 20 feet by 30 feet with average depth
of 20 inches.

C-18. East Fork Big Caney River. Sec. 31, T. 30 S, R. 9 E. July 27,
Isolated pool 25 feet by 25 feet with an average depth of 15 inches.

M-1. Middle Caney Creek. Sec. 23, T. 33 S, R. 10 E. July 4. Intermit-
tent pools. Average width 45 feet, average depth 15 inches. Water stained
Ijrown. Oil fields nearby but no sludge or surface film of oil noted. Bottom
rubble and bedrock.

M-2. Pool Creek. Sec. 25, T. 33 S, R. 10 E. May 26. Pool 120 feet by
40 feet below hmestone ledge approximately 12 feet high forming Butcher's
Falls. Other smaller pools sampled. Water clear. Bottom bedrock and
rubble.

E-1. Elk River. Sec. 12, T. 31 S, R. 11 E. July 9. Four intermittent
pools seined. Average width 32 feet, average depth 13 inches. Bottom bed-
rock, rubble, and mud. Water turbid.

E-2. Elk River. Sec. 3, T. 31 S, R. 11 E. June 28. Intermittent pools
below and above sandstone ledge approximately 6 feet high forming "falls"
at Elk Falls. Average width 33 feet, average depth 15 inches. Bottom bed-
rock, rubble and mud. Water slightly turbid.

E-3. Elk River. Sec. 21, T. 30 S, R. 11 E. June 28. Two small pools,
10 feet by 30 feet viath average depth of 6 inches. Bottom bedrock.

E-4. Elk River. Sec. 12, T. 30 S, R. 10 E. June 28. One long pool 500
feet by 50 feet with a variety of depths and bottom conditions ranging from
mud to bedrock. Average depth 18 inches. Water turbid and pools unshaded.

E-5. Elk River. Sec. 32, T. 29 S, R. 10 E. August 30. Intermittent
pools. Average width 21 feet, average depth 20 inches. Bottom nibble.
Water clear.

E-6. Elk River. Sec. 23, T. 29 S, R. 9 E. August 30. Small isolated
pools. River mostly dry. Bottom bedrock. Water slightly turbid with gray-
green "bloom."

E-7. Wildcat Creek. Sec. 11, T. 31 S, R. 10 E. Volume of flow less
than one cfs. Average width 20 feet, average deptli 18 inches. Domestic
sewage pollution from town of Moline suspected.

ANNOTATED LIST OF SPECIES

Lepisosteus osseus oxyurus ( Linnaeus ) : Stations A-1, W-2, W-3,
G-2, G-3, G-4, C-1, C-2, C-3, C-5, C-8.

Of 34 longnose gar taken, 27 were young-of-the-ycar. The lat-
ter were from shallow isolated pools (bedrock bottom at C-1, C-3,
C-4; gravel bottom at C-6). At station W-1 in moderate flood con-
ditions several young-of-the-year were found in the most sheltered
water next to the banks.

The longnose gar was found only in the lower parts of the streams
surveyed (but were observed by me in smaller tributaries of these
streams in years when the streams had a greater volume of flow).
A preference for downstream habitat is suggested in several other
surveys: Cross (1950:134, 1954a:307) on the South Fork of the

Fishes of Arkansas River Basin 363

Cottonwood and on Stillwater Creek; Cross and Moore (1952:401)
on the Poteau and Fourche Maline rivers; Moore and Buck
(1953:21) on the Chikaskia River.

Lepisosteus platostomus Rafinesque: One shortnose gar (K. U.
3157) has been taken from the Arkansas River in Cowley County.
This gar was taken by Mr. Richard Rinker on a bank line on April
10, 1955, at station A-3.

Dorosoma cepedianum ( Le Sueur ) : Stations W-3, G-4, C-4, C-5,
M-1, E-1, E-4.

In smaller streams such as the Elk and Caney rivers adult gizzard
shad seemed scarce. They were more common in collections made
in larger rivers (Walnut, Verdigris, and Neosho). In impound-
ments of this region shad often become extremely abundant.
Schoonover (1954:173) found that shad comprised 97 per cent by
number and 83 per cent by weight of fishes taken in a survey of
Fall River Reservoir.

Carpiodes carpio carpio (Rafinesque): Stations A-1, A-2, A-3,
W-3, G-1, C-3.

Hubbs and Lagler (1947:50) stated that the river carpsucker was
"Mostly confined to large silty rivers." Of the stations listed above
C-3 least fits this description being a large shallow pool about % acre
in area having bedrock bottom and slightly turbid water. The
other stations conform to conditions described by Hubbs and
Lagler (loc. cit.).

Carpiodes velifer (Rafinesque): SBS. Three specimens of the
highfin carpsucker (K. U. 177-179) were collected on July 11,
1912, from an unspecified location on Elk River in Elk County.

Ictiobus bubalus ( Rafinesque ) : Stations W-3, G-1, G-2, C-1, C-3,
C-4, C-6, E-1, E-2, E-3.

The smallmouth buff^alo shared the downstream proclivities of
the river carpsucker. In half of the collections (G-2, C-1, E-1,
E-2, E-3) only large juveniles were taken; in the other half only
young-of-the-year were found. In one pool at station C-1 hundreds
of young buffalo and gar were observed. This large shallow pool
was 100 X 150 feet, with an average depth of 8 inches. The bot-
tom consisted of bedrock. Station C-6 was a small pool with bed-
rock bottom, eight feet in diameter, with an average depth of only
4 inches. Station E-3 was also a small isolated pool with bedrock
bottom and an average depth of 6 inches.

Ictiobus niger ( Rafinesque ) : Station C-5.

Only two specimens of the black buffalo were taken. An adult

364 University of Kansas Publs., Mus. Nat. Hist.

was caught on spinning tackle, with doughballs for bait. The
second specimen was a juvenile taken by seining one mile below
Station C-5 on September 22.

Ictiobiis cyprinella ( Valenciennes ) : Station G-2.
Two Juvenal bigmouth buffalo were taken in a shallow pool,
along with several juvenal smallmouth buffalo.

Moxostoma aureolum pisolabrttm Trautman and Moxostoma
carinatum ( Cope ) : SBS.

Two specimens of Moxostoma aureolum pisolabrum (K. U.
242-243) and one specimen of Moxostoma carinatum (K. U. 223)
were taken from an unspecified locality on Elk River in Elk County
on July 11, 1912. There are no other records for any of these fish
in the collection area. M. aureolum pisolabrum has been taken in
recent years in eastern Kansas (Trautman, 1951:3) and has been
found as far west as the Chikaskia drainage in northern Oklahoma
by Moore and Buck (1953:21). That occasional northern redhorse
enter the larger rivers of the area here reported on seems probable.

M. carinatum has been reported only a few times from Kansas.
The only recent records are from the Verdigris River (Schelske,
1957:39). Elkins (1954:28) took four specimens of M. carinatum
from cutoff pools on Salt Creek in Osage County, Oklahoma, in
1954. This recent record suggests that occurrences in southern Kan-
sas are probable,

Moxostoma erythrurum ( Rafinesque ) : Stations G-5, G-7, G-10,
G-12, C-4, C-5, C-6, C-8, C-10, C-11, C-12, C-13, C-15, E-1, E-2,
E-4 (C-131, C-133, C-136).

The golden redhorse was common in several of the streams sur-
veyed, and utilized the upland parts of streams more extensively
than any of the other catostomids occurring in the area. M. ery-
thrurum and Ictiobus hubalus were taken together at only two sta-
tions. In no case was 7. bubalus taken from a tributary of Grouse
Creek or of Big Caney River. In contrast M, erythrurum reached
its greatest concentrations in such habitat, although it was always
a minor component of the total fish population. Stations C-5 and
E-2 were the lowermost environments in which this redhorse was
taken.

The largest relative number of golden redhorse was found at
station G-12 on Crab Creek where 7.5 per cent of the fishes taken
were of this species. This station consisted of intermittent pools
averaging one foot in depth. Bottoms were bedrock and rubble
and the water was clear and shaded. The fish were consistentlv

Fishes of Arkansas River Basin 365

taken in the deeper, open part of the pool where aquatic vegetation,
which covered most of the pool, was absent.

Another station at which M. enjthrunun was abundant was C-12
on Cedar Creek. Here a long, narrow, clear pool was the habitat,
with average depth of 17 inches, and bottom of bedrock.

Minytrema melanops ( Rafinesque ) : Stations G-10, C-4, C-12,
E-1.

Occurrences of the spotted sucker were scattered. At stations
C-4 and G-10 single specimens were taken. At station E-1 (July 9)
one specimen was taken at the mouth of a small tributary where
water was turbid and quiet. This specimen (K. U. 3708) was the
largest (9?8 inches total length) found, and possessed pits of lost
tubercles.

Cyprinus carpio Linnaeus: Stations A-1, W-1, W-2, W-3, W-4,
G-3, G-4, G-6, G-8, C-3, C-5, E-4.

Carp were taken most often in downstream habitat. No carp
were taken above station C-5 on Big Caney River.

The earliest date on which young were taken was July 7, when
46 specimens, approximately /2 inch in total length, were taken from
the Walnut River at station W-1. The small carp showed a prefer-
ence for small shallow pools; adults were found in deeper pools.

Hybopsis aestivalis tetranemus (Gilbert): Station A-3.

Only one specimen of tlie speckled chub was taken. The species
has been recorded from nearby localities in the Arkansas River
and its tributaries both in Kansas and Oklahoma. Its habitat seems
to be shallow water over clean, fine sand, and it occurs in strong
current in mid-channel in the Arkansas River. Suitable habitat
does not occur in other parts of the area covered by this report.

Notropis blennius (Girard): Stations A-1, A-2, A-3.

The river shiner was taken only in the Arkansas River and in
small numbers. In all instances N. blennius was found over sandy
bottom in flowing water. Females were gravid at station A-1 on
June 14. To my knowledge there are no published records of this
shiner from the Arkansas River Basin in Kansas. In Oklahoma this
species prefers the large, sandy streams such as the Arkansas River.
Cross and Moore (1952:403) found it in the Poteau River only
near the mouth.

Notropis boops Gilbert: Stations G-5, G-7, C-3, C-5, C-8, C-9,
C-10, C-11, C-12, C-15, C-16, E-4, E-5, M-1, M-2.

Widespread occurrence of the bigeye shiner in this area seems
surprising. Except for this area it is known in Kansas only from

366 University of Kansas Publs., Mus. Nat. Hist.

the Spring River drainage in the southeastern corner of the state
(Cross, 1954b: 474). N. hoops chose habitats that seemed most
nearly like Ozarkian terrain. The largest relative number of bigeye
shiners was taken at C-11 in a clear stream described in the discus-
sion of Notropis ruheUus. At this station IV. hoops comprised 14.11
per cent, and N. hoops and N. ruheUus together comprised 24.78
per cent of all fish taken.

At station G-7 on Grouse Creek the percentage of N. hoops was
7.15. Here, as at station C-11, water was clear. At both stations
MyriophijUum heterophylhim was abundant and at G-7 Nehimho
lutea was also common. At G-7 N. hoops seemed most abundant
in the deeper water, but at C-11 most shiners were found in the
shallower part of a large pool.

Two other collections in which N. hoops were common were
from Spring Creek. It is a small, clear Flint Hills brook running
swiftly over clean gravel and rubble. It had, however, been inter-
mittent or completely dry in its upper portion throughout the
winter of 1955-'56 and until June 22, 1956. In collections at C-15
on June 28, N. hoops formed 6.5 per cent of the fish taken. Farther
upstream, at C-16 on July 9, in an area one mile from the nearest
pool of water that existed prior to the rains of June 22, N. hoops
made up 7.2 per cent of the fish taken.

In streams heading in the hilly area of western Elk County, the
relative abundance of Notropis hoops decreased progressively down-
stream. On upper Elk River percentages were lower than on up-
per Grouse Creek and upper Big Caney River.

Hubbs and Lagler (1947:66) characterize the habitat of this
species as clear creeks of limestone uplands. There are numerous
records of the bigeye shiner from extreme eastern Oklahoma. It
has been reported as far west as Beaver Creek in Osage County,
Oklahoma. Beaver Creek originates in Cowley County, Kansas,
near the origin of Cedar Creek and Crab Creek. Drought had left
a few pools of water in Beaver Creek in Kansas at the time of my
survey. The fish-fauna seemed sparse and N. hoops was not among
the species taken. Of interest in considering the somewhat isolated
occurrence of the bigeye shiner in the Flint Hills area of Kansas is a
record of it by Ortcnburger and Hubbs (1926:126) from Panther
Creek, Comanche County, Oklahoma, in the Wichita Mountain
area of that state.

Notropui huchanani Meek: Stations G-1, E-4 (C-1.31).

At station G-1 the ghost shiner was taken in small numbers in the
shallow end of a long pool (150x40 feet.) The three individuals
taken at station E-4 were in an isolated pool (50x510 feet) aver-

Fishes of Arkansas River Basin 367

aging 1/2 feet in depth. Water was turbid, and warm due to lack
of shade.

The habitat preferences of this species and of the related species
N. volucellus have been described as follows by Hubbs and Orten-
burger (1929b: 68): "It seems probable that volucellus when oc-
curring in the range of buchanani occupies upland streams, whereas
buchanani is chiefly a form of the large rivers and adjacent creek
mouths." The results of this survey and impressions gained from
other collections, some of which are unpublished, are in agreement
with this view. A collection on the Verdigris River at Inde-
pendence, Kansas, directly downstream from the mouth of the
Elk River, showed N. buchanani to be common while N. volucellus
was not taken. At station E-5 upstream from E-4, however, N.
volucellus was taken but N. buchanani was not found.

In the upper Neosho basin. Cross (1954a:310) took N. volucellus
but not N. buchanani. Other collections have shown N. buchanani
to be abundant in the lower Neosho River in Kansas. Moore and
Paden ( 1950:85) observe that N. buchanani was found only near the
mouth of the Illinois River in Oklahoma and was sharply segregated
ecologically from N. volucellus that occupied a niche in the clear
main channels in contrast to the more sluggish waters inhabited by
N. buchanani.

Notropis camurus ( Jordan and Meek ) : Stations C-3, C-4, C-5,
C-6, C-7, C-8, C-9, C-10, C-11, C-12, C-13, E-1, E-5 (C-131).

Highest concentrations of the bluntface shiner were found close
to the mouths of two tributaries of Big Caney River: Rock Creek
and Otter Creek. On Rock Creek (Station C-4) this shiner was
abundant in a shallow pool below a riffle where water was flowing
rapidly. Many large males in breeding condition were taken
(June 3). The species formed 20.2 per cent of the fish taken.

On Otter Creek (Station C-13) the species was common in shal-
low bedrock pools below riffles. It formed 12.1 per cent of the fish
taken.

At station C-5, N. camurus was characteristically found in an area
of shallow pools and riffles. At station C-10 it was found in clear
flowing water over rubble bottom and in small coves over mud
bottom. At C-11 (July 26) N. camurus was taken only in one
small pool with rapidly flowing water below a riffle. In this pool
N. camurus was the dominant fish. At station C-12, on April 2,
N. camurus was abundant in the stream, which was then clear and
flowing. On August 24, it was not taken from the same pool, which
was then turbid and drying.

The frequent occurrence of this species in clear, flowing water

368 Unrtersity of Kansas Publs., Mus. Nat. Hist.

seems significant. Cross (1954a:309) notes that the bluntface
sliiner prefers moderately fast, clear water. Hall (1952:57) found
N. caniurus onh' in upland tributaries east of Grand River and not
in lowland tributaries west of the river. Moore and Buck ( 1953:22)
took this species in the Chikaskia River, which was at that time a
clear, flowing stream. They noted that in Oklahoma it seems to be
found only in relatively clear water.

N. camurus did not seem to ascend the smaller tributaries of Big
Caney River as did N. riibelhis and N. hoops even when these
tributaries were flowing,

Notropis deliciosus missuriensis (Cope): Stations A-1, A-2, A-3,
W-1, W-2, W-3 (C-136).

Sand shiners seemed to be abundant in the Arkansas River, rare
in the Walnut River and absent from other streams surveyed. This
shiner was most abundant in shallow, flowing water in the Arkan-
sas River; in backwaters, where Gambusia affinis prevailed, N.
deliciosus formed only a small percentage of the fish population.

Notropis girardi Hubbs and Ortenburger: Stations A-2 and A-3.

At station A-2 the Arkansas River shiner made up 14.6 per cent
of all fish taken. At A-2, it was found only in rapidly-flowing
water over clean sand in the main channels. It was absent from
the shallow, slowly-flowing water where N. deliciosus missuriensis
was abundant. At A-3 IV. girardi made up 22 per cent of the total
catch, and again preferred the deeper, faster water over clean-swept
sand. Failure to find N. girardi at station A-1 is not understood.

Females were gravid in both collections (August 25 and 27).
In neither collection were young-of-the-year taken. Moore
(1944:210) has suggested that N. girardi requires periods of high
water and turbidity to spawn. Additional collecting was done at
station A-3 on December 22, 1957. A few adults were taken in
flowing water but no young were found.

In this area, N. girardi showed no tendency to ascend tributaries
of the Arkansas River. Not far to the west, however, this pattern
changes as shown by Hubbs and Ortenburger (1929a:32) who
took this fish at seven of ten stations on the Cimarron, Canadian,
and Salt Fork of the Arkansas. N. girardi was taken only in the
lowermost stations on both Stillwater Creek (Cross, 1950:136) and
the Chikaskia River (Moore and Buck, 1953:22). In the next major
.stream west of the Chikaskia, the Medicine River, N. girardi seems
to occur farther upstream than in the Chikaskia. (Collection C-5-51
by Dr. A. B. Leonard and Dr. Frank B. Cross on Elm Creek near
Medicine Lodge on July 20, 1951. )

Fishes of Arkansas River Basin 369

Notropis lutrensis (Baird and Girard): Stations A-1, A-2, W-1,
W-2, W-3, W-4, G-1, G-2, G-4, G-5, G-8, G-9, G-10, G-11, G-12,
G-13, G-14, G-15, G-16, B-1, B-2, B-3, C-1, C-2, C-3, C-4, C-5,
C-6, C-9, C-10, C-11, C-12, C-13, C-14, M-1, E-1, E-2, E-4, E-7
(E&F, C-131, C-133, C-136).

The red shiner was taken in every stream surveyed. The relative
abundance seemed to be greatest in two types of habitat which
were separated geographically. The first habitat was in large
rivers such as the Arkansas and Walnut. In the Arkansas River the
red shiner consistently made up 20 per cent to 25 per cent of the
catch. On the Walnut River percentages ranged from 10 per cent
( station W-3 ) to 45 per cent ( station W-2 ) ,

The second habitat in which numbers of N. lutrensis reached
high proportions was in the upper parts of the most intermittent
tributaries. At the uppermost station in Silver Creek this species
formed 30 per cent of the fish taken. In Crab Creek the following
percentages were taken in six collections from mouth to source:
20.6%, 26.1%, 25%, 85%, 14.6%, and 1%. In the mainstream of Grouse
Creek the highest percentage taken was 19.27 near the mouth at
station G-1. In middle sections of Grouse Creek this species was
either absent or made up less than 2 per cent of the fish taken.

At no station on Big Caney River was the red shiner abundant.
The smallest relative numbers were found at upstream stations, in
contrast to collections made on tributaries of Grouse Creek. This
distributional pattern possibly may be explained by the severe con-
ditions under which fish have been forced to live in the upper
tributaries of Grouse Creek. Water was more turbid, and pools
were smaller than in Big Caney. These factors possibly decimate
numbers of the less hardy species permitting expansion by more
adaptable species, among which seems to be N. lutrensis. In the
upper tributaries of Big Caney River conditions have not been so
severe due to greater flow from springs and less cultivation of the
watershed in most places. Under such conditions N. lutrensis
seems to remain a minor faunal constituent.

Notropis percobromus (Cope): Stations A-1, A-2, W-1, W-2,
W-3, G-1.

At station W-1 the plains shiner constituted 20 per cent of the
fish taken. The river was flowing rapidly with large volume at the
time of this collection, and all specimens were taken near the bank
in comparatively quiet water over gravel bottom. At station W-3,
below Tunnel Mill Dam at Winfield, N. percobromus comprised
18.7 per cent of the fish taken, second only to Lepomis humilis in

o

70 UNn'ERSiTY OF Kansas Publs., Mus. Nat. Hist.

relative abundance. Immediately below the west end of the dam,
plains shiners were so concentrated that fifty or more were taken
in one haul of a four-foot nylon net. The amount of water over-
flownng the dam at this point was slight. Water was shallow
(8-12 inches) and the bottom consisted of the pitted apron or of
fine gravel. At the east end of the dam where water was deeper
(1-3 feet) and the flow over the dam greater, large numbers of
Lepomis humiJis were taken while 'N. percobromus was rare.

In the Arkansas River smaller relative numbers of this shiner
were obtained. At station A-2, it formed 4.68 per cent of the total.
At this station N. percobromus was taken with N. lutrensis in water
about 18 inches deep next to a bank where the current was sluggish
and tangled roots and detritus offered some shelter.

At station G-1 on Grouse Creek the plains shiner made up 7.68
per cent of the fish taken. The habitat consisted of intermittent
pools with rubble bottoms at this station, which was four miles
upstream from the mouth of the creek. The plains shiner seems
rarely to ascend the upland streams of the area.

Notropis rubellus (Agassiz): Stations C-3, C-5, C-6, C-7, C-8,
C-10, C-11, C-12, C-13, C-14 (J&J).

No fish in these collections showed a more persistent preference
than Notropis rubellus for clear, cool streams. All collections of
the rosyface shiner were in the Big Caney River system, but at only
four stations in this system was it common. At station C-11 the
highest relative numbers (10.6 per cent) were obtained. This site
possessed the most limpid water of any station on the mainstream
of Big Caney. Aquatic plants (Myriophyllum heterophyllum and
Potamogeton nodosus) were common. Other fishes that fliourished
at this station were N. boops, N. camurus, Campostoma anomalum,
and Etheostoma s])ectabile. The water temperature was 86° at
surface and 80° at bottom whereas air temperature was 97°.

N. rubellus was common at all stations in Otter Creek, the clear,
upland character of which has been discussed. In May and June
only adults were found. On September 1, examination of several
pools in upper Otter Creek revealed numerous young-of-the-year
in small spring-fed pools.

Literature is scarce concerning this shiner in Kansas. Cross
(1954a:308) stated that it was abundant in the Soutli Fork of the
Cottonwood River and was one of those fishes primarily associated
with the Ozarkian fauna, rather than with the fauna of the plains.
Elliott (1947) found N. rubellus in Spring Creek, a tributary of

Fishes of Arkansas River Basin 371

Fall River which seems similar to Otter Creek in physical features.
Between the Fall River and Big Caney River systems is the Elk
River, from which there is no record of the rosyf ace shiner. Perhaps
its absence is related to the intermittent condition of this stream at
present. The Elk River is poor in spring-fed tributaries, which seem
to be favorite environs of the rosyface shiner.

N. rubellus was taken by Minna Jewell and Frank Jobes in Silver
Creek on June 30, 1925 ( UMMZ 67818 ) . The shiner was not found
in any stream west of the Big Caney system in my collections.

In Oklahoma, Hall (1952:57) found N. rubellus in upland tribu-
taries on the east side of Grand River and not in the lowland
tributaries on the west side, Martin and Campbell (1953:51)
characterize N. rubellus as preferring rifHe channels in moderate
to fast current in the Black River, Missouri. It is the only species
so characterized by them which was taken in my collections.
Moore and Paden (1950:84) state "Notropis rubellus is one of the
most abundant fishes of the Illinois River, being found in all habi-
tats but showing a distinct preference for fast water . . ."

Notropis topeka ( Gilbert ) : Two specimens ( formerly Indiana
University 4605) of the Topeka shiner labeled "Winfield, Kansas"
are now at the University of Michigan Museum of Zoology. Col-
lector and other data are not given. Evermann and Fordice
(1886:185) noted that two specimens of N. topeka were taken from
Sand Creek near Newton in Harvey County, but do not list it from
Cowley County near Winfield. They deposited their fish in the
museum of Indiana University.

Notropis umbratilis (Girard): Stations G-1, G-3, G-4, G-7, G-8,
G-9, G-12, G-14, B-2, B-3, C-1, C-2, C-3, C-4, C-5, C-6, C-7, C-8,
C-9, C-10, C-11, C-12, C-13, C-14, C-15, C-16, E-1, E-2, E-4, E-5,
M-1, M-2 (J&J, C-131, C-132).

The redfin shiner flourished in all the streams surveyed except
the Arkansas and Walnut Rivers. N. umbratilis has been found in
upland tributaries of the Walnut River, some of which originate in
terrain similar to that in which Elk River, Big Caney River, and
Grouse Creek originate. ( Collection C-26-51 by Cross on Durechon
Creek, October 7, 1951.) This suggests downstream reduction in
relative numbers of this species, a tendency which also seemed to
exist on both Big Caney River and Grouse Creek. N. umbratilis
was the most abundant species in Big Caney River except at the
lowermost stations where it was surpassed in relative abundance by
N. lutrensis and Gambiisia affinis.

372 University of Kansas Publs., Mus. Nat. Hist.

N. umhmtilis was a pool-dweller, becoming more concentrated
in the deeper pools as summer advanced. In May and early June,
large concentrations of adult N. timbratilis were common in the
shallow ends of pools together with N. ntbeUtis, N. hoops, Pirne-
phales nototus, and Timephales teneUus. By July and August,
only young of the year were taken in shallow water, and adults
were scarcely in evidence.

Notropis volucellits ( Cope ) : Stations G-5, G-8, C-3, C-5, C-7,
C-8, C-9, C-10, M-1, E-4, E-5.

The mimic shiner was a minor element in the fauna, 2.02 per cent
at station C-5 being the largest percentage taken. In the Big
Caney River system N. vohtceUiis was taken only in the main
stream. In the Grouse Creek drainage it was found at two stations
in the upper part of the watershed, where water is clearer, gradient
greater, and pools well-shaded and cool.

In the Elk River the mimic shiner was taken only in the upper
part of the main stream. The dominant shiner in situations where
N. volucellus was taken was, in all cases, N. timbrafilis. Elliott
(1947) found N. volucellus in Spring Creek, a tributary of Fall
River. Farther north in the Flint Hills region, N. volucellus was
reported by Cross ( 1954a: 310).

Notemigonus crijsoleucas (Mitchell): Station W-5.

This isolated record for the golden shiner consisted of nine
specimens collected on June 6 in Timber Creek, a tributary of the
Walnut River. Most of the creek was dry. N. crtjsoleucas was
taken in one pool with dimensions of 8 feet by 4 feet with an aver-
age depth of 4 inches. This creek is sluggish and silt-laden, even
under conditions of favorable precipitation. Hubbs and Ortcn-
burger (1929b:89) observed that the golden shiner prefers sluggish
water. Hall (1952:58) took the golden shiner only in the low-
land tributaries west of Grand River and not east of the river in
upland tributaries.

Phenacobius mirabilis Girard: Stations W-3, C-3.

In no case was the suckermouth minnow common; it never com-
prised more than 1 per cent of the fish population.

Pimephales notatus (Rafinesque): Stations W-4, G-5, G-7, G-9,
G-12, G-13, B-3, C-1, C-2, C-3, C-4, C-5, C-6, C-7, C-8, C-9, C-10,
C-11, C-12, C-13, C-14, C-15, C-16, C-17, C-18, M-1, M-2, E-1, E-2,
E-4, E-5, E-7 (J&J, C-131, C-132, C-133).

This was much the most abundant of the four species of
Pimephales in this area. It was taken at 33 stations as compared
with 10 for P. tcnellus, 8 for P. promelas, and 3 for P. vigHax.

Fishes of Arkansas River Basin 373

The bluntnose minnow was taken almost everywhere except in
the main stream of the Arkansas and Walnut rivers and in lower
Grouse Creek. P. notatus seemed to prefer clearer streams of the
FHnt Hills part of my area. There was a marked increase in per-
centages taken in the upland tributaries of both Caney River and
Grouse Creek. In the Elk River, too, higher concentrations were
found upstream.

The highest relative numbers of bluntnose minnows were taken
at station G-12 on Crab Creek, station C-12 on Cedar Creek and
station C-16 on Spring Creek. At G-12, this minnow was abundant
in the deeper isolated pools. Males in breeding condition were
taken on June 9. In Cedar Creek the population of bluntnose
minnows was observed periodically in one pool in which they were
dominant. This pool was 100 feet by 50 feet, shallow, and with
bedrock bottom. At its upper end, however, there was a small
area of heavily-shaded deeper water. Throughout the spring
bluntnose minnows were found in large schools in the shallow
area. As the summer progressed they were no longer there, but
seining revealed their presence in the deeper, upper end.

At station C-16 on Spring Creek on July 9 male P. notatus were
taken in extreme breeding condition, being light brick-red in color
and with large tubercles.

Pimephales tenellus (Girard): Stations G-1, C-2, C-3, C-5, C-6,
C-7, C-8, M-1, E-2, E-4 (C-131 C-133).

The mountain minnow was never taken far from the mainstream
of Big Caney, Middle Caney, or Elk River, In this respect it
differed from P. notatus, which reached large concentrations in the
small upland tributaries. On the other hand, P. tenellus was not
so abundant as P. vigilax in the silty larger streams. In no collection
was the mountain minnow common. The highest percentages were
2.4 per cent (Station C-5), and 2.1 per cent (Station C-7) on Big
Caney River. These stations consisted of clear, flowing water over
rubble bottoms. Males at C-7 (June 16) were in breeding
condition.

Moore and Buck (1953:23) reported finding this species among
rocks in very fast water rather than in the quiet backwaters fre-
quented by P. vigilax. Other records of the mountain minnow
from the Flint Hills indicate that it seeks areas of maximum gradient
and flow; in this distributional respect it is like Notropis camurus.
The two species are recorded together from other streams in this
region such as the Chikaskia (Moore and Buck, 1953:23), Cotton-
wood (Cross, 1954a:310), and Spring Creek, tributary of Fall
River (Elliott, 1947). It is conceivable that a preference for flow-

374 University of Kansas Publs.^ Mus. Nat. Hist.

ing water might explain its restriction to the medium-sized, less
intermittent streams in this area. The only tributary which the
species seemed to ascend to any extent was Otter Creek, which is
seldom intermittent downstream.

Pimephales vigilax perspicuus (Girard): Stations A-3, C-1, C-4.

The parrot minnow was found only in downstream habitats.
Collection C-4 (June 3) on Rock Creek was made about M mile
from the mouth of this tributary of Big Caney and the creek here
had almost the same character as the river proper. The presence
of other channel fishes such as Ictiohus huhalus indicates the down-
stream nature of the creek. Some males of P. vigilax in breeding
condition were taken in this collection.

At C-1, only one specimen was found in a turbid, isolated pool
with bedrock bottom. At A-1 only one parrot minnow was taken;
it was in deep, fairly quiet water near the bank.

Other collections outside the three-county area revealed the
following: In the Neosho River, several parrot minnows were
found in quiet backwaters and in shallow pools. In the Verdigris
River three were taken directly under water spilling over the dam
at this station, while others were found, together with P. promelas,
in the mouth of a small creek that provided a backwater habitat
with mud bottom.

Cross and Moore (1952:405) found this species only at stations
in the lower portion of the Poteau River. Farther west the minnow
may ascend the smaller sandy streams to greater distances. Moore
and Buck (1953:23) took parrot minnows at six of 15 stations on
the Chikaskia River and found the species as far upstream as
Drury, Kansas. Elliott (1947), in comparing the South Ninnescah
and Spring Creek fish faunas, found only P. vigilax and P. promelaa
on the sandy, "flatter" Ninnescah and only P. nofattis and P. tenellus
on Spring Creek, an upland, Flint Hills stream in Greenwood
County,

Pimephales promelas Rafinesque: Stations A-2, A-3, W-3, W-4,
G-9, B-1, M-1, E-4 (E&F, C-136).

Occurrences of the fathead minnow were scattered, but included
all streams sampled except Big Caney.

Three of the collections were in small intermittent streams where
conditions were generally imfavorable for fishes and in one instance
extremely foul. Two of these stations had turbid water and all
suffered from siltation.

In Middle Caney Creek the species was rare but in the Elk River

Fishes of Arkansas River Basin 375

(June 28) more than 100 specimens, predominantly young, were
taken. This station consisted of a large isolated pool with a variety
of bottom types. Water was turbid and the surface temperature was
high (93° F.). In different parts of the pool the following numbers
of specimens were taken in single seine-hauls : 15 over shallow bed-
rock; 35 over gravel (IM feet deep); 50 over mud bottom (1 foot
deep ) .

P. promelas was found also in the large, flowing rivers : Arkansas,
Walnut, Verdigris, and Neosho. The species was scarce in the
Arkansas River, and was found principally in muddy coves. In
the Walnut (W-3), this minnow comprised 7.65 per cent of the
fish taken and was common in quiet pools.

Campostoma anotnalum Rafinesque: Stations W-4, G-4, C-1, C-3,
C-5, C-6, C-7, C-8, C-9, C-10, C-11, C-12, C-13, C-14, C-15,
C-16, C-17, C-18, B-3 (E&F, C-131, C-136).

Although the stoneroller was found in most streams surveyed,
it was taken most often in the Big Caney system, where it occurred
at 16 of the 18 stations. In contrast, it was represented at only one
of 17 stations on Grouse Creek. High percentages were found in
three creeks — Cedar, Otter, and Spring. As noted above, these
streams are normally clear, swift and have steep gradients and
many rubble and gravel riffles. On these riffles young stonerollers
abounded. Station C-16 on Spring Creek typifies the habitat in
which this species was most abundant. The stream has an average
width of 10 feet and depth of a few inches. The volume of flow
was less than 1 cubic foot per second but turbulence was great.
Water was clear and the bottom was gravel and rubble. Following
rains in June, stonerollers quickly occupied parts of Spring Creek
(upstream from C-16) that had been dry throughout the previous
winter.

On April 2 many C anomalum and Etheostoma spectabile were
taken in shallow pools and riffles in an extensive bedrock-riffle area
on Cedar Creek near station C-12. Most of the females were gravid
and the males were in breeding condition. On June 6 these pools
were revisited. Flow had ceased and the pools were drying up.
Young-of-the-year of the two species were abundant, but only a few
mature stonerollers were taken. On August 24, prolonged drought
had drastically altered the stream and all areas from which stone-
rollers and darters had been taken were dry. Seining of other pools
which were almost dry revealed no stonerollers.

Collections on May 31, June 15, and June 16 in Otter Creek re-

376 University of Kansas Publs., Mus. Nat. Hist.

vealed large numbers of stonerollers. They were found in riffle
areas, in aquatic vegetation, and especially in detritus alongside
banks. Most of the specimens were young-of-the-year.

Anguilla bostoniensis (Le Sueur): An American eel was caught
by me in Grouse Creek in 1949.

Gambusia affinis (Baird and Girard): Stations A-1, A-2, A-3,
W-1, \V-2, W-3, W-4, W-5, G-1, G-2, G-3, G-4, G-5, G-7, G-8,
G-9, C-1, C-2, C-3, C-4, C-6, C-15, E-1.

Mosquitofish occurred widely but in varied abundance. Huge
populations were in the shallow sandy backwaters and cut-off pools
of the Arkansas River. In the shallow pools of several intermittent
streams such as station G-8 on Silver Creek this fish also flourished.

G. offinis was taken at every station in the Arkansas, Walnut and
Grouse systems except those stations on two upland tributaries of
Grouse Creek (Crab Creek and Grand Summit Creek). The
mosquitofish was not observed in the clear upland tributaries of
Big Caney, nor on upper Big Caney River itself in May, June, and
July. On September 3, however, Gambusia were taken at station
C-15 on Otter Creek and others were seen at station C-14 on the
same date.

Hubbs and Ortenburger (1929b:99) and Cross and Moore
(1952:407) observed that G. affinis usually was absent from small
upland tributaries, even though it was abundant in lower parts of
the same river systems.

Fundulus kansae (Garman): Stations A-2, A-3, Evermann and
Fordice as Fundulus zebrinus.

At station A-2, seven plains killifish were taken together with a
great many Notropis deliciosus and Gambusia affinis in a shallow,
algae-covered channel with slight flow and sand bottom. At sta-
tion A-3 many young killifish were taken in small shallow pools on
December 22. Fundulus kansae has been found in the lower part
of the Walnut River Basin, especially where petroleum pollution
was evident. Eastward from the Walnut River plains killifish have
not been taken.

Fundulus notatus (Rafinesque) : Stations B-1, G-1, G-2, G-3, G-4,
G-5, G-7, G-8, G-10, G-11, G-14, C-1, M-1, E-1, Evermann and
Fordice as JAjp^onectes notatus.

The black-banded topminnow was not taken in the Arkansas
River but was common in the Walnut and Grouse systems. It
was common also in Middle Caney, but in Big Caney and Elk River
it was taken only at the lowermost stations.

Fishes of Arkansas River Basin 377

This species did not seem to ascend far into smaller tributaries
of Grouse Creek. In Crab Creek it was taken at the lower two of
six stations and in Grand Summit Creek at the lower of two stations.

The highest relative numbers were taken at stations G-3 (17.5
per cent), G-4 (24 per cent), G-10 (25.75 per cent) and G-11 (41.52
per cent), on Crab Creek and Grouse Creek. Both upstream and
downstream from these stations, which were within five miles of
each other, the relative abundance dropped off sharply. The bot-
toms at these stations were mostly rubble and mud, and water was
turbid at three of the stations. At G-10 (June 24) and G-11 (July
16) young-of-the-year were abundant.

Ictalurus melas (Girard): Stations W-2, W-3, W-4, W-5, B-1,
B-2, B-3, G-1, G-2, G-3, G-4, G-5, G-8, G-9, G-11, G-12, G-13, G-14,
G-17, C-1, C-9, C-11, C-12, C-14, C-15, C-17, C-18, E-1, E-2, E-4,
E-5, E-6, N-1, Evermann and Fordice as Amekirus melas (C-133).

The black bullhead was taken at slightly more than half of the
stations, and probably was present at others. Larger numbers
were taken in Grouse Creek than in any other sti-eam system. In
many small, shallow pools in the Grouse Creek system young black
bullheads shared dominance with Gambusia affinis in the late sum-
mer. I. melas was also abundant in isolated pools at the extreme
upper ends of Crab Creek, Beaver Creek and Grand Summit Creek.
I. melas was most common in areas with silty bottoms. Tlie species
seemed scarce in the main stream of Big Caney River but was
common in some of its tributaries.

Pylodictis olivaris (Rafinesque): Stations A-3, G-1, C-5.

Flathead catfish were taken by angling at stations A-3 and C-5.
At station G-1 (September 5) a flathead catfish five inches long
was taken in the four-foot nylon net.

Ictalurus punctatus ( Rafinesque ) : Stations A-3, W-2, W-3, G-2,
C-5, E-4.

Channel catfish from stations W-3, A-3, and C-5 were taken on
hook and Hne. At station G-2 (August 29) twenty young-of-the-
year were seined from the shallow narrow end of a large pool. All
collections of both I. punctatus and F. olivaris were in the larger
streams surveyed.

Ictalurus natalis (LeSueur): Stations G-1, G-2, G-3, G-4, G-8,
G-11, C-12, C-14, C-15.

The yellow bullhead was taken at only 9 stations, compareH
with 33 stations for the black bullhead. /. natalis was represented

3_7079

378 University of Kansas Publs., Mus. Nat. Hist.

in 7 of 17 stations in the Grouse Creek system but in only 3 of 18
stations in the Big Caney system. Of the seven records from Grouse
Creek four were from the main stream. At every station where
yellow bullheads were taken, black bullheads were found also
and were abundant, usually several times more abundant than
7. natalis.

At G-11 on Crab Creek (July 16), 7. natalis made up 3.8 per cent
of the fish taken. All were young-of-the-year, existing in a tiny,
gravelly pool containing not more than five gallons of water, and
were the only fish present. Young yellow bullheads were also found
in small pools with gravel bottoms at station G-4 on September 7.

Labidesthes sicculiis (Cope): Stations G-1, G-2, G-3, G-7, G-10,
B-2, C-1, C-2, C-3, C-4, C-5, C-12, E-1, E-2, E-3, E-7, M-1 (E&F,
C-131).

The brook silversides was taken, sometimes abundantly, in all
stream systems except the Walnut and Arkansas. At station G-7
on July 8, 41.8 per cent of the fish taken were of this species. L. sic-
culus was most abundant in large pools where the bottom was pre-
dominantly bedrock and gravel. The highest concentrations were
in the mainstreams of Big Caney, Grouse, and Elk Rivers. Brook
silversides were taken rarely in the smaller tributaries of these
streams.

Percina phoxocephala ( Nelson ) : Stations C-2, C-3, C-5, G-1
(C-133).

Slenderhead darters were scarce, and were found only over
gravel bottoms. Specimens were taken from flowing and quiet
water, and from both shallow and deep water.

Larger numbers of P. phoxocephala were taken by the writer in
other collections made during 1956 on the Neosho and Verdigris
Rivers over bottoms of rubble or gravel. Restriction of this darter
to the larger streams follows a pattern observed by Cross
(1954a: 313) who noted it was absent from smaller riffles in minor
tributaries. Elliott (1947), however, took one specimen of P.
phoxocephala in Spring Creek, a tributary of Fall River.

Percina caprodes carbonaria ( Baird and Girard ) : Stations G-3,
G-4, G-7, G-12, C-5, C-6, C-7, C-9, C-12, C-13, C-14 (J&J, C-131,
C-133).

The logperch was generally distributed in the Caney, Elk, and
Grouse systems. This species usually comprised less than 1 per cent
of the fish taken; however, at station G-12 it formed 3.76 per cent of
the total.

Fishes of Arkansas River Basin 379

In many instances the logperch was taken over submerged gravel
bars, often along the edges of the larger pools. At 8 of 13 stations
where the logperch was taken, the golden redhorse was also found.
At every station where logperch were found, Notropis umbratilis
was taken and Pimephales notatus also occurred at all but three
of these stations.

Percina copelandi ( Jordan ) : Stations C-4, C-5, C-6, C-8, G-1
(C-131,C-133,J&J).

Channel darters were collected over bottoms of rubble or gravel,
both in flowing streams and in isolated pools. Although P. cope-
landi was found only in Big Caney River and at the lowermost
station on Grouse Creek ( G-1 ) in this survey, this species has been
taken previously from Elk River (K. U. 3463 and K. U. 3197) and
from Silver Creek. Notropis camurus occurred everywhere that
H. copelandi was found. In several instances the two species were
taken in the same seine-haul.

Etheostoma spectabile pulchellum ( Girard ) : Stations W-4, G-1,
G-4, G-5, C-6, C-9, C-11, C-12, C-13, C-14, C-15, C-16, C-17, C-18,
E-1, E-5. Evermann and Fordice as Etheostoma coeruleum (C-131,
C-132).

The habitat preferences of the orangethroat darter seemed similar
to those of Campostoma anomalum. There were sixteen stations
at which both species were taken, seven where only E. spectabile
pulchellum was taken and six where only C. anomalum was taken.
The largest relative numbers of both species were found in the
same small, clear upland tributaries of Big Caney River. On May
31, collections from riffles at station C-15 (upper Otter Creek) con-
sisted almost entirely of these two species. On September 1 at this
station the stream was intermittent, but even the tiniest pools
abounded with young darters and stonerollers.

Gravid females and males in breeding condition were taken in
riffles in Cedar Creek on April 2. During June numerous young
and adult orangethroat darters were taken in Cedar Creek, in
partly decayed leaves which lined the banks. On June 15 in Otter
Creek young darters were abundant in streamside detritus and in
clear, shallow, rubble riffles. At station C-11 a few darters were
taken on rubble riffles; however, large numbers were found in-
habiting thick mats of Potamogeton foliosus Raf., which grew in
shallow water. Many darters {Etheostoma spectabile pulchellum
and Percina phoxocephala ) were taken in September along gravelly
banks at stations C-2 and C-3 by disturbing small rocks and leaf-
litter along the shores. Young orangethroat darters seemed to seek

380 University of Kansas Publs., Mus. Nat. Hist.

out sheltered areas and in some cases were found in sluggish, even
foul, water (Stations W-4, B-1 and G-12). Moore and Buck
(1953:26) note that the orangethroat darter is able to thrive in
Oklahoma in rather sluggish and even intermittent waters which
reach quite high summer temperatures.

Unlike other darters taken in this survey, the orangethroat darter
was common to abundant at several stations and was found at a
great many more stations than any other darter. The comparatively
great tolerance of this species to varying habitats, suggested by this
survey, is also reflected by its widespread distribution in Kansas.

Micropterus salmoides salmoides (Lacepede): Stations B-1, G-4,
G-5, G-7, G-12, C-1, C-3, E-1, E-2, E-3.

Most of the largemouth bass taken were young-of-the-year. In
Big Caney River this species seemed rare, being found at only two
downstream stations compared with eight stations at which M.
punciulatus was taken.

Many ponds in the Flint Hills have been stocked with largemouth
bass. At present largemouth bass are frequently caught by hook
and line in Crab Creek (Station G-12); however, Mr. A. C. Metcalf,
who has fished this stream for approximately 45 years, states that he
took no bass in the creek prior to the building and stocking of large
ponds on nearby ranches.

Micropterus pimctulatus (Rafinesque): Stations C-4, C-5, C-6,
C-7, C-8, C-10, C-14, C-15, E-2, E-5 (C-133).

The spotted bass was taken only in tributaries of the Verdigris
River, where it seemed more numerous than the preceding species.
It has been reported from other Verdigris tributaries such as Fall
River (Elhott, 1947) and is common eastward from the Verdigris
Basin. A spotted bass (K. U. 3467) was taken by Cross on the
Little Walnut River in Butler County on April 5, 1955. This seems
to be the only record of this species from the Walnut River Basin
at the present time.

Pomoxis annularis (Rafinesque): Stations W-3, W-5, G-1, G-2,
G-5, G-10, G-11, G-12, C-1, C-2, C-4, C-5, C-6, M-1, E-1, E-2, E-4,
E-5 (C-136).

White crappie were found in almost all habitats and were taken
in all rivers except the Arkansas. The relative abundance of this
species was greater at downstream than at upstream stations on
Grouse Creek, Big Caney, and Elk River. Schools of young crappie
were frequently found and the factor of chance in taking or failing
to take a school of crappie prevented confident appraisal of
abundance. White crappie usually sought quiet waters. Often they

Fishes of Arkansas River Basin 381

were found in backwaters and many times schools were taken over
bottoms where mud and detritus had been deposited. It was not
uncommon to take Pomoxis annularis and Ictalurus melas in the
same seine-haul in such areas.

Pomoxis nigromaculatus (LeSueur): Station C-1.

Black crappie were taken in Otter Creek on May 29 and Septem-
ber 3. Several ponds in eastern Cowley County are stocked with
black crappie, but none was taken from streams into which these
ponds drain.

Lepomis cyanellus (Rafinesque): Stations W-3, W-4, W-5, B-1,
B-2, B-3, G-1, G-2, G-3, G-4, G-5, G-8, G-9, G-10, G-12, G-13, G-14,
G-15, G-16, G-17, C-1, C-2, C-3, C-4, C-5, C-6, C-7, C-8, C-9, C-10,
C-11, C-12, C-13, C-14, C-16, C-17, C-18, E-1, E-4, E-5, E-6, E-7,
M-2 (C-131, C-132, C-133, C-136, E&F).

The green sunfish was taken at 45 of 60 stations, which is the
greatest number recorded for one species. The only stream from
which it was not obtained was the Arkansas River. Green sunfish
constituted a minor but consistent part of the fauna in Big Caney
River except for some intermittent pools on small tributaries, where
it was high in relative abundance. It usually comprised approxi-
mately 4 per cent of the fish taken at stations on Grouse Creek. In
some intermittent bibutaries of Grouse Creek and Elk River percent-
ages also were high.

Funk and Campbell (1953:74) observed that L. cyanellus held a
definite but minor place in all collections made on the Black River
in Missouri. This pattern was also observed by the writer in col-
lections made on the Neosho and Spring Rivers in southeastern
Kansas. This seems to indicate that the Big Caney River popula-
tions (exclusive of the upstream stations in intermittent streams)
follow a pattern commonly found in southeastern Kansas and prob-
ably in the Ozark region,

Lepomis humilis (Girard): Stations A-3, W-2, W-3, W-4, W-5,
G-1, G-2, G-3, G-4, G-5, G-7, G-8, G-9, G-10, G-11, G-12, G-14, G-15,
C-1, C-2, C-3, C-4, C-5, C-6, C-7, C-8, C-9, C-10, C-11, C-12, C-13,
C-16, C-17, B-2, B-3, E-1, E-2, E-4, E-5, E-6, E-7, M-1. (C-131,
C-132, C-133, C-136, J&J, E&F.)

The orangespotted sunfish was found in every stream surveyed,
although only one specimen was taken from the Arkansas River.

The largest relative number of this species (44.6) was taken at
station G-1. Percentages at other stations on Grouse Creek and its
tributaries progressively dechned in an upstream direction.

In Big Caney River representation of L. humilis in collections

382 University of Kansas Publs., Mus. Nat. Hist.

varied from 1.56 per cent at station C-1 to 23.47 per cent at station
C-7. This SLinfish was usually the dominant species in collections
made from the Elk River, where the relative abundance ranged
from 10 to 30 per cent.

The orangespotted sunfish is widespread in Kansas and seems to
be a diagnostic constituent of the Plains Fauna. Moore and Buck
( 1953:26) found it "very common" in the Chikaskia River in Kansas
and Oklahoma. Cross (1950:140) noted that in Stillwater Creek it
seemed to be the most tolerant and consequently the most abundant
of the stream's centrarchids. Moore and Paden (1950:91) note
that L. hinnUis is most common in muddy waters and found in over-
flow pools, backwaters, and oxbow lakes. This species is frequently
found in farm ponds in the area surveyed, which further suggests a
wide range of habitat tolerance.

Lepomis megalotis breviceps ( Baird and Girard ) : Stations W-3,
W-4, W-5, B-1, B-2, G-1, G-4, G-5, G-7, G-8, G-9, G-10, G-11, G-12,
G-13 (all Big Caney River stations except C-18), E-1, E-2, E-3,
E-4, E-5, E-6, M-1, M-2 (C-131, C-132, C-133, J&J, E&F).

In Big Caney River the longear sunfish shared dominance with
the redfin shiner (Notropis iimbratilis) at almost every station.
The average of its relative abundance at all stations in the Big
Caney system was 16.5 per cent. It was also abundant at several
stations on Grouse Creek and made up 43.25 per cent of all fish
taken at station G-4.

Cross (1950:140) observed that L. megalotis breviceps increased
in Stillwater Creek probably as a result of clearer water and stabi-
lized water level.

In collections made west of the area treated here (Moore and
Buck, 1953:26; Elliott, 1947) the longear sunfish is less abundant
than in Big Caney River and Grouse Creek.

Lepomis macrochirns (Rafinesque) : Stations W-3, G-3, G-4, G-5,
C-3, C-5, E-1, E-2 (C-131, C-132, C-133).

The bluegill was, in all cases, a minor constituent in the fish
fauna. No clear pattern of habitat preference can be deduced. In
the Verdigris River at Independence (collection AM-53, August 22,
1956) bluegills were common in quiet pools and coves below a
low-water dam. Moore and Paden (1950:91) note that L. macro-
chirus prefers quiet waters and Hubbs and Lagler (1947:94) state
that it is "generally restricted to the quieter pools."

The bluegill is widely-stocked in impoundments of the area
treated here.

Fishes of Arkansas River Basin 383

Aplodinotus grunniens (Rafinesque): Stations C-4, E-2.

The dearth of stations from which the freshwater drum is re-
ported may indicate diflBculty in taking this species with seines,
rather than scarcity. Both collections were at downstream stations.
At station C-4 three half-grown drum were taken. Fishermen take
"drum" at least as far upstream as station C-5 on Big Caney River.
In the Elk River one specimen was taken in a 20-foot seine below
a dam at Elk Falls.

FISHES OF DOUBTFUL OR POSSIBLE OCCURRENCE

In addition to the species listed above, the following species
have been reported nearby and may occur within the area surveyed.

Lepisosteus prodiictiis (Cope) — This gar has not been reported
from Kansas. It has been taken at several points in the northern
half of Oklahoma and as far west as Canton Reservoir by Buck
and Cross (1951), A specimen of the spotted gar was taken by
Elkin (1954:28) in Salt Creek in Osage County, Oklahoma.

Polyodon spathula (Walbaum) — The paddlefish has never been
reported from the Arkansas River system in Kansas. Several reports
by fishermen were traced by the writer, but authentication was not
achieved. One mounted specimen was examined in a sporting
goods store in Arkansas City. This fish was said to have been
taken on the Arkansas River south of Arkansas City but information
on the date and method of capture were vague. Mr. Darrell Wheat
of Arkansas City reported taking four paddlefish below a dam at
Oxford, Kansas, in 1948 and 1949.

Hiodon alosoides (Rafinesque) — One specimen (K. U. 3095) of
the goldeye was taken in 1953 on the Arkansas River near Oxford
in Sumner County. Fishermen also report taking this fish occa-
sionally in the Walnut River in Cowley County.

Noturus flavus (Rafinesque) — The stonecat was taken in the
Verdigris system by R. D. Lindsay in 1911 (K. U. 2058) and more
recently by Cross in Montgomery County (C-120) and Schelske
(1957:46) in Wilson and Montgomery Counties. The close prox-
imity of these collection areas to lower portions of the Elk River
indicate probable occurrence in Elk River and other Verdigris
tributaries.

Noturus nocturnus (Jordan and Gilbert) — The freckled madtom
has been taken on all sides of the area studied making its occurrence
therein highly probable. This madtom has been taken in Beaver

384 University of Kansas Publs., Mus. Nat. Hist.

Creek in Osage County, Oklahoma (OAM 4771); from a tributary
of the Walnut River in Sedgwick County by Cross (1954); from
the Chikaskia River (Moore and Buck, 1953:24); and from several
localities on the Verdigris River (Schelskc, 1957:47).

Etheostoma cragini (Gilbert) — One Cragin's darter (K. U. 3470)
was taken by Cross in the Arkansas River near the Sumner-Cowley
county line (Sec. 25, T31S, R2E). Records of this darter are few
and widely scattered geographically. Several collections from north-
eastern Oklahoma are noted by Moore and Cross (1950:144).

Etheostoma whipplii (Girard) — Schelske (1957:38) reports the
redfin darter from the Verdigris River three miles southeast of
Benedict, Kansas. Dr. George Moore of Oklahoma A. & M. College
states that it has been taken in the Verdigris drainage in Oklahoma
at several locations.

Etheostoma zonale arcansanum (Jordan and Gilbert) — Two
banded darters (K. U, 3213) have been reported by Schelske
(1957:49) from Fall River near Neodesha, Kansas. Because a
tributary of Fall River enters Elk County its presence in this and
other Verdigris tributaries in the area seems possible. This darter
has been reported from only one other stream in Kansas, Shoal Creek
in Cherokee County, where it has been collected often.

Roccus chrysops (Rafinesque) — The white bass has been stocked
in Hulah Reservoir on Big Caney River in Oklahoma. To date it
has not been reported from the Big Caney in Kansas. White bass
are common in many reservoirs of Kansas and Oklahoma and have
been taken in rivers in both states. Mr. Clement Gillespie of Ar-
kansas City, Kansas Forestry, Fish and Game Commission wildlife
protector for the area, states tliat two hundred young of R. chrysops
were released in Grouse Creek several years ago under auspices of
the Commission. The fish has not been reported by fishermen since
that time to the knowledge of Mr. Gillespie or of the writer.

Lepomis microlophus (Gunther) — One redear sunfish was taken
on Salt Creek in Osage County, Oklahoma, by Elkin (1954:28).
Because this species has been stocked widely in Oklahoma its even-
tual occurrence in Kansas seems probable.

Chaenobryttus gulosus (Cuvier) — The warmouth has been taken
south of the collection area in Osage County on Salt Creek by Elkin
(1954:28).

FAUNAE COMPARISONS OF DIFFERENT STREAMS

The faunas of Elk River, Big Caney River, and Grouse Creek
were generally similar. These streams and most of their tributaries
originate in the same hilly area of eastern Cowley County and west-

Fishes of Aeka.nsas River Basin 385

em Elk and Chautauqua counties; their similarities and differences
have been pointed out.

The following species were taken in all of these streams:

Lepisosteus osseus Fundulus notatus

D'orosoma cepedianum Gamhusia affinis

Ictiobus bubalus Ictalurus melas

Moxostoina erythrurum Ictalurus punctatus

Minytrema melanops Etheostoma spectabile

Cyprinus carpio Percina caprodes

Campostoma anomalum Micropterus salmoides

Notropis boops Vomoxis annularis

Notropis lutrensis Lepomis cyanellus

Notropis umbratilis Lepomis humilis

Notropis volucellus Lepomis megalotis

Pimephales notatus Lepomis macrochirus

Fimephales tenellus Labidesthes sicculus

No species was found in Elk River to the exclusion of Big Caney
and Grouse Creek. Fish taken exclusively in Grouse Creek were
Ictiobus cyprinella at station G-2 and Notropis percobromus at sta-
tion G-1. The following species were taken only in Big Caney
River: Ictiobus niger, Notropis rubellus, Phenacobius mirabilis,
Pimephales vigilax, and Pomoxis nigromaculatu^.

Notropis buchanani and Pimephales promelas were taken in
Grouse Creek and Elk River, but not in Big Caney River, although
the watershed of Big Caney lies largely between these two streams.
Three species, Notropis camurus, Micropterus punctulatus, and
Aplodinotus grunniens, were found in Elk River and Big Caney but
not in Grouse Creek. Ictalurus natalis, Pylodictis olivaris, and Per-
cina phoxocephala were taken in Big Caney River and Grouse Creek
but not in Elk River. Percina copelandi was taken by Cross on Elk
River in 1954 and 1955 (K. U. 3464 and K. U. 3197).

Forty species were taken in Big Caney River, 35 in Grouse Creek
and 31 in Elk River. Collections were made from only six stations
on Elk River as compared with 18 from Big Caney and 17 from
Grouse Creek.

Twenty-four species were taken in the Walnut River system, only
one of which (Notemigonus crysoleucas) was taken exclusively
there.

In the Arkansas River 18 species were found, four of which did
not occur elsewhere. These were Hybopsis aestivalis, Notropis
blennius, N. girardi, and Fundulus kansae.

Table 5 Hsts the number of stations in each of the streams sur-
veyed from which each species was taken.

386

Unrtrsity of Kansas Publs., Mus. Nat. Hist.

Table 5. — Species of Fishes Collected and Number of Stations in Each
Stream System at Which Each Species Was Found.

Total number
of stations

Arkansas

River
3 stations

Walnut
River
5 stations

Grouse

Creek
17 stations

Big Caney

River
18 stations

>- 2

cj a

> o

HO

Middle

Caney
2 stations

Beaver

Creek
3 stations

L. osseus

1

Seen

2

3

1
1
1

3
1
1

2

1

6
3
1
4

Seen
2

D. cepedianum

Carpiodes carpio

/. bubahis

1

3

/. cyprinella

/. niqer

2

10
3
2

14

M. erythrurum

4

1
4
1

3

1
1
2

M. melonops

Cyprinus carpio

1

4
1

C. anonialutn

1

H. aestivalis

1
2

N. blennius

A^. boops

2
1

14
13'

2

1
2

2

N. biichanani

N . caiuurus

A'^. deliciosus

3
2
3

3

N . girardi

N. luirensis

N. rubellus

4

13

14
11

5

1

3

N. percobromus

3

3

1
8
2

N . umbratilis

18
5

4
2

2

1

2

A'^. voluceUus

A'^. crysoleucas

1

2

1
1
2

H. placila

3

P. tnivabilis

1

1

18
.. .^..

7
1

P. notatus

5

1

1
1

2
1

1
2

1

P. promelas

2

1

1

P. vigilax

P. tenellus

1

10

F. notatus

4

1

F. kansae

2
3
1

G. affinis . .

5
4

8
12
6
1
1
4
1
1
5
4

8

9

3

5

1

17

5

4

8

2

7

1

1

17

17

18

3

1

7

1
5

I. me! as

3

I natalis

I . punctalus

1
1

2

"l"

1

P . olivaris

E. spectabilr

2

1

P. copelandi

P. phnxocephala

P. caprodes . . .

1
3
1
4

AT. salmnides

1

M . pxmctidalxis

P. annularis

2

7

1

P. nigromaculatus

L. cyancllus

3
4
3

1

14

13

0

3

5
(i
()
2

T

4

1

1
2

3

L. hiimilis

1

2

L nif'oalotis

2

Li. macrorhirus

A. grunniens

L. sicculus

5

1

1

Fishes of Arkansas River Basin 387

DISTRIBUTIONAL VARIATIONS WITHIN THE
SAME STREAM

An analysis of faunal variations in different parts of the same
stream system was made for Big Caney River and Grouse Creek.
Collecting was more extensive in these streams, and sampling was
done over a wider range of habitat, than in the Arkansas and Wal-
nut rivers.

The fish taken in the first five seine hauls at each station were
counted and the number of each species was recorded as a per-
centage of the total number of fish taken. These percentages were
calculated for the main stream and for each tributary in an attempt
to discern possible intra-stream faunal patterns. In Table 6 lower,
middle, and upper segments of each stream have been segregated
and the average of all stations within each segment is shown.

The results are subject to several sources of error, some of which
are discussed below:

( 1 ) Seining techniques could not be entirely standardized. One
station might present a series of long narrow riflBes and narrow,
shallow pools in which only a small seine could be used effectively;
another station might consist of a large, deep, isolated pool in which
a larger seine was needed for effective sampling. In practice, the
five seine hauls were made with any of several seines ranging from
ten to twenty feet in length.

(2) Seines are species-selective, due partly to the preference of
certain fishes for special habitat niches. Fishes that are often found
under stones or in weedy pools require special collecting techniques
and frequently were not represented in the initial five hauls. If
work subsequent to the first five hauls indicated that such fish were
a prominent part of the fauna at a particular station, these results
were considered before percentages were calculated.

(3) Temporal variations occur in populations at the same sta-
tion. There were both seasonal and diurnal differences in relative
numbers of species taken in these collections. This was noted
especially at station C-5 where collecting was done both at night
and by day. Spawning by certain species during the course of the
study complicated estimates of their relative abundance.

( 4 ) In tabulating percentages of fishes obtained an arbitrary ele-
ment is often unavoidable in deciding whether a station, especially
a station on a tributary, should be considered as part of the lower,
middle, or upper segment of a river system.

Despite these disadvantages it is felt that table 6 has factual basis
permitting some reliable interpretation.

388

Unrtrsity of Kansas Publs., Mus. Nat. Hist.

Table 6. — Relattve Abltndance in Per Cent of Fishes in Collections

From Three Stream Segments,

Big Caney River

G

rouse Creek

Lower

Middle

Upper

Lower

Middle

Upper

Li. osseus

.7
.3
.06
.6

.5

.6

.02
.02

D. cepedianum

Carpiodes carpio

/. bubalus

1.0

1.4

.1

.45

/. cyprinella

/. niger

.01

.2

.1

.7
.6
.6

M. erythrurum

M. melanops

1.1
.01

1.0

.03

.5
.1
.2
.1
1.3

1.1

Cyprinus carpio

C. anomalum

N. boops

1.3

""'.bi

5.9
.6

18.0
5.1

N, buchanani

N. camurus

6.4
8.8

5.5
1.0

.4
.5

N. lulrensis

6.4
1.1

11.4

15 2

N. percobromus

N. rubellus

.4
17.6
.3
.3
3.5
.8

i.4

28.3

.4

3.9
15.4

N. umbralilis

N. volucellus

2.5

3.9
.3

5.5

P. mirabilis

P. notalus

5.7

13.0

.9

6 6

P. vigilax

P. promelas

2.9

P. tenelliis

.7

14.6

.1

.9

.5
.4

""'4'"

.01
20.8
6.6
5.6
.5
.01
.4
.4
.01
.1
.2

G. affinis

10.2

17.2

2.3

.8

1.0

F. nolntus

1 4

I. melas

2.2

2.4
.5

18 0

/. natalis

P. olivaris

.01

.3

1.9

.8

.1

.4

.00

.5

3.9

3.4

10.6

12.4

.3

.1

7.1

/. punctatus

E. spectabile

4.9
.1

18.0

.3

.3

P. copelandi

P. phoxocephala

P. cavrodes

.6

.2

.2
1.1

4

M. sahnoides

.3

M. punctidalus

1.7

.8

.8

13.1

22.3

.4

"6'6'
1.8
12.0

P. annularis

2.9

5.2

31.4

3.6

.2

4.2

1.8

17.7

14.0

1.3

.3

L. cyanellus

30.5

L. huniilis

14.8

L. megalotis

1.7

L. macrochirus

A. griinniens

L. sicculus

1.6

.4

7.7

10.2

Big Caney River
The "lower segment" of Big Caney River is immediately upstream
from Hulah Reservoir, and is not the lowermost portion of the
entire river basin, but merely the lower part of the river in the area
studied. A conspicuous characteristic of the lower segment was
the general restriction of the deep-bodied suckers and the carp to

Fishes of Arkansas River Basin 389

this part of the stream. Other fishes that were most common in the
lower section were Pimephales vigilax, Vercina phoxocephala, Gam-
busia affinis, and Aplodinotus grunniens. Labidesthes sicculus
and Lepisosteus osseus ranged into the middle section of the stream,
but were present in larger numbers downstream. Ictalurus punc-
tatus, Pomoxis annularis, and Lepomis macrochirus were taken
chiefly in downstream habitats; however, stocking has confused the
distributional pattern of these species. Notropis lutrensis, although
found throughout the system, progressively declined in numbers
taken in the middle and upper sections. Approximately 18 species
were usually taken in downstream collections.

No species were found exclusively in the middle section of the
Big Caney system. Micropterus pimctulatus, Notropis umbratilis,
and Lepomis megalotis tended to be most common in the middle
section of the main stream. These three species were taken to-
gether at stations C-5, C-6, C-8, and C-10.

The upper section yielded no species that did not occur also in
another section. Fishes most abundant in the upper section in-
cluded: Campostoma anomalum, Etheostoma spectabile, Notropis
boops, Notropis rubellus, Pimephales notatus, and Lepomis cyanel-
lus. Ictalurus natalis also seemed more common upstream than in
lower parts of the basin.

Campostoma anomalum was one of the most common fishes taken
at many of the stations on small upland tributaries. In downstream
collections its relative abundance was less, although it was often
concentrated on riffles.

In the Big Caney system as a whole Notropis umbratilis
was the most abundant species. Several species were present
throughout the system in proportions varying, sometimes greatly,
from station to station. Lepomis megalotis and Lepomis humilis
were erratic in occurrence, and the numbers of Notropis camurus
and Ictalurus melas varied without pattern.

Grouse Creek
The fauna of the main stream of Grouse Creek fluctuated more
in number and kinds of fish from station to station than did the
fauna of Big Caney River. Again, the deep-bodied suckers showed
downstream proclivities. In addition, Notropis buchanani, Pime-
phales tenellus, Percina copelandi, Percina phoxocephala, Notropis
percobromus and Pylodictis olivaris were taken only at the lower-
most station (G-1). At stations G-2 and G-3 the creek is sluggish
and often turbid, meandering between high mud banks in a flood
plain. At these stations Fundulus notatus, Gambusia affinis. La-

390 Uniaersity of Kansas Publs., Mus. Nat. Hist.

hidesthes sicculus, Ictahmis melas, and Lepomis humilis were the
most common fishes. Shiners ( Notropis spp. ) and Lepomis mega-
lotis were rarely taken. Hall (1953:36) states that Gamhusia af-
finis, Fundulus notatus, and Labidesthes dcculus are usually as-
sociated with overflow pools, oxbows, and vegetated backwaters.

Those fishes mentioned in the preceding paragraph remained
common in the middle section of the stream. In addition Notropis
lutrensis, Notropis umhratilis, and Lepomis megalotis were im-
portant members of the fauna.

In the uppermost section shiners {Notropis spp.) were common.
In the few upstream stations that were still in good condition with
clear flowing water, the fauna resembled that of the upstream sta-
tions on Big Caney River. Most upstream stations on Grouse
Creek were located on highly intermittent streams that are treated
below.

FAUNAS OF INTERMITTENT STREAMS

Because of severe, protracted drought, most of the streams studied
had ceased to flow by the close of the survey period. However,
the duration of intermittency varied greatly in different streams, as
did its effect in terms of the number and sizes of residual pools,
water temperatures, pollution, and turbidity. Crab Creek, Beaver
Creek, and a small unnamed tributary of Grouse Creek were
severely affected by intermittency. Their faunas are discussed be-
low.

In Crab Creek six collections were made from points near the
mouth to the uppermost pool in which water was found. Pools
near the mouth were as large as thirty feet in width and ninety
feet in length, while those that were uppermost were shallow
puddles averaging ten feet in length and five feet in width. The
uppermost station was situated in bluestem pasture without benefit
of shade from trees.

The species taken and their relative abundances based on five
seine hauls at each station are shown in Table 7. At the uppermost
pool (G-17) only small green sunfish were found. At G-16, next
downstream, this species was joined by large numbers of black
bullheads and a few redfin shiners and red shiners. G-13 was
similar to G-16, but two additional species occurred there. G-12
was a clear, deep pool much larger than any at the stations up-
stream. Here, seven species were added to the fauna, and the per-
centages of Ictalurus melas and Lepomis cyanellus were much less.
At G-10 Fundulus notatus, Labidesthes sicculus, and Minytrema

Fishes of Arkansas River Basin

391

Table 7. — Percentages of Fishes Taken on Crab Creek.

Stations

G-10

G-11

G-12

G-13

G-16

G-17

Minytrerna melanops. .

8.7
20.0
25.7

Labidesthes sicculus . . .

1.0
41.0

3.8
11.8

9.9

Fundulus notatus

Ictalurus nalalis

.43
1.9
8.5
1.9
1.9
3.8
7.0
7.0

34.0

.5

4.7

25.0
1.9

Pomoxis annularis ....

8.8
15.45

Lepomis himilis

Microplerus salmoides
Etheostoma speciabile . .
Percina caprodes

1.0

Moxostoma erythrurum

1.6
5.7

"2;3'"

Lepomis megalotis. . . .
Pimephales notatus . . .

2.0

9.0

29.0

9.0

14.0

34.0

Ictalurus melas

5.3

49.0
1.0
1.0

49.0

Notropis umbratilis . . .

Notropis hitrensis

Lepomis cyanellus ....

20.6
1.0

26.0

ioo'o '

melanops appeared. Nevertheless, fewer species (10) were cap-
tured here than at station G-12 upstream.

A series of collections similar to that on Crab Creek was carried
out along 1/2 miles of Beaver Creek on July 22, 1956. Nine pools
were sampled (Table 8) of which number nine was the uppermost

Table 8. — Fish Taken in Nine Pools on Upper Beaver Creek (Progress-
ing From Downstream to Upstream).

Notropis
umbratilis

Notropis
lutrensis

Lepomis
humilis

Lepomis
cyanellus

Ictalurus
melas

Pools:
1

2

5 adults
2 adults

4 adults
7 young

4 adults
1 adult

adults
abundant

6 adults

7 adults
4 adults

2 adults

young
abundant

young
abundant

3 juveniles

young
abundant

1 juvenile

3

2 juveniles

4

young

5

abundant

6 . . .

28 young

7 . ..

8

1 adult

9

1 adult

392

llNn-ERsiTY OF Kansas Pxibls., Mus. Nat. Hist.

point where water was found (except for farm ponds). Mainly
young of Lcpomis cyanellus and Ictahirus melas were found in
the uppermost stations, as on Crab Creek. Only adults of Notropis
hitrensis and Notropis mnhratilis were taken.

In another small intermittent tributary of Grouse Creek two
collections (G-14 and G-15) were made. One was from several
isolated pools near the source of the creek and tlie other was VA
miles upstream from the mouth. The two stations were approxi-
mately four miles apart. Table 9 indicates approximate percent-
ages of fish taken in five seine hauls at these stations.

Table 9. — Fishes Taken in a Tributary of Grouse Creek.

Species

Upstream
station

Dov\Tistream
station

Ictalurus melas

45%

48%

5%

2%

Liepomis humilis

40%

Notropis lutrensis

Lepomis cyanellus

30%
20%

Fundulus notatus

10%

At two other stations, only Lepomis cyanellus w^as found. One
of these stations consisted of several small spring-fed pools in a
dry arroyo tributary to Little Beaver Creek. Around these small
"oases" rushes and smartweeds grew and blackbirds were nesting
in the rushes. Although green sunfish up to eight inches in length
were common in the shallow pools, no other species was found.
The second station (C-17) on the East Fork Big Caney River is of
special interest. The pool was isolated, had dimensions of about
25 X 25 feet, and had an average depth of 15 inches. The water
was foul; cows had been fed fodder in a sheltered area above the
pool during the preceding winter and the entire bottom was cov-
ered to a depth of 6 inches to 1 foot with a detritus of decomposing
fodder, cattle feces, and leaves. The water became almost inky in
consistency when the bottom was stirred and its odor was offensive.
A thick gray-green bloom lay on the surface. This bloom was full
of bubbles indicating gases rising from the bottom muds. One
hundred fifty-three green sunfish, all less than 5 inches in length,
were taken in one seine-haul at this station.

EAST-WEST DISTRIBUTION

In the Arkansas River system in Kansas there are marked differ-
ences between fish faunas of the western and eastern parts of the

Fishes of Arkansas River Basin

393

state. This can be illustrated by comparison of Spring River in
Cherokee County with the Cimarron River in southwestern Kan-
sas. Single collections from Spring River or its tributaries usually
contain 25 or more species of fish. Collections from the Cimarron
rarely contain more than five or six species. Many of those fishes
found in Spring River are characteristic of an Ozarkian fauna, and
some are endemic to the Ozark uplands. Fish found in the Cimar-
ron or Arkansas in western Kansas are members of a plains fauna
of wide distribution. There is minghng of these two faunal groups
across the state, with the number of Ozarkian species diminishing
westward, and certain plains species diminishing eastward. A
number of species such as Moxostoma duquesnii and Notropis
spilopterus are limited, on the basis of present records, to Spring
River and its tributaries in Kansas. Others have not been taken
west of the Neosho drainage. The Verdigris River provides the
next major avenue of westward dispersal followed by Caney River,
Grouse Creek, and the Walnut River. West of the Walnut River
system Ozarkian species have been almost always absent from col-
lections. The Chikaskia River is somewhat exceptional. Moore
and Buck ( 1953 ) reported from this river several species that seem
more typical of eastern faunal associations. Table 10 indicates the
stream system in which the present westernmost records are located
for a number of fishes found in the Arkansas River system in
Kansas.

Table 10. — Present Westernmost Records of Some Fishes in the
Arkansas Rtver Basin in ICansas.

Spring River
Cottus carolinae
Dionda nubila
Etheostoma blenniaides
Etheostoma gracile
Etheostoma nigrum nigrum
Etheostoma punctulatum
Etheostoma saxatile
Hypentelium nigricans
Moxostoma duquesnii
Notropis spilopterus
Noturus exilis

Neosho River

Cycleptus elongatus
Etheostoma chlorosomum
Etheostoma flabellare lineolatum
Hybopsis amblops
Hybopsis biguttata
Hybopsis x-punctata
Notropis zonatus pilsbryi

Verdigris River
Etheostoma whipplii
Etheostoma zonale arcansanum
Percina copelandi
Moxostoma carinatum
Notropis boops
Notropis volucellus
Noturus miurus

Chikaskia River

Ictalurus natalis
Percina phoxocephala
Labidesthes sicculus
Lepomis megalotis breviceps
Micropterus punctulatus
Moxostoma aureolum pisolabrum
Moxostoma erythrurum
Notropis camurus
Pimephales notatus
Pimephales tenellus
Noturus noctumus

4—7079

394 University of Kansas Publs., Mus. Nat. Hist.

The westernmost records for seven species are in the area studied.

1. Lepisosteus platostomus.

2. Carpiodes velifer.

3. Moxostoma carinattim.

4. Minytrema meJanops. One specimen taken at station G-10
near the mouth of Crab Creek constitutes the present westernmost
record. A specimen has been taken by Cross (C-24-51) in the
headwaters of the Walnut River.

5. Notropis hoops. The westernmost record is station G-5 on
Grouse Creek. This fish has been reported sHghtly west of this in
Oklahoma on Big Beaver Creek in Kay County (number 4776,
Oklahoma A & M College Museum of Zoology).

6. Notropis volucellus. Two specimens were taken at station
G-8 on Silver Creek.

7. Percina copelandi. The westernmost record is from station
G-1, two miles above the mouth of Grouse Creek.

The easternmost occurrences of four species are in the area
studied. These species are Hybopsis aestivalis tetranemus ( Station
A-2), Notropis blennius (Station A-1), Notropis girardi (Station
A-2), and Fundtdiis kansae (Station A-2 and Walnut River).
These fish are associated with the Arkansas River proper and its
sandy western tributaries. In Oklahoma, these fish are found in
the Arkansas River as it proceeds eastward and in the downstream
portions of some of its tributaries. These fish show little tendency
to ascend the streams of the Flint Hills.

SUMMARY

The fish fauna of the area studied is transitional between the
Ozarkian and Great Plains faunas.

Fluctuation in water level seemed especially important in de-
termining distribution of fishes in the area studied. Variable climate
characteristic of the region studied causes recurrent floods and in-
termittency in streams. Both of these conditions have probably
been accentuated by man's modifications of tlie habitat. The effects
of intermittency were most strikingly demonstrated in small creeks
of the uplands. The number of species of fish in the highly inter-
mittent streams was small — especially in the uppermost pools
sampled — but the actual number of fish was often high even though
the number of species was low. In several instances the only
fishes found in these isolated pools were Lepoinis cyanellus and
Ictalurus melas. Tliis phenomenon of concentrated numbers of
individuals of a few species would indicate the presence of limiting

Fishes of Arkansas River Basin 395

factors that allow only those species most tolerant of the particular
factor to flourish.

Soon after rains restored flow in these intermittent creeks
L. cyanellus and I. melas appeared in parts of the channels that
had previously been several miles from the nearest water. Rapid
upstream movements of other species after rains was also noted.

It was impossible to ascertain the precise effects of gradient and
bottom-type on distribution, but certain species such as Notropis
blennius, Notropis girardi, and Ftindulus kansae were taken only
in streams with sandy bottoms. Notropis deliciosus and Hybog-
nathus placita were most abundant over sandy bottoms.

The high gradient of upland tributaries in the Flint Hills area
produced turbulence and bottoms predominantly of rubble. A
fauna of which Etheostoma spectahile and Campostoma anomalum
were characteristic existed in these waters while they were flowing.
As flow decreased and intermittency commenced, qualitative and
quantitative changes in the fish faunas were observed. Gradient did
not change during drought, but turbulence did. Because turbu-
lence varies with water level as well as gradient, the effect of
gradient on fish distribution ultimately is linked to climate.

Probably the small number of fish taken on the Walnut River
in comparison with other eastern Kansas rivers (Verdigris, Neosho)
results, in part, from the long-term pollution of the stream noted by
Clapp (1920:33) and Doze (1924). No percid fishes, black bass,
or madtom catfish were taken on the Walnut in Cowley County
and the species of Notropis numbered only three.

Four faunal associations seem to be recognizable in the area.

Arkansas River Fauna

This fauna contained Notropis girardi, Notropis blennius, Hybop-
sis aestivalis tetranemus, and Fundulus kansae which, in this area,
did not seem to wander far from the sandy main stream of the
Arkansas. Minnows abounded; Notropis lutrensis and N. deliciosus
missuriensis predominated; and Notropis girardi, N. percobromus,
and Hybognathus placita were common. In quiet backwaters,
coves, and shallow pools Gambusia affinis occurred in great num-
bers. Lepisosteus osseus seemed to be the most important
predator.

Lower Walnut River Fauna

The Walnut River in Cowley County supported large popula-
tions of deep-bodied suckers, carp, and gar. Notropis lutrensis
and N. percobromus were characteristic minnows. Lepomis

396 University of Kansas Publs., Mus. Nat. Hist.

himiilis abounded at some stations. The fauna of the main stream
of the Walnut River was somewhat intermediate between that of
the Arkansas River and that of the three streams considered below.
Fifteen of the species common to the Big Caney, Elk, and Grouse
systems were also taken in the Walnut River main stream. Thirteen
species were common to the Walnut and Arkansas rivers. Seven
species were common to all these streams.

Caney-Elk-Grouse Main Stream Fauna
This fauna includes fishes living not only in the main streams but
also in the lower parts of the larger tributaries of these streams.
The fauna was comparatively rich: in the main stream of Big Caney
River 39 species were taken, in Grouse Creek 35 species, in the
Walnut River main stream 21 species, and in the Arkansas River
19 species. It has been pointed out that large rivers such as the
Walnut and Arkansas have been subjected to greater direct and
indirect modification by man, possibly resulting in a less diverse
fauna than would otherwise occur in these streams. At present,
there is a paucity of ecological niches in the upland tributaries and
large rivers, as compared with streams of intermediate size. Fishes
typical of the Caney-EUc-Grouse association were Notropis um-
hratilis, Lepomis megalotis, Lepomis htimilis, Labidesthes sicculus,
Ftindulus notatus, and the two species of Micropterus ( Micropterus
punctulatiis was not taken in Grouse Creek).

Upland Tributary Fauna

Tributary faunas were divisible into two categories: (1) Those
nf the Walnut River and Grouse Creek ( intermittency was severe,
species were few, with Ictalurus melas and Lepomis cyanelltts
predominating); (2) those of Big Caney River (stream-flow was
more stable, and eastern fishes, some of which have Ozarkian
aflBnities, occurred in greater abundance than in any other part of
the area surveyed). In the latter streams Campostoma anomaJum
and Etheostoma sjyectabile usually were dominant. Pimephales
notatus, Notropis vohicellus, N. camurus, N. boops, and N. rubellus
characteristically occurred. Notropis lutrensis was sparsely repre-
sented in flowing tributaries. Notropis umhratilis, which seems to
prefer habitats intermediate between those of Notropis lutrensis
and Ozarkian shiners, was usually represented. Deep-bodied suck-
ers and carp were not taken in upland tributaries but Moxostoma
erythrurum was common and Minytrema melanops was taken.

The kinds and numbers of shiners (Notropis) taken at different

Fishes of Arkansas River Basin 397

points along Grouse Creek seem significant. N. liitrensis and N.
timhratilis occurred throughout the stream but were rare in slug-
gish areas where populations of Gamhusia affinis, Fundulus notatus,
and Labidesthes sicculus flourished. At the lowermost station
Notropis percobromus and IV. buchanani were taken; these were
not present in other collections. In the uppermost stations where
water remained plentiful, N. boops and N. volucellus were taken,
and N. rubellus has been recorded.

In the broader distributional sense those fishes that seemed most
tolerant of intermittency (Lepomis cyanellus, Lepomis humilis,
Ictalurus melas, Notropis lutrensis) are widely distributed in the
Arkansas River Basin, and are common in the western part of the
Arkansas River Basin. Species less tolerant of intermittency are
Notropis boops, Notropis camurus, Notropis rubellus, Notropis
volucellus, and Pimephales tenellus; they have not been taken far
west of the area studied, and become more common east of it.

LITERATURE CITED

Bass, N. W.

1929. The geology of Cowley County, Kansas. Kansas Geol. Survey Bull.,
12:1-203, 23 figs., 12 pis.

BlEBEB, R. P.

1932. Frontier life in the army, 1854-1861. Southwest Historical Series,
2:1-330.

Bbeitkelman, J.

1940. A collection of fishes in the State University Museum. Trans.
Kansas Acad. Sci., 43:377-384.

Buck, H., and Cross, F. B.

1951. Early limnological and fish population conditions of Canton Reser-
voir, Oklahoma, and fishery management recommendations. A
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Research Foundation, Oklahoma A&M College. 110 pp., 17 figs.

Caldwell, M. B.

1937. The soutliern Kansas boundary survey. Kansas Hist. Quart., 6:339-
377.

Clapp, a.

1920. Stream pollution. Kansas Fish and Game Department Bull., 6:33.

Cross, F. B.

1950. Effects of sewage and of a headwaters impoundment on the fishes
of Stillwater Creek in Payne County, Oklahoma. Amer. Midi. Nat.,
43 (1):128-145, Ifig.

1954a. Fishes of Cedar Creek and the south fork of the Cottonwood
River, Chase County, Kansas. Trans. Kansas Acad. Sci. 57:303-
314.

1954b. Records of fishes little-known from Kansas. Trans. Kansas Acad.
Sci. 57:473-479.

398 University of Kansas Publs., Mus. Nat. Hist.

Choss, F. B., and Moore, G. A.

1952. The fishes of the Poteau River, Oklahoma and Arkansas. American
Midi. Nat, 47 (2):396-412.

Doze, J. B.

1924. Stream pollution. Bien. Report. Kansas Fish and Game Dept.
5:1-42.

Elkin, R. E.

1954. The fish population of two cut-off pools in Salt Creek. Osage
County, Oklahoma. Proc. Oklahoma Acad. Sci., 35:2.5-29.

Elliott, A.

1947. A preliminary survey and ecological study of the fishes of the South
Ninnescah and Spring Creek. Unpublished thesis, Kansas State
College.

Evermann, B. W., and Fordice, M. W.

1886. List of fishes collected in Harvey and Cowley counties, Kansas.
Bull. Washburn Lab. Nat. Hist., 1:184-186.

Flora, S. D.

1948. Climate of Kansas. Rept. Kansas State Board Agric. 67:xii-320.
Illus.

Foley, F. C, Smrha, R. V., and Metzler, D. F.

1955. Water in Kansas. A report to the Kansas State Legislature as
directed by the Kansas State Finance Council. University of Kan-
sas, pp. 1-216— A1-J6.

Frye, J. C, and Leonard, A. B.

1952. Pleistocene geology of Kansas. Bull. Kansas Geol. Surv., 99:1-230.
17 figs., 19 pis.

FxJNK, J. L., and Campbell, R. S.

1953. The population of larger fishes in Black River, Missouri. Univ.
Missouri Studies, 26:69-82.

Gates, F. C.

1936. Grasses in Kansas. Rept. Kansas State Board Agric, 55 (220-A):
1-349, frontispiece, 270 figs., 224 maps.

Graham, I. D.

1885. Preliminary list of Kansas fishes. Trans. Kansas Acad. Sci., 9:69-

78.

Hale, M. E., Jr.

1955. A survey of upland forests in the Chautauqua Hills, Kansas. Trans.
Kansas Acad. Sci., 58:165-168.

Hall, G. E.

1952. Observations on the fishes of the Fort Gibson and Tenkiller reser-
voir areas, 1952. Proc. Oklahoma Acad. Sci., 33:.55-63.

1953. Preliminary observations on the presence of stream-inhabiting
fishes in Tenkiller Reservoir, a new Oklahoma impoundment.
Proc. Oklahoma Acad. Sci., 34:34-40.

Hoyle, W. L.

1936. Notes on faunal collecting in Kansas. Trans. Kansas Acad. Sci.,
39:283-293.

Fishes of Arkansas River Basin 399

HuBBS, C. L., and Ortenburger, A. I.

1929a. Further notes on the fishes of Oklahoma with descriptions of new
species of cyprinidae. Publ. Univ. Oklahoma Biol. Surv., 1
(2): 17-43.

1929b. Fishes collected in Oklahoma and Arkansas in 1927. Publ. Univ.
Oklahoma Biol. Surv., 1 (3):47-112, 13 pis.

HuBBS, C. L., and Lagler, K. F.

1947. Fishes of the Great Lakes Region. Cranbrook Inst. Sci. Bull.,
26 (Revised Edition) :i-xi-l-186, illus.

Jewett, J. M., and Abernathy, G. E.

1945. Oil and gas in eastern Kansas. Bull. Kansas Geol. Survey, 57:1-
244, 21 figs., 4 pis.

Metzler, D. F.

1952. Water Pollution Report, Walnut River Basin. Department of Sani-
tation, Kansas State Board of Health (Unpublished), 64 pp.

Miller, N. H.

1932. Surveying the southern boundary fine of Kansas. Kansas Hist.
Quarterly, 1:104-139.

Moore, G. A.

1944. Notes on the early hfe history of Notropis girardi. Copeia, 1944
(4):209-214, 4Figs.

Moore, G. A., and Cross, F. B.

1950. Additional Oklahoma fishes with vaHdation of Poecilichthys
parvipinnis (Gilbert and Swain). Copeia, 1950 (2): 139-148.

Moore, G. A., and Paden, J. M.

1950. The fishes of the Illinois River in Oklahoma and Arkansas. Amer.
Midi. Nat., 44:76-95, 1 Fig.

Moore, G. A., and Buck, D. H.

1953. The fishes of the Chikaskia River in Oklahoma and Kansas. Proc.
Oklahoma Acad. Sci., 34:19-27.

Moore, R. C.

1949. Divisions of the Peimsylvanian system in Kansas. Bull. Kansas
Geol. Survey, 83: 1-203, 37 Figs.

Moore, R. C, Frye, J. C, Jewett, J. M., Lee, W., and O'Conner, H. G.

1951. The Kansas rock column. Bull. Kansas Geol. Survey, 89:1-132,
52 Figs.

Mooso, J.

1888. The life and travels of Josiah Mooso. Telegram Post, Winfield,
Kansas, pp. 1-400.

Ortenbxjrger, a. I., and Hubbs, C. L.

1926. A report on the fishes of Oklahoma, with descriptions of new
genera and species. Proc. Oklahoma Acad. Sci., 6:132-141.

SCHELSKE, C. L.

1957. An ecological study of the fishes of the Fall and Verdigris rivers
in Wilson and Montgomery counties, Kansas, March 1954, to
February 1955. Emporia State Research Studies, 5 (3):31-56.

400 Uni\trsity of Kansas Publs., Mus. Nat. Hist.

ScHOONOvER, R., and Thompson, W. H.

1954. A post-impoundment study of the fisheries resources of Fall River
Reservoir, Kansas. Trans. Kansas Acad. Sci., 57:172-179.

Tr.\utman, M. B.

1951. Moxostoma aureolum pisolabrum, a new subspecies of sucker from
the ozarkian streams of the Mississippi River System. Occ. Papers
Mus. Zool. Univ. Michigan, 534:1-10, 1 pi.

Transmitted December 19, 1958.

D

27 7079

(Continued from inside of front cover)

Vol 8 1. Life history and ecology of the five-lined skink, Evimeces fasciatus. By Henry
S. Fitch. Pp. 1-156, 26 figs, in text. September 1, 1954.

2. Myology and serology of the Avian Family Fringillidae, a taxonomic study.
By William B. Stallcup. Pp. 157-211, 23 figures in text, 4 tables. November
15, 1954.

3. An ecological study of the collared lizard (Crotaphytus collaris). By Henry
S. Fitch. Pp. 213-274, 10 figures in text. February 10, 1956.

4. A field study of the Kansas ant-eating frog, Gastrophryne ohvacea. By Henry
S. Fitch. Pp. 275-306, 9 figures in text. February 10, 1956.

5. Check-list of the birds of Kansas. By Harrison B. Tordoff. Pp. 307-359, 1
figure in text. March 10, 1956.

6. A population study of the prairie vole (Microtus ochrogaster) in northeastern
Kansas. By Edwin P. Martin. Pp. 361-416, 19 figures in text. April 2,
1956.

7. Temperature responses in free-living amphibians and reptiles of northeastern
Kansas. By Henry S. Fitch. Pp. 417-476, 10 figures in text, 6 tables. June
1, 1956.

8. Food of the crow, Corvus brachyrhynchos Brehm, in south-central Kansas. By
Dwight Piatt. Pp. 477-498, 4 tables. June 8, 1956.

9. Ecological observations on the woodrat, Neotoma floridana. By Henry S.
Fitch and Dennis G. Rainey. Pp. 499-533, 3 figures in text. June 12, 1956.

10. Eastern woodrat, Neotoma floridana: Life history and ecology. By Dennis
G. Rainey. Pp. 535-646, 12 plates, 13 figures in text. August 15, 1956.'
Index. Pp. 647-675.
Vol. 9. 1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 fig-
ures in text. December 10, 1955.

2. Additional records and extensions of ranges of mammals from Utah. By
Stephen D. Diurant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80.
December 10, 1955.

3. A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rol-
lin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.

4. Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming.
By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956.

5. The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6
figures in text. May 19, 1956.

6. Additional remains of the multituberculate genus Eucosmodon. By Robert
W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.

7. Mammals of Coahuila, Mexico. By RoUin H. Baker. Pp. 125-335, 75 figures
in text. June 15, 1956.

8. Comments on the taxonomic status of Apodemus peninsulae, with description
of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 337-346,
1 figure in text, 1 table. August 15, 1956.

9. Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp.
347-351. August 15, 1956.

10. A new bat (Genus Leptonycteris) from Coahuila. By Howard J. Stains.
Pp. 353-356. January 21, 1957.

11. A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexica
By Robert J. Russell. Pp. 357-361. January 21, 1957.

12. Geographic variation in the pocket gopher, Thomomys bottae, in Colorado.
By Phillip M. Youngman. Pp. 363-387, 7 figmes in text. February 21,
1958.

13. New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones,
Jr. Pp. 385-388. May 12, 1958.

14. Pleistocene bats from San Josecito Cave, Nuevo Leon, Mexico. By J. Knox
Jones, Jr. Pp. 389-396. December 19, 1958.

15. New Subspecies of the rodent Baiomys from Central America. By Robert L.
[Packard. Pp. 397-404. December 19, 1958.

More numbers will appear in volume 9.
VoL 10. 1. Studies of birds killed in noctvumal migration. By Harrison B. TordofiF and
Robert M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 1956.

2. Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.

3. The forest habitat of the University of Kansas Natural History Reservation.
By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures
in text, 4 tables. December 31, 1956.

(Continued on outside of back cover)

(Continued from inside of back cover)

4. Aspects of reproduction and development in the prairie vole (Microtus ochro-
gaster). By Henry S. Fitch. Pp. 129-161, 8 figures in text, 4 tables. Decem-
ber 19, 1957.

5. Birds found on the Arctic slope of northern Alaska. By James W. Bee.
Pp. 163-211, pis. 9-10, 1 figure in te.\t, March 12, 1958.

6. The wood rats of Colorado: distribution and ecology. By Robert B. Finley,
Jr. Pp. 213-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958.
More number will appear in volume 10.

Vol. 11. 1. The systematic status of the colubrid snake, Leptodeira discolor Giinther.
By William E. Duellman. Pp. 1-9, 4 figs. July 14, 1958.

2. Natural history of the six-lined raceninner, Cnemidophorus sexUneatus. By
Henry S. Fitch. Pp. 11-62, 9 figs., 9 tables. September 19, 1958.

3. Home ranges, territories, and seasonal movements of vertebrates of the
Natural History Reservation. By Henry S. Fitch. Pp. 63-326, 6 plates, 24
figures in text, 3 tables. December 12, 1958.

4. A new snake of the genus Geophis from Chihuahvia, Mexico. By John M.
Legler. Pp. 327-334, 2 figures in text. January 28, 1959.

5. A new tortoise, genus Gopherus, from north-central Mexico. By John M.
Legler. Pp. 335-343, 2 plates. April 24, 1959.

6. Fishes of Chautauqua, Cowley and Elk counties, Kansas. By Artie L.
Metcalf. Pp. 345-400, 2 plates, 2 figures in text, 10 tables. May 6, 1959.

More numbers will appear in Volume 11.

jv n L-

m.mi zcoi

JUN181959

University of Kansas PuBLicAxiAai*
Museum of Natural History

wmm

Volume 11, No. 7, pp. 401-442, 2 plates, 4 figs, in text, 5 tables
May 8, 1959

Fishes of the Big Blue River Basin,

Kansas

BY

W. L. MINCKLEY

University of Kansas
Lavv^rence

■^^959

UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY

Institutional libraries interested in publications exchange may obtain this
series by addressing the Exchange Librarian, University of Kansas Library,
Lawrence, Kansas. Copies for individuals, persons working in a particular
field of study, may be obtained by addressing instead the Museum of Natural
History, University of Kansas, Lawrence, Kansas. There is no provision for
sale of this series by the University Library which meets institutional requests,
or by the Museum of Natural History which meets the requests of individuals.
However, when individuals request copies from the Museum, 25 cents should
be included, for each separate number that is 100 pages or more in length, for
the piupose of defra>'ing the costs of wTapping and mailing.

* An asterisk designates those numbers of which the Museum's supply (not the Library's
supply) is exhausted. Numbers published to date, in this series, are as follows:
Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950.
*Vol, 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140
figures in text. April 9, 1948.
Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, and distribution. By Rol-
lin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951.
*2, A quantitative study of the nocturnal migration of birds. By George H.
Lowery, Jr. Pp. 361-472, 47 figm-es in text. June 29, 1951.

3. Phylogenv of the waxwings and allied birds. By M. Dale Arvey. Pp. 473-
530, 49 figures in text, 13 tables. October 10, 1951.

4. Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and
Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10,
1951.

Index. Pp. 651-681.
*Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31

figures in text. December 27, 1951.
Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953.

*Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D.
Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952.
VoL 7. *1. Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, 73 figures in text,
57 tables. August 25, 1952.

2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry
S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 figures in text. Augusf
24, 1953.

3. The silky pocket mice (Perognathus flavus) of Mexico. By RoUin H. Baker.
Pp. 339-347, 1 figme in text. February 15, 1954.

4. North American jumping mice (Genus Zapus). By Philip H. Krutzsch. Pp.
349-472, 47 figures in text, 4 tables. April 21, 1954.

5. Mammals from Southeastern Alaska. By Rollin H. Baker and James S.
Findley. Pp. 473-477. April 21, 1954.

6. Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479-
487. AprU 21, 1954.

7. Subspeciation in the montane meadow mouse. Microtus montanus, in Wyo-
ming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text.
July 23, 1954.

8. A new subspecies of bat (Myotis velifer) from southeastern CaUfomia and
Arizona, By Terry A. Vaughan. Pp. 507-512. July 23, 1954.

9. Mammals of the San Gabriel mountains of California. By Terry A. Vaughan.
Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954.

10. A new bat (Genus Pipistrellus ) from northeastern Mexico. By Rollin H.
B.iker. Pp. 583-586. November 15, 1954.

11. A new subspecies of pocket mouse from Kansas. By E. Raymond Hall. Pp.
587-590. November 15, 1954.

12. Geographic variation in the pocket gopher, Cratogeomys castanops, in Coa-
huila, Mexico. By Robert J. Russell and Rollin H. Baker. Pp. 591-608.
March 15, 1955.

13. A new cottontail (Sylvilagus floridanus) from northeastern Mexico, By RoUiil
H. Baker. Pp. 609-612. April 8, 1955.

14. Taxonomy and distribution of some American shrews. By James S. Findley,
Pp. 613-618. June 10, 1955.

15. The pigmy woodrat, Neotoma goldmani, its distribution and systematic posi-
tion. By Dennis G. Rainey and Rollin H. Baker. Pp. 619-624. 2 figures in
text. June 10, 1955.

Index. Pp. 625-651.

(Continued on inside of back cover)

University of Kansas Publications
Museum of Natural History

Volume 11, No. 7, pp. 401-442, 2 plates, 4 figs, in text, 5 tables
May 8, 1959

Fishes of the Big Blue River Basin,

Kansas

BY
W. L. MINCKLEY

University of Kansas

Lawrence

1959

Untveksity of Kansas Publications, Musexjm of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 11, No. 7, pp. 401-442, 2 plates, 4 figs, in text, 5 tables
Published May 8, 1959

MUS. COM?. ZOOl

JUN181959

UNIVERSilY

University of Kansas
Lawrence, Kansas

A Contribution From
The State Biological Survey of Kansas

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA. KANSAS

1969

27-7080

Fishes of the Big Blue River Basin,
Kansas

BY
W, L. MINCKLEY

CONTENTS

PAGE

Introduction 403

Acknowledgments 404

Tuttle Creek Dam and Reservoir 404

Big Blue River Basin 404

Geology of the basin 405

Climate, population, and land-use 406

Physical features of streams 407

Previous records of fishes 410

Methods and materials 410

Collecting stations 412

Annotated list of species 414

Hybrid combinations 431

Relative abundance and discussion of species 431

Creel census 435

Recommendations 437

Summary 438

Literature cited 438

INTRODUCTION

The Big Blue River in northeastern Kansas will soon be im-
pounded by the Tuttle Creek Dam, located about five miles north
of Manhattan, Kansas. Since the inception of this project by the
U. S, Army Corps of Engineers much argument has arisen as to
the values of the dam and reservoir as opposed to the values of
farmland and cultural establishments to be inundated (Schoewe,
1953; Monfort, 1956; and Van Orman, 1956). Also, there has
been some concern about the possible eflFects of impoundment on
the fish-resources of the area, which supports "a catfish fishery
that is notable throughout most of the State of Kansas and in some
neighboring states (U. S. Fish and Wildlife Service, 1953:9)."
The objectives of my study, conducted from March 30, 1957, to

(403)

404 University of Kansas Publs., Mus. Nat. Hist.

August 9, 1958, were to record the species of fish present and their
relative abundance in the stream system, and to obtain a measure
of angler success prior to closure of the dam. These data may
be used as a basis for future studies on the fish and fishing in the
Big Blue River Basin, Kansas.

ACKNOWLEDGMENTS

I thank Messrs. J. E. Deacon, D. A. Distler, Wallace Ferrel, D. L. Hoyt,
F. E. Maendcle, C. O. Minckley, B. C. Nelson, and J. C. Tash for assistance
in the field and for valuable suggestions. Dr. J. B. Elder, Kansas State
College, arranged for loan of specimens, and Mr. B. C. Nelson supplied data
on Notropis deliciosus (Girard) in Kansas, and on specimens in the University
of Michigan Museum of Zoology.

I thank the many landowners who allowed me access to streams in the
Big Blue River Basin. The U. S. Army Corps of Engineers, Kansas City
District, also allowed access in the reservoir area, and furnished information
and some photographs. Mr. J. C. Tash did chemical determinations on my
water samples.

Dr. Frank B. Cross guided me in this study and in preparation of this re-
port. Drs. E. Raymond Hall and K. B. Armitage oflFered valuable suggestions
on the manuscript. Equipment and funds for my study were furnished by
the State Biological Survey of Kansas, and the Kansas Forestry, Fish and
Game Commission granted necessary permits.

TUTTLE CREEK DAM AND RESERVOIR

The data on Tuttle Creek Dam and Reservoir that follow were
furnished by Mr. Donald D. Poole, U. S. Army Corps of Engineers,
Kansas City District. The dam, an earth-fill structure, will be
7,500 feet in length, with a maximum height of 157 feet
above the valley floor. Release of water will be from beneath the
west end of the dam, through two tunnels 20 feet in diameter that
have a capacity of 45,000 cubic feet per second; however, releases
exceeding 25,000 c. f. s. are not planned. The gated spillway is
located at the east end of the dam. Freeboard will be 23 feet
at the top of flood-control pool.

The reservoir will have a maximum pool of 2,280,000 acre-feet ca-
pacity, a 53,500-acre surface area, and 368 miles of shoreline.
The present operational plan provides for a conservation pool
having a surface area of 15,700 acres, a shoreline of 112 miles,
and a length of 20 miles.

BIG BLUE RIVER BASIN

Big Blue River and its tributaries, a sub-basin of the Kansas
River System, drain approximately 9,600 square miles, of which
2,484 miles are in Kansas (Colby, et al, 1956:44). The head-

Fishes of Big Blue Basin

405

waters of the Big Blue River are in central Hamilton County,
Nebraska, near the Platte River (Fig. 1). The stream flows gen-
erally south and east for 283 miles to its confluence with the Kansas
River near Manhattan, Kansas. Little Blue River, the largest
tributary to the Big Blue, rises in eastern Kearney and western
Adams counties, Nebraska, and flows southeast for 208 miles to
join the Big Blue near Blue Rapids, Kansas (Nebraska State Plan-
ning Board, 1936:628). The Big Blue River Basin varies in width
from 129 miles in the northwest, to approximately ten miles near
the mouth (Colby, et al, 1956:44).

GEOLOGY OF THE BASIN

In Kansas, outcrops of Pennsylvanian and Cretaceous age occur
along the extreme eastern and western sides of the Big Blue River
Basin, respectively, whereas Permian beds (overlain by Pleistocene
deposits) occur throughout most of the remainder of the water-
shed (see Moore and Landes, 1937), The Big Blue and Little
Blue rivers and their tributaries have deeply incised the Permian
beds of the Flint Hills in Kansas, exposing limestones and shales

NEBRASKA

MISSOURI
RIVER

— 40

KANSAS

Fig. 1. Big Blue River Basin, Kansas and Nebraska.

—h

406 University of Kansas Publs., Mus. Nat. Hist.

of the Admire, Council Grove, Chase, and Sumner groups (Wolf-
campian and Leonardian series) (Walters, 1954:41-44). Pleisto-
cene deposits in the Big Blue Basin in Kansas consist of alluvium,
glacial till, and glacial outwash from the Kansan glacial stage,
overlain by loess deposits of Wisconsin and Recent stages (Frye
and Leonard, 1952: pi. 1).

The Big Blue River was formed "in part on the till plain surface
and in part by integration of spillway channels," in the latter portion
of the Kansan glaciation (Frye and Leonard, 1952:192). This
stream, and the Repubhcan River to the west, carried waters from
the areas that are now the Platte, Niobrara, and upper Missouri
River basins (Lugn, 1935:153). Drainage was southward, through
Oklahoma, until establishment of the east-flowing Kansas River
(Frye and Leonard, 1952:189-190). As Kansan ice receded the
Blue and Republican rivers retained what is now the Platte River
Basin. The lower Platte River developed and the surface drainage
became distinct in the lowan ( Tazwellian ) portion of the Wisconsin
glacial stage (Lugn, 1935:152-153). However, according to Lugn
(1935:203) the Platte River Basin contributes about 300,000 acre-
feet of water per year to the Big Blue and Republican rivers by
percolation through sands and gravels underlying the uplands that
now separate the basins.

CLIMATE, POPULATION, AND LAND-USE

Climate of the Big Blue River Basin is of the subhumid con-
tinental type, with an average annual precipitation of 22 inches
in the northwest and 30 inches in the southeast. The mean annual
evaporation from water surfaces exceeds annual precipitation by
approximately 30 inches (Colby, et al., 1956:32-33).

The average annual temperature for the basin is 53° F. (Flora,
1948:148). According to Kincer (1941:704-705) the average
temperature in July, the warmest month, is 78° F., and the
coolest month, January, averages 28° F. Periods of extreme cold
and heat are sometimes of long duration. Length of the growing
season varies from less than 160 days in the northwest to 180 days
in the southeast (Kincer, loc. cit.).

The human population of the Big Blue Basin varies from about
90 persons per square mile in one Nebraska county in the north-
west and one Kansas county in the southeast, to as few as six
persons per square mile in some northeastern counties. The
population is most dense along the eastern border of the basin,
decreasing toward the west. This decrease in population is corre-

Fishes of Big Blue Basin 407

lated with the decrease in average annual precipitation from east
to west (Colby, et al, 1956:80).

The principal land-use in the Big Blue Watershed is tilled
crops, with wheat, sorghums, and corn being most important.
Beef cattle are important in some portions of the basin. Colby,
et al. (1956:24) reported that in 1954 as much as 55 per cent of
the land in some counties near the mouth of the Big Blue River
was in pasture. Only one Nebraska county had less than 15 per
cent in pastureland.

PHYSICAL FEATURES OF STREAMS

Streams of the Big Blue River Basin are of three kinds: turbid,
sandy-bottomed streams, usually 150 to 300 feet in width; relatively
clear, mud-bottomed streams, ten to 60 feet in width; and clear,
deeply incised, gravel-bottomed streams, usually five to 30 feet
in width.

Sand-bottomed Streams. — The Big Blue and Little Blue rivers
represent this kind of stream. The bottoms of these rivers consist
almost entirely of fine sand; nevertheless, their channels are pri-
marily deep and fairly uniform in width, rather than broad, shal-
low, and braided as in the larger Kansas and Arkansas rivers in
Kansas (Plate 11, Fig. 1). In the Big Blue River, gravel occurs
rarely on riffles, and gravel-rubble bottoms are found below dams
(Plate 11, Fig. 2). The Big Blue flows over a larger proportion
of gravelly bottom than does the Little Blue.

Big Blue River rises at about 1,800 feet above mean sea level
and joins the Kansas River at an elevation of 1,000 feet above
m. s. 1. The average gradient is 2.8 feet per mile. Little Blue River,
originating at 2,200 feet, has an average gradient of 5.3 feet per
mile, entering the Big Blue at 1,100 feet above mean sea level
(Nebraska State Planning Board, 1936:628, 637). The Little Blue
is the shallower stream, possibly because of the greater amount
of sandy glacial deposits in its watershed and the swift flow that
may cause lateral cutting, increased movement, and "drifting"
of the sandy bottom.

For approximately a 50-year period, stream-flow in the Big Blue
River at its point of entry into Kansas (Barnston, Nebraska)
averaged 603 cubic feet per second, with maximum and minimum
instantaneous flows of 57,700 c. f. s. and one c. f. s. The Little Blue
River at Waterville, Kansas, averaged a daily discharge of 601
c. f. s. (maximum 50,400, minimum 28). Below the confluence
of the Big Blue and Little Blue rivers, at Randolph, Kansas, the

408

University of Kansas Publs., Mus. Nat. Hist.

average daily discharge was 1,690 c.f.s. (maximum 98,000, mini-
mum 31) (Kansas Water Resources Fact-finding and Research
Committee, 1955:27).

The turbidity of the Big Blue River, as determined by use of
a Jackson turbidimeter, varied from 27 parts per million in winter
(January 10, 1958) to as high as 14,000 p. p. m. (July 12, 1958).
The Little Blue River has similar turbidities, with high readings
being frequent. In the summer of 1957, pH ranged from 7.2 to
8.4 in the Big Blue River Basin — values that correspond closely
with those of Canfield and Wiebe (1931:3) who made 25 deter-
minations ranging from 7.3 to 8.3 in the streams of the Nebraskan
portion of this basin in July, 1930. Surface temperatures at var-
ious stations varied from 38' F. on January 10, 1958, to 90° F.
in backwater-areas on July 19, 1957. The average surface tem-
perature at mid-day in July and August, 1957, was approximately
86.5° F.

Chemical determinations were made on water-samples from
my Station 4-S on the Big Blue River, and Station 50-S on the
Little Blue ( Table 1 ) . These samples were taken from the surface
in strong current. Determinations were made by methods described
in Standard Methods for the Examination of Water and Sewage,
10th edition, 1955.

Table 1. — Chemical Determinations in Milligrams Per Liter at Five
Stations in the Big Buje River Basin, Kansas, 1958.

Station

AND

Date

o 5

° 2

(0

CO

-a

ca

u.

-C

^

2

a.

3

u

w

^

en

0)

2

"5
o

a

00

«

-a
a

tn
O
J3
Ph

4-S
August 9. . . .

50-S
Augu.gt 9. . . .

35-M
August 9. . . .

11-G

Julys

18-G
July 22

0.0

154

16

28

3.5

.083

.250

0.0

125

24

20

2.5

.669

.427

0.0

3G6

15

108

9.4

.220

.750

0.0

272

15

CO

4.5

.060

.625

0.0

183

10

60

1.6

.938

.293

.225

.240

.080

140

.240

Fishes of Big Blue Basin 409

The banks of both the Big Blue and Little Blue rivers support
narrow riparian forests comprised primarily of elm, Ulmus ameri-
canus, Cottonwood, Populus deltoides, sycamore, Platanus occiden-
talis, and willow, Salix spp. Maple, Acer sp., oak, Querctis spp.,
and ash, Fraxinus sp. occur where the rivers flow near steep, rocky
hillsides. Many of the hills are virgin bluestem prairies (Andropo-
gon spp. ) , but the floodplains are heavily cultivated.

Mud-bottomed Streams. — Streams of this kind are present in
the watershed of the Black Vermillion River that enters Big Blue
River from the east. The area east of the Big Blue River and
north of the Black Vermillion River is till plains, where rehef sel-
dom exceeds 100 feet (Walters, 1954:12). Streams in this portion
of the basin, and streams entering the Little Blue River from the
west (Mill Creek and Horseshoe Creek systems), tend to have
V-shaped channels, fewer riffles than the Little Blue and Big Blue
rivers and in the gravelly streams (to be described later), and
have bottoms of mud or clay, with few rocks (Plate 12, Fig. 1).
However, in the extreme headwaters of most western tributaries
of the Little Blue River (in Washington and RepubHc counties)
sandy bottoms predominate. The Black Vermillion River flows
on a broad floodplain and is a mud-bottomed, sluggish stream,
with an average gradient of approximately one foot per mile.
Fringe-forests of elm, cottonwood, sycamore, and willow persist
along most of these stream-courses.

Notwithstanding the mud bottoms, the water in this kind of
stream in the Big Blue Basin remains clearer than that of the Big
Blue and Little Blue rivers. Heavy algal blooms were noted in
the Black Vermillion River and Mill Creek, Washington County,
in 1957 and 1958. Temperatures at Stations 45-M and 46-M on
Mill Creek, Washington County, averaged 85.5° F. on July 31, 1957.
Chemical characteristics of a water-sample from Station 35-M,
Black Vermillion River, are in Table 1.

Gravel-bottomed streams. — Most streams of this kind are tribu-
tary to the Big Blue River; however, streams entering Black Ver-
million River from the south are also of tliis type (Plate 12, Fig. 2).
The streams are "characteristically a series of large pools (to 100
feet in length and more than two feet in depth ) connected by short
rifiles and smaller pools" (Minckley and Cross, in press). The
average gradients are high: Carnahan Creek, 33 feet per mile;
Mill Creek, Riley County, 21 feet; Clear Creek, 16 feet per mile.
Stream-flow is usually less than five cubic feet per second. In sum-
mer, these streams may become intermittent, but springs and

410 University of Kansas Publs.^ Mus. Nat. Hist.

subsurface percolation maintain pool-levels (Minckley and Cross,
loc. cit.).

The average temperatures of these small streams (79.5° to
81.0° F. in July and August, 1957 ) were lower than temperatures
in stream-types previously described. Turbidities were usually
less than 25 p. p. m. The chemical properties of water-samples from
two of these streams (Stations 11-G and 18-G) are listed in Table 1.

PREVIOUS RECORDS OF FISHES

The earliest records of fishes from the Big Blue River Basin are those of
Cragin (1885) and Graham (1885) in independently published lists of the
fishes of Kansas. Meek (1895) recorded fishes collected in 1891 "from both
branches of the Blue River, a few miles west of Crete, Nebraska." Evcrmann
and Cox (1896) reported five collections from the Nebraskan part of the
basin. Their collections were made in October, 1892, and August, 1893, and
the stations were: in 1892, Big Blue River at Crete; in 1893, Big Blue River
at Seward, Lincoln Creek at Seward and York, and Beaver Creek at York.

Canfield and VViebe (1931) obtained fish from 18 localities in Nebraska
in July, 1930; however, their major concern was determination of water quality.
Their stations were: Big Blue River at Stromsburg, Polk Co.; Surprise and
Ulysses, Butler Co.; Staplehurst, Seward, and Milford, Seward Co.; Crete and
Wilber, Saline Co.; Beatrice, Blue Springs, and Barnston, Gage Co.; Little Blue
River at Fairbury, Jefferson Co.; Hebron, Thayer Co.; Sandy Creek at Alex-
andria, Thayer Co.; West Fork of Big Blue River at Stockham, Hamilton Co.;
McCool Junction, York Co.; Beaver Crossing, Seward Co.; and Beaver Creek
at York, York Co.

Breukelman ( 1940 ) and Jennings ( 1942 ) listed fishes from the University
of Kansas Museum of Natural History and the Kansas State College Museum,
respectively, including some specimens collected from the Big Blue River
System in Kansas. Because records in these two papers pertain to collections
that were widely spaced in the basin and in time, the specific localities are not
given herein. One of Jennings' ( loc. cit. ) records, Scaphirhynchus platorynchus
(Rafinesque), was cited by Bailey and Cross (1954:191). More recently,
Minckley and Cross (in press) recorded several localities, and cited some
papers mentioned above, in a publication dealing vnth Notropis topeka
(Gilbert) in Kansas.

Information on the fishes of the Nebraskan portion of the Big Blue River
Basin was compiled, and additional localities were reported, in a doctoral
thesis by Dr. Raymond E. Johnson, entitled The Distribution of Nebraska
Fishes, 1942, at the University of Michigan.

METHODS AND MATERIALS

Collection of Fishes

The gear and techniques used are listed below:

Entrapment Devices. — Hoop and fyke nets and wire traps were used
for 288 trap/net hours in 1957. The nets were not baited, and were set
parallel to the current, with the mouths downstream. Hoop nets were IJi to
three feet in diameter at the first hoop, with a pot-mesh of one inch; fyke nets

Fishes of Big Blue Basin 411

were three feet at the first hoop, pot-mesh of one inch; wire traps, with an
opening at each end, were 2Ji feet in diameter and covered with one-inch-mesh,
galvanized chicken wire.

Gill Nets. — Experimental gill nets were set on three occasions in areas
with little current. These nets were 125 feet in length, with % to two inch
bar-mesh in 25-foot sections.

Seines. — Seining was used more than other methods. An attempt was
made to seine all habitats at each station. In swift water, seine-hauls were
usually made downstream, but in quiet areas seining was done randomly.
Haul-seines six to 60 feet in length, three to eight feet in depth, and with
meshes of Js to H inch were used. For collection of riffle-fishes, the seine was
planted below a selected area and the bottom was kicked violently by one
member of the party, while one or two persons held the seine, raising it when
the area had been thoroughly disturbed. Seining on riffles was done with a
four-foot by four-foot bobbinet seine.

RoTENONE. — Rotenone was used in pools of smaller streams, mouths of
creeks, borrow-pits, and cut-off areas. Both powdered and emulsifiable
rotenone were used. The rotenone was mixed with water and appUed by
hand, or into the backwash of an outboard motor.

Electric Shocker. — The electrical unit used in this study generated 115
volts and 600 to 700 watts, alternating current. The shocking imit consisted
of two booms, each with two electrodes, mounted on and operated from a
slowly moving boat. Fish were recovered in scape nets, or in many cases
were identified as they lay stunned and were not collected.

Estimation of Relative Abundance

Data on relative abundance of fishes were obtained by counts of seine
hauls at 29 of the 59 stations, counts of rotenoned fish at seven stations, and
results with the electric shocker at nine stations. Counts were usually made
in the field; however, in some collections all fish were preserved and counted
in the laboratory. Some fish (or "swirls" presumed to be fish) observed
while shocking were not identified and are not included in the calculations.
However, all fish positively identified while shocking are included.

Age and Growth of Fishes

Fish from selected size-gi'oups were aged in this study. Scales for age-
determinations were removed from positions recommended by Lagler (1952:
108). Scales were placed in water between glass slides and were read on a
standard scale-projection device.

Pectoral spines of catfish were removed from one or both sides, sectioned,
and read by methods described by MarzoLf (1955:243-244).

Calculation of length at the last annulus for both scale-fish and catfish was
made by direct proportion. All measurements are of total length to the nearest
tenth of an inch unless specified otherwise.

Creel Census

From April 6 to May 28, 1957, a creel census was taken below Tuttle Creek
Dam. From June 16 to July 24, 1958, I periodically visited the main points of
access to the Big Blue River, beginning approximately eight miles downstream
from Tuttle Creek Dam and ending six miles upstream from the maximal ex-

412

University of Kansas Publs., Mus. Nat. Hist.

tension of the reservoir at capacity level. Access-points consisted of 11 bridges,
two power dams, and three areas where county roads approached the river.
Eleven eight-hour da\'s were spent in the 1957 census and 22 checks in 15 days
were made in 1958. An equal number of morning (6:00 a. m. to 12:00 noon)
and afternoon (12:00 noon to 8:30 p.m.) checks were made.

Fishermen contacted were asked the following questions: home address (or
residence at the time of the fishing trip); time they started fishing; kind of fish
sought; number and kinds of fish in possession; and baits used. Also, the
number of poles and ty-pe of fishing (from the bank, from boat, etc.) were
recorded. Fishes caught were examined to confirm identifications. About 80
per cent of all fishermen seen were contacted.

Fish per man-hour, as used in this report, refers to the average number of
fish of all species caught by one fisherman in one hour. Fisherman-day is tlie
average time spent fishing in one day by one person. Because some fishermen
used more than one pole, the data are also expressed as catch per pole-hour.

COLLECTING STATIONS

In the list that follows, stations are numbered consecutively from the mouth
of the Big Blue Ri\er, listing stations on each tributary as it is ascended. The
letters following station-numbers indicate the general type of stream: S =
sandy; M = muddy; and G = gravelly. The Big Blue River is the boundary
between Riley and Pottawatomie counties, Kansas, along part of its length.
Stations in this area have been designated Riley County. The legal description
of each station is followed by the date(s) of collection, and each station is
plotted in Figure 2.

10 0 10 20
I' 1 1

scale of miles

L

15

Index Mop

J L

• Vr/ ^ix \iZ/ \!3' \LiJj3' \Z/ \ly V' *•/ v^j-v v^ i >6J/ v^

96

Fig. 2. Collection stations in the Big Blue River Basin, Kansas, 1957 and 1958.

PLATE 11

Fig. 1. Bitf Blue River at Station o-S. U. S. Army Corps oi Eiigiinns iilioto-

graph No. 563697.

Fig. 2. Big Blue River at Oketo, Marshall County, Kansas. U. S. Army Corps
of Engineers, photograph No. 67516.

PLATE 12

Fi(.. 1. lilatk Vcnnillioii Ri\fr, appro\iiiiat(l\- one mile uiistri'am iioiii its
nioiith. Pluitograph by Uohcrt G. Wehh.

^.".i

•i

„f^

i

Fic. 2. Carnaliaii (^ii'ik at Station ll-(;. I'hotogiapli In Holxrt i'.. Webb.

Fishes of Big Blue Basin 413

1-S: Pottawatomie Co., mouth of Big Blue River, Sec. 16, T. lOS, R. 8E,

June 20, 1933.
2-S: Riley Co., Big Blue River, Sec. 4, T. lOS, R. 8E, June 6, 12, and 14,

1957.
3-S: Riley Co., Big Blue River, E ¥z. Sec. 30, T. 9S, R. 8E, Mar. 30, Apr. 6,

July 15, 16, 17, Aug. 14, and Dec. 26, 1957; Apr. 26, June 20, and Aug.

5, 1958.
4-S: Riley Co., Big Blue River at Rocky Ford Dam, W )^, Sec. 30, T. 9S,

R. 8E, Aug. 14, 1957; and Aug. 5, 1958.
5-G: Pottawatomie Co., Mclntire Creek, Sec. 12, T. 9S, R. 7E, July 14, 1958.
6-S: Riley Co., Big Blue River and adjacent borrow-pit, Sec. 24, T. 9S, R.

7E, July 18 and 19, 1957; and July 11, 1958.
7-G: Riley Co., Tuttle Creek, Sec. 10, T. 9S, R. 7E, Aug. 5, 1958.
8-S: Riley Co., Big Blue River, Sec. 10, T. 9S, R. 7E, Aug. 14, 1957.
9-G: Riley Co., Mill Creek, Sec. 4, T. 9S, R. 7E, July 20 and 25, 1958.
10-G: Riley Co., Mill Creek, Sec. 2, T. 9S, R. 6E, Aug. 13, 1957.
11-G: Pottawatomie Co., Carnahan Creek, Sec. 22, 27, and 34, T. 8S, R. 7E,

Aug. 1, 1957; and July 8, 1958.
12-G: Pottawatomie Co., unnamed tributary to Carnahan Creek, Sec. 15, T.

8S, R. 7E, Mar. 19, 1956 (collection made before my formal study was

begun ) .
13-G: Pottawatomie Co., Carnahan Creek, Sec. 36, T. 7S, R. 7E, Aug. 13,

1957.
14-S: Riley Co., Big Blue River, Sec. 18, T. 8S, R. 7E, Mar. 22, 1958.
15-S: Riley Co., Big Blue River, Sec. 7, T. 8S, R. 7E, Apr. 3, and June 12,

1958.
16-G: Riley Co., unnamed creek. Sec. 1, T. 8S, R. 6E, July 10, and Aug. 5,

1958.
17-G: Riley Co., unnamed creek. Sec. 10, T. 8S, R. 6E, June 26, 1958.
18-G: Riley Co., Fancy Creek, Sec. 14, T. 7S, R. 6E, July 29, 1957.
19-G: Riley Co., Walnut Creek, Sec. 20, T. 7S, R. 6E, June 26, 1958.
20-G: Riley Co., Fancy Creek, Sec. 2, T. 7S, R. 5E, Mar. 13, 1957; and June

26, 1958.
21-G: Riley Co., Schoolhouse Branch, Sec. 35, T. 6S, R. 5E, July 22, 1958.
22-G: Riley Co., Fancy Creek, Sec. 33, T. 6S, R. 5E, June 1, 1957.
23-G: Riley Co., West Branch Fancy Creek, Sec. 32 and 33, T. 6S, R. 5E, June

1 and 3, 1957.
24-G: Clay Co., West Branch Fancy Creek, Sec. 32 and 33, T. 6S, R. 4E, July

22, 1958.
25-S: Riley Co., Big Blue River, Sec. 5, T. 7S, R. 7E, Aug. 7, 1958.
26-G: Riley Co., Swede Creek, Sec. 21, T. 6S, R. 7E, Mar. 22, 1958.
27-G: Pottawatomie Co., unnamed creek. Sec. 14, T. 6S, R. 7E, Sept. 10, 1957.
28-G: Pottawatomie Co., Bluff Creek, Sec. 6, T. 6S, R. 8E, Oct. 6, 1957.
29-G: Pottawatomie Co., Bluff Creek, Sec. 15, T. 6S, R. 8E, June 29, 1958.
30-M: Marshall Co., Black Vermillion River, Sec. 9, T. 5S, R. 8E, Mar. 5,

1958.
31-G: Pottawatomie Co., Clear Creek, Sec. 3, T. 6S, R. 9E, July 14, 1958.
32-G: Pottawatomie Co., unnamed creek. Sec. 14, T. 6S, R. 9E, July 14, 1958.
33-M: Marshall Co., Robidoux Creek, Sec. 20, T. 2S, R. 9E, July 23, 1958.
34-M: Marshall Co., Litde Timber Creek, Sec. 10, T. 4S, R. 9E, Oct. 6, 1957.
35-M: Marshall Co., Black Vermillion River, Sec. 15, T. 4S, R. 9E, Aug. 9,

1958.
36-M: Marshall Co., unnamed creek. Sec. 8, T. 4S, R. 9E, Oct. 6, 1957.

414 Unin'ersity of Kansas Publs., Mus. Nat. Hist.

37-M: Marshall Co., Black Vermillion River, Sec. 11, T. 4S, R. lOE, Oct. 6,
1957.

38-S: Marshall Co., Big Blue River, Sec. 18, T. 5S, R. 8E, Aug. 8, 1958.
39-S: Marshall Co., Big Blue River, Sec. 20, T. 4S, R. 7E, xMay 29, 1958.
40-M: Washington Co., Coon Creek, Sec. 27, T. 4S, R. 4E, July 22, 1958.
41-S: Marshall Co., Little Blue River, Sec. 9, 16, and 17, T. 4S, R. 6E, June

27, 1958.
42-S: Washington Co., Little Blue River, Sec. 21, T. 3S, R. 5E, Aug. 8, 1958.

43-S: Washington Co., Little Blue River, Sec. 5 and 8, T. 3S, R. 5E, July 30,
1957.

44-S: Washington Co., Little Blue River, Sec. 36, T. IS, R. 4E, July 31, 1957.

45-M: Washington Co., Mill Creek, Sec. 35 and 36, T. IS, R. 4E, July 31,
1957.

46-M: Washington Co., Mill Creek, Sec. 4, T. 2S, R. 4E, July 31, 1957.

47-M: Washington Co., Spring Creek, Sec. 11 and 12, T. 2S, R. 3E, June 19,
1958.

48-M: Washington Co., Mill Creek, Sec. 28, T. 2S, R. 2E, June 19, 1958.

49-M: Repubhc Co., Mill Creek, Sec. 8 and 17, T. 2S, R. IW, July 23, 1958.

50-S: Washington Co., Little Blue River, Sec. 5, T. IS, R. 4E, Aug. 9, 1958.

51-M: Republic Co., Rose Creek, Sec. 20, T. IS, R. 2W, July 23, 1958.

52-S: Marshall Co., Big Blue River, Sec. 6, T. 4S, R. 7E, Aug. 6, 1958.

53-S: Marshall Co., Big Blue River, Sec. 18, T. 3S, R. 7E, July 29 and 30,
1957; May 28, and Aug. 6, 1958.

54-G: Marshall Co., Hop Creek, Sec. 13 and 18, T. 3S, R. 7E, May 28, 1958.
55-M: Marshall Co., Spring Creek, Sec. 29, T. 2S, R. 8E, July 9, 1958.
56-S: Marshall Co., Big Blue River at Marysville Dam, Sec. 20, T. 2S, R. 7E,
June 16, 1958.

57-M: Marshall Co., Horseshoe Creek, Sec. 6, T. 2S, R. 7E, July 1, 1958.
58-G: Marshall Co., unnamed creek, Sec. 2, T. IS, R. 7E, July 1, 1958.
59-G: Marshall Co., Mission Creek, Sec. 3, T. IS, R. 8E, Nov. 30, 1957.

ANNOTATED LIST OF SPECIES
Forty-eight species were obtained in this survey and five others
have been recorded in hterature or are deposited in museums:
KSC = Kansas State College Museum; and UMMZ = University
of Michigan Museum of Zoology. Specimens, unless designated
otherwise, are in the University of Kansas Museum of Natural
History (KU).

In this list, the scientific name of each species is followed by
the common name, citations of previous records, and the stations
where the species was obtained. I follow Bailey (1956:328-329)
in treating Lepisosteus osscus (Linnaeus), Catostomus commer-
sonnii (Laccpcde), Semotilus atromaculatus (Mitchill), Notropis
lutrensis (Baird and Girard), Pimcphales promclas Rafinesque, Ic-
talurus melas (Rafinesque), Ictalurus punctatus (Rafinesque), and
Lepomis macrochinis Rafinesque, in binomial form only.

Scaphirhynchtts platorynchus (Rafinesque), shovelnose sturgeon:

Fishes of Big Blue Basin 415

Jennings (1942:364) as Scaphirhynchus platorhynchus (Rafi-
nesque); Bailey and Cross (1954:191). Stations 3-S and 4-S.

Shovelnose sturgeon were found only in the lower portion of the
Big Blue River. On April 20, 1957, many were seen in fishermen's
creels at Stations 3-S and 4-S. One male and two females that I
examined on that date were ripe or nearly so; eggs seemed well
developed and milt flowed freely from the male. After April, 1957,
none was collected or observed until April 26, 1958, when one
specimen was obtained while shocking. Forbes and Richardson
(1920:27) reported that shovelnose sturgeon spawn in Illinois
between April and June, and Eddy and Surber (1947:80) reported
spawning in May and early June in Wisconsin and Minnesota.

Lepisosteus platostomus Rafinesque, shortnose gar: Jennings
(1942:364). Stations 3-S and 4-S.

I saw shortnose gar at various times in 1956 and 1957 at Rocky
Ford Dam on the Big Blue River (Station 4-S). One was seen
while shocking at Station 3-S on December 26, 1957.

Lepisosteus osseus (Linnaeus), longnose gar: Jennings (1942:
364) as Lepisosteus osseus oxyurus Rafinesque. Stations 1-S, 2-S,
3-S, 4-S, 6-S, 8-S, 9-G, 15-S, 18-G, 25-S, 41-S, 44-S, 52-S, and 53-S.

Longnose gar were abundant in the mainstream of the Big Blue
River but usually evaded capture. This species, and the shortnose
gar, resided in the larger rivers, with L. osseus being taken in only
two creeks near their mouths. In periods of high water, gar moved
into the flooded creeks, but returned to the river as stream-levels
subsided.

Young-of-the-year L. osseus, averaging 21.5 mm. in total length
(range 13 to 30 mm.), were taken on June 14, 1957, and larger young
(estimated 60 to 70 mm. total length) were taken on June 27, 1958.

Dorosoma cepedianum (LeSueur), gizzard shad: Jennings
(1942:364). Stations 1-S, 3-S, 4-S, 6-S, 8-S, 44-S, 45-M, and 53-S.

Most gizzard shad were young-of-the-year, taken on July 16 and
17, 1957, at Stations 3-S and 4-S. Twenty specimens from Station
6-S that were in their second summer of life were from 3.8 to 5.9
inches total length at the last annulus ( average 4.3 ) . This species
was usually found in quiet water and was most abundant near the
mouth of the Big Blue River.

Hiodon alosoides (Rafinesque), goldeye. Stations 3-S, 4-S, and
53-S.

I caught five specimens of H. alosoides from the Big Blue River,

416 Unfv'ersity of Kansas Publs., Mus. Nat. Hist.

and another specimen, obtained by Dr. R. B. Moorman in 1954, is
at Kansas State College (KSC 4984).

One goldeye that I caught on April 20, 1956, prior to the be-
ginning of my study, was a ripe female measuring 15.5 inches total
length. The fish was beginning its seventh summer of Hfe.

Cycleptus elongatus LeSueur, blue sucker. The blue sucker is
included on the basis of a single specimen (KSC 2917) collected
by I. D. Graham and labeled "Blue River." No other data are with
the specimen; however, most fishes deposited at Kansas State Col-
lege by Graham are dated "1885" or "1886" and were caught near
"Manhattan" (Riley County).

Ictiobus cyprinella (Valenciennes), bigmouth buffalo. Stations
3-S, 6-S, and 30-M.

Bigmouth buffalo were rare, and were taken only in quiet parts
of larger streams, and in the borrow-pit at Station 6-S.

Ictiobus niger (Rafinesque), black buffalo. Stations 3-S, 41-S,
and 53-S.

Only four individuals of 7. niger were taken. All were large
adults (more than 20 inches in total length), and all were shocked
in the deeper, swifter areas, where the channel narrowed.

Ictiobus bubalus (Rafinesque), smallmouth buffalo. Stations
1-S, 3-S, 6-S, 7-G, 18-G, 38-S, 41-S, 43-S, 46-M, and 53-S.

This species was found in relatively quiet waters in the main
channel, in cut-off areas, and in creek-mouths. The ages and total
lengths of 30 individuals obtained at Station 6-S were (average
followed by number of fish in parentheses): I, 2.4 (11); II, 4.4
(14); and III, 6.6 (5).

Canfield and Wiebe (1931:6-7, 10) recorded 'buffalo-fish" and
"buffalo" from the Big Blue Basin in Nebraska; however, no spe-
cific designation was given.

Carpiodes forbesi Hubbs, plains carpsucker. Station 3-S.

This represents the first record known to me of the plains carp-
sucker from Kansas. The specimen (KU 4180), 430 mm. in stand-
ard length, has the following characters: lower lip without a
median, nipple-like projection; dorsal fin-rays, 25; lateral-line scales,
38; diameter of orbit into distance from anterior nostril to tip of
snout, 1.1; body-depth into standard length, 3.3; and head-length
into standard length, 3.9. The specimen was taken while shocking
a wide, shallow channel, over sand bottom.

Carpiodes carpio carpio (Rafinesque), river carpsucker: Jen-
nings (1942:364). Stations 1-S, 2-S, 3-S, 4-S, 5-G, 6-S, 7-G, 8-S,

Fishes of Big Blue Basin 417

9-G, 11-G, 14-S, 15-S, 18-G, 19-G, 23-G, 25-S, 27-G, 28-G, 30-M,
38-S, 39-S, 41-S, 42-S, 43-S, 44-S, 45-M, 50-S, 52-S, and 53-S.

The river carpsucker occurred at most stations on the larger
streams, and in many of the smaller tributaries. In smaller streams
C. c. carpio frequented the largest pools, in or near the floodplains
of larger streams. A marked preference for still water, soft, silty
bottoms, and areas with drift or other cover was apparent; however,
the species also occurred in open waters with moderate to swift
currents.

The sizes attained by the river carpsucker at different ages were
(averages followed by number of fish in parentheses): I, 1.9 (10);
II, 3.9 (5); III, 5.3 (8); IV, 7.7 (5); V, 11.9 (2); VI, 11.6 (7);
VII, 12.8 (6); VIII, 13.1 (1); IX, 14.9 (2); X, 15.8 (8); and XI,
17.6 ( 1 ) . These averages are significantly less than those reported
by Buchholz (1957:594) for the river carpsucker in the Des
Moines River, Iowa.

Examination of the gonads of river carpsucker in summer, 1957,
indicated that spawning occurred in late July. Young-of-the-year,
averaging 21 mm. in total length, first appeared in my collections
on July 30, 1957.

Carpiodes velifer (Rafinesque), highfin carpsucker: Meek
(1895:135); Evermann and Cox (1896:389).

The highfin carpsucker was not taken in my survey. Meek
(1895:135) reported "this small sucker [C. velifer] . . . com-
mon in Blue River at Crete," characterizing the specimens as hav-
ing "Dorsal rays, 24 to 30; scales in the lateral-line, 36 to 41; head
3^ to 4; and depth 2/2 to 3." The ranges in the number of dorsal
rays and the number of scales in the lateral-line are higher than
usual in C. velifer, or in C. c. carpio, which is now common in the
Big Blue River Basin. Both species normally have 33 to 37 lateral-
line scales and 27 or fewer dorsal rays (Bailey, 1956:352-353;
Moore, 1957:79; and Trautman, 1957:81-82). The other charac-
ters listed by Meek would fit the young and some adults of either
species, or possibly a composite including C. forbesi.

Graham (1885:72) and Cragin (1885:107) reported Ictiobm
velifer {=: Carpiodes velifer) from "Eureka Lake," Riley County,
Kansas. This lake, which no longer exists, was in the Kansas River
Valley, about ten miles upstream from the mouth of the Big Blue
River, Other, more recent records from the Kansas River Basin,
in the vicinity of the Big Blue River, are: Maple Leaf Lake, Riley

2—7080

418 University of Kansas Publs., Mus. Nat. Hist.

Co., Oct. 4, 1925; Deep Creek, Riley Co., no date; Wildcat Creek,
Riley Co., Sept. 7, 1923; and Wildcat Creek, Riley Co., Sept. 29,
1925 (UMMZ 122187-90). Most of the collections were made by
Minna E. Jewell (Nelson, personal communication).

Moxostoma aureolwn (LeSueur), northern redhorse: Cragin
(1885:108) as Moxostoma macrolepidottun LeSueur; Meek (1895:
136) as Moxostoma macrolepidofvm dtiqiiesnei (LeSueur); Ever-
mann and Cox (1896:394-395); and Jennings (1942:364) as Mox-
ostoma erythrurum (Rafinesque). Stations 41-S, 43-S, 44-S, and
53-S.

I collected three northern redhorse from the Big Blue River
Basin, and another specimen was seined in the mouth of Mill Creek,
Riley County (my present Station 9-G) by the Kansas State Col-
lege class in fisheries management in 1954 (KSC 5068). I reidentify
as M. aureolum the two specimens recorded by Jennings ( loc. cit. )
as M. erythrurum.

The subspecific status of M. aureolum in the Kansas River Basin is
to be the subject of another paper.

Catostomus commersonnii (Lacepcde), white sucker: Canficld
and Wiebe (1931:8) as "common suckers"; and Breukelman (1940:
380). Stations 7-G, 11-G, 12-G, 13-G, 16-G. 18-G, 19-G, 23-G,
29-G, 31-G, 53-S, 57-M, and 58-G.

The white sucker occurred primarily in upland streams of the
Flint Hills, with one occurrence in muddy habitat, and one in the
main stream of the Big Blue River. Young C. comtnersonnii were
often taken in riffles, but adults were in the larger, deeper pools.
The ages and total lengths at the last annulus for 12 white suckers
were: 1,2.8 (4); 11,3.9 (6); HI, 8.2 (1); and IV, 9.2 (1).

Cyprinus carpio Linnaeus, carp: Canfield and Wiebe (1931:
5-8, 10) as "carp." Stations 1-S, 2-S, 3-S, 4-S, 6-S, 7-G, 8-S, 15-S,
16-G, 18-G, 23-G, 24-G, 25-S, 27-G, 30-M, 35-M, 38-S, 41-S, 42-S,
43-S, 44-S, 45-M, 52-S, 53-S, and 56-S.

Carp occurred throughout the basin. The habitat of this spe-
cies closely approximated that of the river carpsucker; however,
carp were more often taken in moderate to swift water than were
C. c. carpio.

The ages and average lengths at the last annulus for 40 carp from
the Big Blue River Basin were: 1,2.3 (4); 11,4.7 (10); HI, 7.0 (10);
IV, 9.0 (3); V, 11.3 (4); VI, 18.6 (1); VIT, 18.9 (3); VIII, no fish;
IX, 20.6 (3); X, 19.1 (2); XI, 21.1 (1); XII, 22.0 (1); and XIII,
24.1 (2).

Fishes of Big Blue Basin 419

Carassius auratus (Linnaeus), goldfish. Station 4-S.

I saw goldfish seined from Station 4-S by anglers obtaining bait
on April 20, 1957. Goldfish were commonly used for bait at Sta-
tions 4-S and 54-S.

Semotilus atromaculatus (Mitchill), creek chub: Evermann and
Cox (1896:399); and Jennings (1942:364) as Semotilus atromacu-
latus atromaculatus (Mitchill). Stations 5-G, 7-G, 10-G, 11-G,
12-G, 13-G, 16-G, 17-G, 18-G, 23-G, 24-G, 27-G, 28-G, 29-G, 31-G,
32-G, 33-M, 34-M, 36-M, 37-M, 40-M, 46-M, 47-M, 48-M, 49-M,
50-S, 53-S, 54-G, 55-M, 56-S, 57-M, 58-G, and 59-G.

Creek chubs were found in all habitats in the Big Blue River
Basin, but were abundant only in the headwaters of muddy streams
and in clear upland creeks.

Chrosomus erythrogaster (Rafinesque), southern redbelly dace:
Jennings (1942:365). Stations 11-G, 12-G, 13-G, 16-G, 27-G, 29-G,
and 53-S.

This colorful species occupied the headwaters of the clear,
spring-fed creeks where it was abundant. Only one specimen was
taken in muddy or sandy habitat ( at the mouth of a small creek at
Station 53-S ) , where it may have been washed by floods just prior
to my collecting.

Hybopsis storeriana (Kirtland), silver chub. Station 3-S.

One specimen of H. storeriana (KU 3810) was seined in swift
water near a sandbar on April 6, 1957, and another was taken at
the same locality on April 26, 1958.

Hybopsis aestivalis (Girard), speckled chub: Meek (1895:137);
and Evermann and Cox (1896:409), both as Hybopsis hyostomus
Gilbert. Stations 3-S, 4-S, 14-S, 25-S, 38-S, 39-S, 50-S, and 56-S.

This species was restricted to wide, swift parts of the Big Blue
and Little Blue rivers, and was found over clean, sometimes shifting,
sand bottoms. On May 29, 1958, three males in breeding condi-
tion were collected and on June 16, 1958, a large series of both
male and female H. aestivalis, all with well-developed gonads, was
collected. The water temperature was 77.0° F. Hubbs and Orten-
burger (1929:25-26) reported that Extrarius tetranemus (Gilbert)
(=: Hybopsis aestivalis tetranemus) spawns in summer especially
in early July. Cross (1950:135) reported a single pair of H. a.
tetranemus that he considered in breeding condition on June 9, 1948.

Breukelman (1940:380) recorded speckled chubs in the Kansas
River Basin as Extrarius {= Hybopsis) aestivalis: sesquialis X
tetranemus; however, the name sesquialis is a nomen nudum, and

420 Uxn-ERSiTY OF IC\NSAS PuBLS., Mus. Nat. Hist.

the status of this species in the Kansas River Basin is yet to be
elucidated.

Phenacobius mirabilis (Girard), plains suckermouth minnow:
Meek (1895:136); and Evennann and Cox (1896:408). Stations
2-S, 3-S, 4-S, 5-G, 6-S, 7-G, 8-S, 9-G, 11-G, 16-G, 18-G, 25-S, 26-G,
27-G, 35-M, 38-S, 39-S, 40-M, 42-S, 47-M, 50-S, 52-S, 53-S, 54-G,
and 56- S.

Fhenacohiiis mirabilis was widespread in the basin, occurring
most frequently on rifHes over bottoms of clean sand or gravel.
Young-of-the-year were usually taken in backwaters.

Notropis percobromus (Cope), plains shiner. Stations 3-S and

4-S.

The plains shiner occurred only in the lower part of the main
stream of the Big Blue River.

Notropis rubellus (Agassiz), rosyface shiner. Station 5-G.

One rosyface shiner ( KU 4195 ) was taken. This species was pre-
viously reported from only two localities in the Kansas River Basin:
in the Mill Creek Watershed, Wabaunsee County, and Blacksmith
Creek, Shawnee County as Notropis rtibrifrons (Cope) (Gilbert,
1886:208). Mill Creek and Blacksmith Creek are northward-flow-
ing tributaries of the Kansas River that arise in the Flint Hills.
Graham (1885:73) also recorded N. rubellus (as N. rubrifrons)
from the "Kansas and Missouri Rivers"; however, I suspect that his
specimens were Notropis percobromus, a species not generally
recognized in Graham's time (see Hubbs, 1945:16-17). Notropis
rubellus is now abundant in the Mill Creek Watershed ( Wabaunsee
County), but, except for my specimen No. 4195, has not been taken
recently in other streams in the Kansas River Basin.

Notropis umbratilis umbratilis (Girard), redfin shiner. Station
3-S.

One specimen of N. u. umbratilis was captured near a sandbar
on March 26, 1958. The absence of this species in Flint Hills
streams of the Big Blue River Basin is unexplained; redfin shiners
occur commonly in southern tributaries of the Kansas River both
upstream and downstream from the mouth of the Big Blue River.
In Kansas this species is usually associated with tlie larger pools
of clear, upland streams.

Canfield and Wiebe (1931:6-8) may have referred to this species
in recording "black-fin minnows" from the Nebraskan portion of
the Big Blue River Basin.

Fishes of Big Blue Basin 421

Notropis cornutus frontalis (Agassiz), common shiner. Stations
4-S, 5-G, 7-G, 10-G, 11-G, 12-G, 13-G, 18-G, 22-G, 26-G, 27-G, 28-G,
29-G, 31-G, 32-G, and 59-G.

Common shiners were most abundant in middle sections of the
clear, gravelly creeks.

Notropis lutrensis (Baird and Girard), red shiner: Meek (1895:
136); and Evermann and Cox (1896:404-405). All stations ex-
cepting 1-S, 17-G, 30-M, and 51-M.

Red shiners were the most widespread species taken in my sm*-
vey, occurring in all habitats, and in all kinds of streams. On two
occasions I observed what apparently was spawning behavior of
this species. Both times the specimens collected were in the
height of breeding condition, stripping in the hand easily, and
often without pressure. At the first locality (Station 29-G) no at-
tempt was made to obtain eggs, but by disturbing the bottom at
the second (55-M) I found eggs that were thought to be those of
red shiners. The eggs were slightly adhesive, clinging to the hand
and to the bobbinet seine.

On June 29, 1958, at Station 29-G, red shiners appeared to be
spawning in an open-water area measuring about 15 by 15 feet,
over nests of Lepomis cyanellus Rafinesque and L. humilis ( Girard ) .
No interspecific activity was noted between the sunfish and the red
shiners. Water temperature at this station was 73.4° F., and the
bottom was gravel, sand, and mud. Observations were made from
a high cut-bank, by naked eye and by use of 7-X binoculars.

The red shiners moved rapidly at the surface of the water, with
one male (rarely two or more) following one female. The male
followed closely, passing the female and causing her to change
direction. At the moment of the female's hesitation, prior to her
turn, the male would erect his fins in display, at the side and a little
in front of the female. After brief display, usually less than two
seconds, the male resumed the chase, swimming behind and around
the female in a spiral fashion. After a chase of two to three feet,
the female would sometimes allow the male to approach closely on
her left side. The male nudged the female on the caudal peduncle
and in the anal region, moving alongside with his head near the
lower edge of the left operculum of the female, thus placing his
genital pore about a head-length behind and below that of the
female. At this time spawning must have occurred; however,
possibly because of the speed of the chase, I observed no vibration
of the fish as described for other species of Notropis at the culmina-

422 University of Kansas Publs., Mus. Nat. Hist.

tion of spawning (Pfeiffcr, 1955:98; Raney, 1947:106; and others).
While the spawning act presumably occurred the pair was in
forward motion in a straight course, for three to five feet, at the
end of which the male moved rapidly away, gyrating to the side
and down. The female then swam away at a slower rate. In
instances when the female failed to allow the male to move along-
side, the male sometimes increased his speed, striking the female,
and often causing her to jump from the water.

Some conflict between males was observed, usually when two
or more followed one female. The males would leave the female,
swerve to one side, and stop, facing each other or side by side. At
this moment the fins were greatly elevated in display. There was
usually a rush on the part of one male, resulting in the flight of
the other, and the aggressive male would pursue for about two feet.
Many times the pursued male jumped from the water.

At Station 55-M, on July 9, 1958, activity similar to that described
above was observed in a small pool near a mass of debris. At this
station I watched from the bank, three feet from the spawning
shiners. Water temperature was not recorded.

The minnows performed the same types of chase and display,
all in open water, as described for Station 29-G. However, at Sta-
tion 55-M, much activity of males occurred near the small deposit
of debris. It seemed that conflict was taking place, with males be-
having as described above, and milling violently about. Examina-
tion of the area revealed nests of L. cijanellus near the debris, and
some of the activity by the shiners may have been raids on nests of
the sunfish. However, females nearing the group of males were
immediately chased by one to four individual males, with one
usually continuing pursuit after a short chase by the group. The
male again moved into position at the lower left edge of the oper-
culum of the female as at Station 29-G.

Another kind of behavior was observed also, in which the
female sometimes stopped. The male approached, erecting his
fins and arching his body to the left. The female also assumed this
arch to the left, and the pair moved in a tight, counter-clockwise
circle, with the male on the inside. After a short period in this
position, the male moved aside in display, and gyrated to the side
and down. Females at both stations moved about slowly, usually
remaining in the immediate vicinity of activity by males, and re-
turning to the area even when pursued and deserted some distance
away.

Fishes of Big Blue Basin 423

Notropis deliciosus (Girard), sand shiner: Meek (1895:136);
Evermann and Cox ( 1896:402), both as Notropis blenniiis (Girard);
and Jennings (1942:365) as Notropis deliciosus missuriensis (Cope).
All stations excepting 1-S, 10-G, 12-G, 17-G, 20-G, 21-G, 22-G,
24-G, 29-G, 30-M, 31-G, 32-G, 33-M, 35-M, 51-M, 55-M, 57-M, 5S-G,
and 59-G.

Nelson (personal communication) has studied the sand shiner in
Kansas, and has found that the Big Blue River is an area of inter-
gradation between the southwestern subspecies ( deliciosus ) and the
plains subspecies {missuriensis). Notropis d. deliciosus prefers
cool, rocky habitat, and occurs in small streams of the Flint Hills,
whereas N. d. missuriensis occupies the sandy, turbid Big Blue and
Little Blue rivers. Intergrades occur most frequently in the Big
Blue River, but are found in all habitats.

Notropis topeka (Gilbert), Topeka shiner: Meek (1895:136);
Evermann and Cox (1896:403); and Minckley and Cross (in press).
Stations 10-G, 11-G, 12-G, 19-G, 31-G, and 32-G.

This species was common locally in the upland streams. Female
Topeka shiners stripped easily at Station 11-G on July 8, 1958, and
adult N. topeka in high breeding condition were collected at Sta-
tion 31-G on July 14, 1958. The water temperature at both stations
was 77.5° F. Evermann and Cox (1896:403-404) recorded female
Topeka shiners "nearly ripe" on June 29, 1893.

Notropis buchanani Meek, ghost shiner. Stations 3-S and 4-S.
Only two specimens of N. buchanani were taken, both on
August 14, 1957. These specimens (KU 3833), a female with
well-developed ova, and a tuberculate male, were near a sandbar
in the main channel. To my knowledge, this is the first published
record of the ghost sliiner from the Kansas River Basin. Mr. James
Booth, State Biological Survey, collected N. buchanani from two
stations on Mill Creek, Wabaunsee County, Kansas, 1953.

Hybognathus nuchalis Agassiz, silvery minnow. Stations 2-S,
3-S, 4-S, 7-G, 8-S, and 16-G.

This species was taken sporadically, but sometimes abundantly,
in the Big Blue River. At Stations 7-G and 16-G a few young-of-
the-year were found.

Bailey (1956:333) does not consider the southwestern Hybog-
nathus placita (Girard) specifically distinct from the northeastern
H. nuchalis, but little evidence of intergradation has been pub-
lished. In Table 2, I have compared measurements and counts

424

University of Kansas Publs., Mus. Nat. Hist.

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426 University of Kansas Publs., Mus. Nat. Hist,

of 50 specimens of Ilybognathus from the Big Blue River, 50 H. n.
placita from the Walnut River, Kansas (Arkansas River Basin),
and 50 H. n. niichaJis from Wisconsin. Measurements and counts
were made by methods described by Hubbs and Lagler ( 1947:8-15)
and measurements are exi^ressed as thousandths of standard length.

Hyhognathiis from the Big Blue River tend to have fewer, larger
scales than H. n. placita from the Walnut River, Kansas, but more
and smaller scales than H. n. niichalis from Wisconsin. In speci-
mens from the Blue River, tlie size of the orbit divided by standard
length, and the width of gape divided by standard length and
width of orbit, are also intermediate bet^veen the Walnut River and
Wisconsin specimens, but tend toward the fonner. Specimens from
the Big Blue River resemble H. n. placita from the Walnut Ri\er
in body shape, robustness, and in the embedding of scales on the
nape.

Pimephales notatus (Rafinesque), bluntnose minnow: Meek
(1895:136); and Evermann and Cox (1896:399). Stations 2-S, 3-S,
5-G, 6-S, 8-S, 9-G, 10-G, 11-G, 12-G, 13-G, 16-G, 19-G, 27-G, 29-G,
53-S, 54-G, and 58-G.

Tlie bluntnose minnow preferred the clearer creeks, with gravel
or gravel-silt bottoms, but occurred rarely in the mainstream of the
Big Blue River. Males and females in high breeding condition
were taken on July 14, 1958. The temperature of the water was
75.5° F.

Pimephales promelas Rafinesque, fathead minnow: Meek (1895:
136); and Evermann and Cox (1896:397-398). All stations ex-
cepting 1-S, 4-S, 12-G, 30-M, 43-S, 44-S, and 56-S.

Small muddy streams were preferred by P. promelas; however,
the fathead minnow was taken in all habitats, and in association
with most other species.

Canfield and Wiebe (1931:6-7) may have recorded P. promelas
from the Big Blue River Basin, Nebraska, as "blackhead minnows."

Campostoma anomalum plumheum (Girard), stoneroller. All
stations excepting 1-S, 2-S, 3-S, 14-S, 15-S, 21-G, 22-G, 28-G, 30-M,
33-M, 34-M, 35-M, 36-M, 37-M, 38-S, 41-S, 44-S, 45-M, 51-M,
52-S, and 55-M.

Stonerollers were usually taken in rifHes with gravel-rubble bot-
toms. Those individuals collected in areas with mud or sand
bottoms were almost invariably in the current, or in the edge of
currents.

Specimens from the Big Blue River Basin have an average of
47.4 scale-rows around the body (range 42-54).

Fishes of Big Blue Basin 427

Ictalurus melas (Rafinesque), black bullhead: Evermann and
Cox (1896:387) as Ameitmis melas (Rafinesque); and Canfield
and Wiebe (1931:5-7, 10) as "bullheads." Stations 2-S, 6-S, 7-G,
11-G, 16-G, 20-G, 22-G, 23-G, 24-G, 28-G, 35-M, 40-M, 51-M, 53-S,
55-M, 56-S, 57-M, and 58-G.

Black bullhead occurred in all habitats, but were less commonly
taken in the Big Blue and Little Blue rivers than in other streams.

Ictalurus natalis ( LeSueur ) , yellow bullhead. Stations 7-G, 9-G,
10-G, 11-G, 17-G, 18-G, 19-G, 34-M, 35-M, 36-M, 37-M, 40-M, 47-M,
48-M, 53-S, and 55-M.

The yellow bullhead inhabited the muddy-bottomed streams and
the upland, gravelly creeks, usually occurring in the headwaters.
I obtained only one I. natalis in the sandy Big Blue River.

Ictalurus punctatus (Rafinesque), channel catfish: Cragin (1885:
107); Meek (1895:135); Evermann and Cox (1896:386); and Can-
field and Wiebe (1931:6-7, 10) as "channel catfish." Stations 1-S,
2-S, 3-S, 4-S, 5-G, 6-S, 7-G, 8-S, 9-G, 11-G, 14-S, 15-S, 16-G, 18-G,
25-S, 27-G, 30-M, 35-M, 38-S, 39-S, 41-S, 42-S, 43-S, 44-S, 46-M,
50-S, 51-M, 52-S, 53-S, and 56-S.

Channel catfish were most common in the larger, sandy streams,
but occurred in other kinds of streams. The ages and calculated
total lengths at the last annulus for 40 channel catfish were: I, no
fish; II, 7.3 (16); III, 10.6 (5); IV, 12.3 (5); V, 13.3 (6); VI, 15.5
(4); VII, 18.0 (3); and VIII, 21.9 (1). These lengths are slightly
lower than averages reported by Finnell and Jenkins (1954:5) in
Oklahoma impoundments.

The length-frequency distribution of 438 channel catfish, col-
lected by rotenone on August 5 and 7, 1958, indicated that two
age-groups were represented. Without examination of spines, I
assigned 265 fish to age-group O (1.3 to 2.9 inches, average 2.5)
and 173 fish to age-group I (3.1 to 5.8 inches, average 4.5). The
average total length of age group I (4.5 inches) is only slightly
higher than the total length at the first annulus reported as average
for Oklahoma (4.0 inches, Finnell and Jenkins, loc. cit.). It seems
unlikely that my yearling fish taken in August, 1958, would have
reached the length at the second anntilus recorded in my study of
spines ( 7.3 inches ) by the end of the 1958 growing season.

From 1952 to 1956, severe drought was prevalent in Kansas,
13robably causing streams to flow less than at any previously re-
corded time (Minckley and Cross, in press). This drought must
have resulted in reduced populations of fishes in the streams. The
channel catfish hatched in 1956 were therefore subjected to low

428 University of Kansas Publs., Mus. Nat. Hist.

competition for food and space when normal flow was resumed in
1957, and grew rapidly, reaching an average total length of 7.3
inches at the second annulus, while channel catfish that were mem-
bers of the large 1957 and 1958 hatches sufiFered more competition
and grew more slowly.

Noturus flavus Rafinesque, stonecat: Jennings (1942:365). Sta-
tions 3-S, 4-S, 6-S, 16-G, 25-S, 28-G, 38-S, 41-S, 42-S, 43-S, 52-S, 53-S,
and 56-S.

Noturus flavus frequented riffles and swift currents along sand-
bars in the Big Blue and Little Blue rivers. Cross (1954:311) re-
ported that "the shale-strewn riffles of the South Fork [of the
Cottonwood River, Kansas] provide ideal habitat for the stonecat."
In my study-area, this species was found not only on rubble-
bottomed riffles, but occurred along both stationary and shifting
sandbars where no cover was apparent.

Pylodictis olivaris (Rafinesque), flathead catfish: Canfield and
Wiebe (1931:7) as "yellow catfish." Stations 3-S, 4-S, 6-S, 8-S,
15-S, 25-S, 38-S, 41-S, 43-S, 44-S, 53-S, and 56-S.

Flathead catfish were found only in the larger rivers. The species
was taken rarely by seine, but was readily obtained by electric
shocker. Data on the age and growth and food-habits of this
species are to be the subject of another paper.

Angtiilla bostoniensis (LeSueur), American eel: Jennings (1942:
365).

American eels are now rare in Kansas, and none was taken in my
survey. The specimen reported by Jennings (he. cit.) is at Kansas
State College (KSC 2916), and was taken by I. D. Graham from
the Big Blue River, Riley County, 1885.

Fundulus kansae Carman, plains killifish. Station 42-S.

The plains killifish was collected by me only at Station 42-S.
Specimens were collected from my Station 4-S by the Kansas State
College class in fisheries management in 1954 (KSC 4985). My
specimens were 11 to 13 mm. in total length.

Roccus chrysops (Rafinesque), white bass. Station 3-S.

That the white bass is indigenous to Kansas is evidenced by
records of Craham (1885:77) and Cragin (1885:111); however,
since that time, and prior to the introduction of this species into
reservoirs in the State, R. chnjsops has rarely been recorded in
Kansas. I collected young white bass at Station 3-S in both 1957
and 1958, and I collected them also in an oxbow of the Kansas River
four miles west of Manhattan, Riley County, Kansas, in the mouth

Fishes of Big Blue Basin 429

of McDowell's Creek, Riley County, and in Deep Creek, Wa-
baunsee County, and I saw other specimens from an oxbow of the
Kansas River on the Fort Riley Military Reservation, Riley County,
Kansas. The apparent increase in abundance of white bass in the
Kansas River Basin must be attributable to introductions in reser-
voirs, with subsequent escape and establishment in the streams.

Micropterus salmoides salmoides (Lacepede), largemouth bass.
Stations 6-S, 11-G, 43-S, and 45-M.

Four largemouth bass were taken. This species has been widely
stocked in farm-ponds and other impoundments in Kansas.

Lepomis cyanellus Rafinesque, green sunfish: Breukelman (1940:
382); and Canfield and Wiebe (1931:5, 7-8, 10) as "green sunfish."
All stations excepting 1-S, 2-S, 4-S, 8-S, 9-G, 15-S, 22-G, 25-S,
30-M, 32-G, 34-M, 38-S, 39-S, 41-S, 42-S, 43-S, 44-S, 45-M, 46-M,
47-M, 50-S, and 52-S.

Green sunfish occurred primarily in the muddy streams. The ages
and total lengths at the last annulus for 25 specimens are as follows :
I, 1.1 (9); II, 2.2 (4); III, 3.1 (7); IV, 5.4 (4); and V, 6.0 (1).
Male green sunfish were seen on nests on June 29, July 1, and
July 9, 1958.

Lepomis humilis (Girard), orangespotted sunfish: Meek (1895:
137); Evermann and Cox (1896:418); Canfield and Wiebe (1931:6)
as "orange spots"; and Breukelman (1940:382). All stations except-
ing 1-S, 9-G, 13-G, 15-G, 17-G, 21-G, 26-G, 34-M, 36-M, 38-M, 43-M,
44-S, 47-M, 50-S, and 52-S.

Lepomis humilis was most common over sand-silt bottoms. Only
two age-groups were found; their calculated total lengths were
I, 1.7 (15); and II, 2.4 (10). Orangespotted sunfish were seen nest-
ing on the same dates as Lepomis cyanellus.

Lepomis macrochirus Rafinesque, bluegill. Stations 7-G, 13-G,
16-G, 24-G, and 59-G.

This species has been widely stocked in Kansas. Only young-of-
the-year and sub-adults were taken, and these were rare.

Pomoxis annularis Rafinesque, white crappie: Canfield and
Wiebe (1931:5-8, 10) as "white crappie." Stations 3-S, 6-S, 8-S,
12-G, 42-S, and 53-S.

White crappie were rare, except in a borrow-pit at Station 6-S.
Ages and calculated total lengths at the last annulus for 50 speci-
mens from 6-S are as follows: I, 3.6 (22); II, 5.0 (14); III, 7.1 (5);
IV, 8.3 (7); and V, 10.7 (2).

Pomoxis nigromaculatus (LeSueur), black crappie. Station 6-S.

430 Untvtersity of Kansas Publs., Mus. Nat. Hist.

One black crappie (KU 4174) was taken. Canfield and Wiebe
(1931:10) noted: "The Bkick Crappie has been planted here [Big
Blue River Basin in Nebraska] by the State, but, apparently, is not
propagating itself."

Stizostedion canadense (Smith), sauger. Station 56-S.

Mr. Larry Stallbaumer, of Marysville, Kansas, obtained a sauger
(KU 4179) while angling on May 25, 1958.

Stizostedion vitrcum (Mitchill), walleye.

Though I failed to obtain the walleye in my survey. Dr. Ray-
mond E. Johnson (personal communication) reported that the
species occurred in the Nebraskan portion of the Big Blue River
in recent years. Canfield and Wiebe (1931:6, 10) reported that
'yellow pike are taken at Crete [Nebraska]," but may have referred
to either the walleye or the sauger.

Perca fiavescens (Mitchill), yellow perch: Canfield and Wiebe
(1931:5-6, 10) as "ring perch" and "yellow perch."

This fish was not taken in my survey. Canfield and Wiebe ( loc.
cit.) reported that the yellow perch "had been planted by the
State [Nebraska]."

Etheostoma nigrum nigrum Rafinesque, johnny darter: Jennings
(1942:365) as Boleosoma nigrum nigrum (Rafinesque). Stations
10-G, 11-G, 12-G, 13-G, 16-G, 29-G, 40-M, 53-S, and 54-G.

The larger pools of gravelly streams were preferred by johnny
darters, but one specimen was taken from the main stream of the
Big Blue River, and the species was abundant in one stream over
hard, sand-silt bottom.

Etheostoma spectabile pulchellum (Girard), orangethroat
darter: Jennings (1942:365) as Poeciliclithys spectahilis pulchclhis
(Girard). Stations 5-G, 7-G, 10-G, 11-G, 12-0, 13-G, 16-G, 17-G,
18-G, 21-G, 23-G, 27-G, 28-G, 29-G, 33-M, 40-M, 49-M, 53-S,
54-G, and 59-G.

The orangethroat darter was less restricted in habitat than the
johnny darter, occurring in all stream-types, but most often in the
rifHes of gravelly streams. Most specimens from muddy or sandy
streams were small.

Aplodinotus grunniens Rafinesque, freshwater drum. Stations
3-S, 4-S, 6-S, 7-G, 8-S, 15-S, 38-S, 39-S, 53-S, and 56-S.

The ages and calculated total lengths at the last annulus for 42
freshwater drum from the Big Blue River were: I, 3.0 (10); II,
5.7 (6); III, 9.4 (7); IV, 12.1 (13); V, 14.0 (3); VI, 15.1 (2); and
VII, 16.3 (1).

Fishes of Big Blue Basin 431

HYBRID COMBINATIONS

I obtained two hybrid fishes in my study-area. One specimen of
Notropis corntitus frontalis X Chrosomus erythrogaster was taken
at Station 29-G. This combination was recorded by Trautman
(1957:114) in Ohio. The other hybrid was Lepomis cyanellus X
Lepomis humilis, captured at Station 24-G. This combination was
first recorded by Hubbs and Ortenburger (1929:42).

Hubbs and Bailey (1952:144) recorded another hybrid combina-
tion from my area of study: Campostoma anomalum plumheiim X
Chrosomus erythrogasier, UMMZ 103132, from a "spring-fed creek
on 'Doc' Wagner's farm, Riley County, Kansas; September 21,
1927; L. O. Nolf [collector]."

RELATIVE ABUNDANCE AND DISCUSSION OF SPECIES

The reladve abundance of different species was estimated by
combining counts of individual fishes taken in 290 seine-hauls, 26
hours and 15 minutes of shocking, and seven samples obtained with
rotenone. At some stations all seine-hauls were counted. At other
stations the seine-hauls in which complete counts were recorded
had been selected randomly in advance; that is to say, prior to
collecting at each station. I selected those hauls to be counted
from a table of random numbers (Snedecor, 1956:10-13). I did
not use the frequency-of-occurrence method as proposed by Starrett
(1950:114), in which the species taken and not the total number
of individuals are recorded for all seine-hauls. However, the fre-
quency of occurrence of each species is indicated by the number
of stations at which it was found, and those stations are listed in
the previous accounts. Table 3 shows the percentage of the total
number of fish that each species comprised in three kinds of streams :
sandy (Big Blue and Little Blue rivers), muddy, and gravelly
streams.

The habitat preferences of some species affect their abundance
in different stream-types. Notropis lufrensis and P. mirahilis seemed
almost ubiquitous. Notropis deliciosus also occurred in all kinds
of streams (rarely in muddy streams); however, this species was
represented by the sand-loving N. d. missuriensis in the Big Blue
and Little Blue rivers, and N. d. deliciosus in the clear, gravelly,
upland creeks (Nelson, personal communication). Because of its
widespread occurrence, and for purposes of later discussion, I refer
to this minnow also as an ubiquitous species in the Big Blue River
Basin.

Carpiodes carpio, Cyprinus carpio, I. punctatus, I. melas, and L.

432

University of Kansas Publs., Mus. Nat. Hist.

Table 3. Relative AsuhTOANCE of Fishes in Per Cent of the Total
Number T.vken, Big Blue Rh'er Basin, Kansas. Trace (Tr. ) Is Used
FOR Values Less Than .05 Per Cent, and Dashes Signify that the
Species Did Not Occur est the Counted Collections Although it May
Have Occurred in Uncounted Collections from the Same Stream-type.
Three Species, C. auratus, N. buchanani, and S. canadense. Were Not
Taken in Counted Collections.

Species

N. lutrensis

/. pundatus

Carpiodes carpio. .

N. deliciosus

/. melas

Cyprinus carpio. .

P. olivaris

L. humilis

/. bubalus

P. mirabilis

H. nuchalis

P. promelas ,

H. aestivalis

A . grunniens . . . .

L. osseus

C. anomalum. . . .

C. commersonnii . .

D. cepedianum. . .
N. percobromus. . .

P. annularis

N. flavus

S. atromaculalus . .

M. aureolum

/. cyprinella

P. notatus

/. niger

H. alosoides

E. spedabile

R. chrysops

L. cyanellus

//. storeriana

L. plalostomus. . . ,

M. salmoides

P. nigromaculalus .

I. natalis

N. ■umbratilis . . . . .

C. forbesi ,

S. plalorynchus . . .

F. kansae

E. nigrum

A'^. rubellus

A^. topeka

N. comutus

C. erylhrogasler . . .
L. macrochirus ...

Sandy streams

Big

Blue
River

43.5
14.0
11.9
8.2
2.5
2.3
1.8
1.7
1.4
1.3
1.2
0.8
0.7
0.5
0.5
0.4
0.4
0.4
0.3
0.3
0.2
0.2
0.1
0.1
0.1
0.1
0.1
0.1
0.1
0.1
Tr.
Tr.
Tr.
Tr.
Tr.
Tr.
Tr.
Tr.

Tr.

Little
Blue
River

55.9
7.0
2.0

28.2

1.9
0.8

0.1
0.7

1.0
0.2

1.0
0.2

Tr.

Tr.
0.4
0.1
0.2

0.1

Tr.

Muddy
streams

27.6
1.2
5.0
3.1
1.3
2.7

9.0

0.3

28.7

2.7
0.1

12.2
0.1

1.4
3.5

1.0

0.1

Gravelly
streams

56.0
4.2
0.5

11.1
0.5
0.2

5.1
Tr.
1.3
Tr.
4.0

0.2

4.6
0.7

2.2

1.6
Tr.

Tr.

0.2

Tr.
1.0
1.0
1.0
1.0

Fishes of Big Blue Basin 433

humilis were widespread, but each was absent or rare in one of
the kinds of streams (Table 3). Carpiodes carpio, Cyprinus carpio,
and J. punctatus occurred most frequently in the sandy streams,
whereas L. humilis was most common in muddy streams. The high
per cent of 1. melas in collections from the Big Blue River is a direct
result of one large population that was taken with rotenone in a
borrow-pit at Station 6-S. In my opinion, this species actually
was most abundant in the muddy streams.

Some fish were almost restricted to the sandy streams, apparently
because of preference for larger waters, or sandy stream-bottoms:
P. olivaris, I. huhalus, H. ntichalis, H. aestivalis, A. grunniens, L.
osseus, D. cepedianum, N. percobromus, P. annularis, N. flavus, M.
aureolum, I. niger, H. alosiodes, and R. chrysops. Other species
that were taken only in the larger rivers, and that are sometimes
associated with streams even larger (or more sandy) than the Big
Blue River are H. storeriana, L. platostomus, M. salmoides, P. ni-
gromaculatus, C. forbesi, S. platorynchus, F. kansae, N. buchanani,
S. canadense, and C. auratus. Ictiobus cyprinella also occurred
more frequently in the larger streams.

The muddy-bottomed streams supported populations composed
primarily of P. promelas, N. lutrensis, and S. atromaculatiis. No
species was restricted to this habitat, but the following were
characteristic there: P. promelas, S. atromaculatus, L. humilis,
L. cyanellus, and I. natalis. Carpiodes carpio, Cyprinus carpio,
C. anomalum, E. spectabile, and E. nigrum were locally common in
muddy streams, but the first two were most frequent in larger,
sandy streams, and the last three in gravelly streams.

In gravel-bottomed, upland streams, N. cornutus, N. rubellus,
N. topeka, and C erythrogaster characteristically occurred; with
the exception of N. rubellus (only one specimen taken), all were
common at some stations. Other species in gravelly creeks were
N. lutrensis, C. anomalum, C. commersonnii, P. notatus, L. macro-
chirus, E. spectabile, and E. nigrum. Although the one specimen of
JV. umbratilis taken in this survey was from the Big Blue River, this
species is more characteristic of the clearer creeks in Kansas.

In order to illustrate the composition of the fauna in some specific
streams in the Big Blue River Basin, I segregated the fishes into
ecological groups, as in the above discussion: ubiquitous types;
species of larger, sandy streams; fishes of muddy streams; and
fishes of clear, gravelly creeks.

The total number of species taken in each of the streams was

3—7080

434

University of Kansas Publs., Mus. Nat. Hist.

divided into the number of species from that stream that were in
each of these units, to give a percentage. The resultant data are
presented graphically in Figure 3,

Fig. 3. Composition of the fauna of the entire Big Blue River Basin, and of
seven streams or stream systems in that basin. "Mill Creek, Wash. Co." refers
to all streams in the Mill Creek System, Washington and Republic counties.
"Bl. Vermillion R. System" includes all streams in that watershed excepting
Clear Creek and one of its tributaries (Stations 31-G and 32-G).

Figure 3 gives a generalized picture of the faunal composition in
difiFerent kinds of streams. However, the fauna of a small tributary
becomes more distinct from the fauna of the larger stream into
which the small stream flows as one moves toward the headwaters
(Metcalf, 1957:92, 95-100). Figure 4 illustrates this in Carnahan

100

c
v
o 50

a>
a.

Fishes of Gravelly Streams

I I I I u

Big Blue
R i y e r

Fishes of Sandy Streams

Stat ion
11 G

Station
13 G

1 mile 1 mile

1 mile

4^ mile

M miles

Fig. 4. Composition of the fauna of the Big Blue River, and of five collecting-
sites pn Carnahan Creek, Pottawatomie County. Lowermost sites are at the

left of the figure.

Fishes of Big Blue Basin 435

Creek. Station 11-G included four sampling-sites, which were ap-
proximately one, two, three, and four miles upstream from the
mouth of Carnahan Creek. Station 13-G (one collection) was
about four miles upstream from the closest sampling-site of Sta-
tion 11-G. Applying the same methods as for Figure 3, my findings
show a gradual decline in the per cent of the fauna represented
by the 'large-river-fishes," and an increase in the segment classified
as "upland-fishes," from downstream to upstream.

CREEL CENSUS

Fifty-three fishermen were interviewed in the 1957 creel census
period, and 152 in 1958. Only those fishermen using pole and line
were interviewed. In the area censused, much additional fishing is
done with set-lines, that are checked periodically by the owners.

In the 1958 census, 22 checks along approximately 80 miles of
river were made, and seven of these trips were made without seeing
one fisherman. The average fishing pressure for the entire area
was estimated at one fisherman per 7.9 miles of stream, or one
fisherman per 15.7 miles of shoreline.

Seven species of fish were identified from fishermen's creels in
1957 and 1958. These, in order of abundance were: channel cat-
fish; carp; freshwater drum; flathead catfish; shovelnose sturgeon;
smallmouth buffalo; and river carpsucker. Shovelnose sturgeon
occurred in fishermen's creels only in April, 1957, and freshwater
drum occurred more frequently in the spring-census of 1957 than
in the summer of 1958.

Sixty-two of the fishermen interviewed in 1958 were fishing for
"anything they could catch," 68 were fishing specifically for catfish,
and 22 sought species other than catfish. The order of preference
was as follows: channel catfish, 21.1 per cent; flathead catfish, 15.1
per cent; unspecified catfish, 12.5 per cent; carp, 9.2 per cent; fresh-
water drum, 1.3 per cent; and unspecified, 40.8 per cent. The kinds
of fish desired by those fishermen checked in 1957 were not
ascertained.

Of all fishermen checked in 1957 and 1958, 165 were men, 17 were
women, and 24 were children. Ninety-three per cent were fishing
from the bank, five per cent were fishing from bridges, and two
per cent were wading. All but two per cent of those checked were
fishing "tightline"; the remainder fished with a cork.

The ten baits most commonly used, in order of frequency, were
worms, doughballs, minnows, Hver, beef-spleen, chicken-entrails,
coagulated blood, crayfish, shrimp, and com.

436

University of Kansas Publs., Mus. Nat. Hist.

For purposes of later comparison the data on angler success
(Table 4) have been divided according to areas: Area I, below
Tuttle Creek Dam; Area II, in the Tuttle Creek Reservoir area; and
Area III, above the reservoir. Areas I and III received the most
fishing pressure, especially Station 4-S (in Area I), and Station
56-S (in Area III).

In Area I, the success ranged from 0.91 fish per fisherman-day
in 1957 to 0.26 fish per fisherman-day in 1958. The 1957 census
was made in April and May, when fishing in warm-water streams
is considered better than in July (Harrison, 1956:203). The 1958
census was from late June through July, and stream-flow in this
period was continuously above normal. Therefore, fewer people
fished the river, and catches were irregular. Catches in 1958
ranged from 0.26 fish per fisherman-day in Area I to 0.44 fish per
fisherman-day in Area III. In 1951, in the Republican River of
Kansas and Nebraska, the average fisherman-day yielded 0.36 fish.
0.09 fish per man-hour, and 0.06 fish per pole-hour (U. S. Fish and
Wildlife Service, 1952:13-14). The average fisherman-day in the
Republican River study was 3.0 hours, whereas the average on the
Big Blue River was 2.2 hours for all areas in 1958 (Table 4).

In the Big Blue River 47.7 per cent of all fishermen were suc-
cessful in Area I in 1957, while only 13.1 per cent were successful
in the same area in 1958 (Table 5). In the Repubhcan River, 24
per cent of the fishing parties were successful (1.64 persons per
party) (U. S. Fish and Wildhfe Service, loc. cit.). The average

Table 4. Angling Success in the Big Blue River, Kansas, 1957 and 1958.

Area, Year,

AND Number

OF Fishermen

Average

length of

fisherman-day

Number

fish per

fisherman-day

Number

fish per

man-hour

Number

fish per

pole-hour*

Area I, 1957
53 fishermen

2.7 hours

0.91

0.33

0.23

Area I, 1958
84 fishermen

2.5 hours

0.26

0.10

0.07

Area II, 1958
27 fishermen

1 . 7 hours

0.37

0.22

0.14

Area III, 1958
41 fishermen

2.4 hours

0.44

0.16

0.11

All areas, 1958
152 fishermen

2.2 hovirs

0.33

0.14

0.09

* Fishermen used an average of 1.44 poles.

Fishes of Big Blue Basin

437

distance that each fisherman had traveled to fish in the Big Blue
River was 15.7 miles. Seventy-nine per cent of the persons con-
tacted lived vdthin 25 miles of the spots where they fished. In
the study on the RepubHcan River, 77 per cent of the parties inter-
viewed came less than 25 miles to fish.

Table 5. Per Cent of Total Fishermen Successful, and Distances
Travelled to Fish, Big Blue River Basin, Kansas, 1957 and 1958. All
Distances Were Measured in Airline Miles.

1957

Area

I

1958

Area

I

1958

Area

II

1958

Area

III

1958

All

areas

Per cent of fishermen successful

Distances traveled to fish,
(averages in parentheses) .

47.1

0-121
(15.6)

13.1

1-197
(20.5)

18.5

0-124
(13.5)

19.5

0-60
(7.4)

15.8

0-197
(15.7)

RECOMMENDATIONS

My primary recommendation is for continued study of the
Tuttle Creek Reservoir, and the Big Blue River above and below
the reservoir, to trace changes in the fish population that result
from impoundment.

Probably the fishes that inhabit the backwaters, creek-mouths,
and borrow-pits in the Big Blue River Basin ( gars, shad, carpsucker,
buflFalo, carp, sunfishes, and white bass ) will increase in abundance
as soon as Tuttle Creek Reservoir is formed. Also, as in eastern
Oklahoma reservoirs (see Finnell, et ah, 1956:61-73), populations
of channel and flathead catfish should increase. Because of the
presence of brood-stock of the major sport-fishes of Kansas ( channel
and flathead catfish, bullhead, bluegill, crappie, largemouth bass,
and white bass), stocking of these species would be an economic
waste: exception might be made for the white bass. It may be
above Tuttle Creek Dam, but was not found there.

I do recommend immediate introduction of walleye, and possibly
northern pike {Esox lucius Linnaeus), the latter species having
been successfully stocked in Harlan County Reservoir, Nebraska,
in recent years (Mr. Donald D. Poole, personal communication).
These two species probably are native to Kansas, but may have
been extirpated as agricultural development progressed. Reservoirs
may again provide habitats suitable for these species in the State.

If Tuttle Creek Reservoir follows the pattern found in most
Oklahoma reservoirs, large populations of "coarse fish" — ^fishes that

438 UxrvTRSiTY of Kansas Publs., Mus. Nat. Hist.

are, however, commercially desirable — will develop (Finnell, et ah,
loc. cit.). To utilize this resource, and possibly to help control
"coarse fish" populations for the betterment of sport-fishing, some
provision for commercial harvest should be made in the reservoir.

SUMMARY

1. The Big Blue River Basin in northeastern Kansas was studied
between March 30, 1957, and August 9, 1958. The objectives were
to record the species of fish present and their relative abundance
in the stream, and to obtain a measure of angling success prior
to closure of Tuttle Creek Dam.

2. Fifty-nine stations were sampled one or more times, using
seines, hoop and fyke nets, wire traps, experimental gill nets,
rotenone, and an electric fish shocker.

3. Forty-eight species of fish were obtained, and five others have
been recorded in literature or found in museums. One species,
Carpiodes forbesi, is recorded from Kansas for the first time.

4. Notropis lutrensis was the most abundant fish in the Big Blue
River Basin, followed by Notropis deliciosus and Ictalurus
punctatus. The most abundant sport-fishes were I. punctatus, I.
melas, and Pylodictis olivaris, respectively.

5. The spawning behavior of Notropis lutrensis is described.

6. A creel census at major points of access to the Big Blue River,
was taken in 1957 (below Tuttle Creek Dam) and in 1958 (above,
in, and below the dam-site). Fishing pressure averaged one fisher-
man per 15.7 miles of shoreline. The average length of the fisher-
man-day averaged 2.2 hours, with an average of 0.33 fish per fisher-
man-day being caught in 1958. The average number of fish per
man-hour in 1958 was 0.14 and 15.8 per cent of the fishermen were
successful. Distances traveled in order to fish ranged from 0 to
197 miles ( airline ) and averaged 15.7 miles.

7. The primary recommendation is that studies be continued, to
document changes that result from impoundment. Because brood-
stock of the major sport-fishes is already present, stocking is un-
necessary, except for walleye and northern pike. Also, I recom-
mend commercial harvest of non-game food-fishes.

Fishes of Big Blue Basin 439

LITERATURE CITED

Badley, R. M.

1956. A revised list of fishes of Iowa, with keys for identification, In

Iowa Fish and Fishing, by J. R. Harlan and E. B. Speaker. Iowa

State Cons. Comm., Des Moines, pp. 325-377.

, and Cross, F. B.

1954. River sturgeons of the American genus Scaphirhynchus: characters,

distribution, and synonymy. Pap. Michigan Acad. Sci., Arts,

and Letters, 39 ( 1953 ) : 169-208.

Breukelman, J.

1940. A collection of Kansas fish in the State University Museum. Trans.

Kansas Acad. Sci., 43: 377-384.

BUCHHOLZ, M.

1957. Age and growth of river carpsucker in Des Moines River, Iowa.
Proc. Iowa Acad. Sci., 64: 589-600.

Canfield, H. L., and Wiebe, A. H.

1931. A cursory survey of the Blue River System of Nebraska. U. S.
Dept. Comm., Bur. of Fisheries, Econ. Circ. 73: 1-10.
Colby, C, Dillingham, H., Erickson, E., Jenks, C, Jones, J., and
Sinclair, R.
1956. The Kansas Basin, Pilot Study of a Watershed. Univ. of Kansas
Press, Lawrence, ix + 103 pp.
Cragin, F. W.

1885. Preliminary list of Kansas fishes. Bull. Washburn Lab. of Nat.
Hist., 1 (3):105-111.
Cross, F. B.

1950. Effects of sewage and of a headwaters impoundment on the
fishes of Stillwater Creek in Pajme County, Oklahoma. Amer.
Midi. Nat., 43 (1):128-145.
1954. Fishes of Cedar Creek and the South Fork of the Cottonwood
River, Chase County, Kansas. Trans. Kansas Acad. Sci., 57 (3):
303-314.
Eddy, S., and Surber, T.

1947. Northern Fishes, with Special Reference to the Upper Mississippi
Valley. Univ. of Minnesota Press, Minneapolis, xii + 276 pp.

Evermann, B. W., and Cox, U. O.

1896. Report upon the fishes of the Missovuri River Basin. Appendix 5.
Rept. U. S. Commissioner of Fish and Fisheries for 1894. pp.
325-429.
Finnell, J. C, and Jenkins, R. M.

1954. Growth of channel catfish in Oklahoma waters: 1954 revision.
Oklahoma Fish Res. Lab. Rept. 41: ii + 1-37.

_^ , Jenkins, R. M. and Hall, G. E.

1956. The fishery resources of the Little River System, McCurtain County,
Oklahoma. Oklahoma Fish. Res. Lab. Rept. 55: ii -f 1-82.
Flora, S. D.

1948. Climate of Kansas. Rept. Kansas State Board of Agri., 67 (285):
xii + 1-320.

440 Untv'ersity of Kansas Publs., Mus. Nat. Hist.

Forbes, S. A., and Richardson, R. E.

1920. The Fishes of Illinois. Nat. Hist. Survey of Illinois. Illinois Print-
ing Co., Danville, cxxxi + 357 pp.
Fbye, J. C, and Leonard, A. B.

1952. Pleistocene geology of Kansas, Kansas Geol. Survey, Bull. 99: 1-
230.
Gilbert, C. H.

1886. Third series of notes on the fishes of Kansas. Bull. Washburn Lab.
of Nat. Hist., 1 (6): 207-211.
Graham, I. D.

1885. Preliminary list of Kansas fishes. Trans. Kansas Acad. Sci., 9: 69-
78.
Harrison, H. M.

1956. Angling for channel catfish. In lovv'a Fish and Fishing, by J. R.
Harlan and E. B. Speaker. Iowa State Cons, Comm., Des Moines.
Pp, 202-212.

HtTBBS, C, L.

1945. Corrected distributional records for Minnesota fishes. Copeia, 1945
(l):13-22.

, and Ortenburger, A, I.

1929. Further notes on the fishes of Oklahoma with descriptions of new
species of Cyprinidae, Pub. Univ. Oklahoma Biol. Survey, 1 (2):
17-43.
.., and Lagler, K. F.

1947. Fishes of the Great Lakes Region. Cranbrook Inst, of Sci., Bull.
26: xi + 1-186.
., and Bailey, R. M.

1952. Identification of Oxygeneum pulverulentum Forbes, from Illinois,
as a hybrid cyprinid fish. Pap, Michigan Acad. Sci., Arts, and
Letters, 37 (1951): 143-152,
Jennings, D.

1942. Kansas fish in the Kansas State College Museum at Manhattan,
Trans. Kansas Acad. Sci., 45: 363-366.
Kansas Water Resources Fact-finding and Research Committee.

1955. Water in Kansas. A Report to the Kansas State Legislatiire . , .
Univ. of Kansas, 1-216 pp,
KiNCER, J. B.

1941, Climate and weather data for the United States. In Climate and
Man, Yearbook of Agri. for 1941, House Doc. 27. pp. 685-699,
Lacler, K. F,

1952. Freshwater Fishery Biology. Wm. C. Brown Co., Dubuque, Iowa.
X + 360 pp,
LuGN, A. L.

1935. The Pleistocene geology of Nebraska. Nebraska Geol. Survey,
Bull. 10, 2nd series: 1-223.
Marzolf, R. C.

1955. Use of pectoral spines and vertebrae for determining age and rate
of growth of the channel catfish. Jour. Wildl. Mgmt., 19 (2): 243-
249.

Fishes of Big Blue Basin 441

Meek, S. E.

1895. Notes on the fishes of western Iowa and eastern Nebraska. Bull.
U. S. Fish Comm., 14 ( 1894) : 133-138.
Metcalf, a. L.

1957. Fishes of Chautauqua, Cowley and Elk counties, Kansas. Univ.
Kansas Publ., Mus. Nat. Hist., 11:345-400.
MiNCKLEY, W. L., and Cross, F. B.

In press. Habitat, distribution, and abundance of Notropis topeka (Gil-
bert) in Kansas. Amer. Midi. Nat.

MONFORT, E.

1956. A layman looks at water. Trans. Kansas Acad. Sci., 59 ( 1 ) : 118-123.
Moore, G. A.

1957. Fishes. In Vertebrates of the United States, by W. Blair, A. Blair,
P. Brodkorb, F. Cagle, and G. Moore. McGraw-Hill Book Co.,
New York, N. Y. pp. 31-210.

Moore, R. C, and Landes, K. K.

1937. Geologic map of Kansas. Scale 1:500,000. Kansas Geol. Survey.
Nebraska State Planning Board.

1936. Water resources of Nebraska. Mimeo. by Nebraska State Planning
Board. Lincoln, xxviii -|- 695 pp.
Pfedffer, R. a.

1955. Studies on the life history of the rosyface shiner, Notropis rubellus.
Copeia, 1955 (2):95-104.

Raney, E. C.

1947. Subspecies and breeding behavior of the cyprinid fish Notropis
procne ( Cope ) . Copeia, 1947 ( 2 ) : 103-109.

SCHOEWE, W. H.

1953. The geography of Kansas, Part III — concluded, hydrogeography.
Trans. Kansas Acad. Sci., 56 (2): 131-190.
Snedecor, G. W.

1956. Statistical Methods. Iowa State College Press, Ames, xiii + 534 pp.
Starrett, W. C.

1950. Distribution of the fishes of Boone County, Iowa, with special ref-
erence to the minnows and darters. Amer. Midi. Nat., 43 (1):
112-127.
Trautman, M. B.

1957. The Fishes of Ohio. Waverly Press, Inc., Baltimore, Md. xvii -f
683 pp.

U. S. Fish and WiLDLrFE Service.

1952. A one-year creel census and evaluation of the Republican River,
Nebraska and Kansas, 1951. Mimeo. by the Staff, Missouri River
Basin Studies, Billings, Mont. 29 pp., Appendix.

1953. A preliminary report on fish and wildlife resources in relation to
the water development plan for the Tuttle Creek Dam and Reser-
voir, Big Blue River, Missouri River Basin, Kansas. Mimeo. by the
Staff, Missouri River Basin Studies, Billings, Mont. 25 pp.

442 Untv'ersity of Kansas Publs., Mus. Nat. Hist.

Van Orman, C. R.

1956. Surface water — its control and retention for use. Trans. Kansas
Acad. Sci., 59 (1):105-110.
Walters, K. L.

1954. Geology and ground-water resources of Marshall County, Kansas.
Kansas Geol. Survey, Bull. 106:1-116.

Transmitted December 19, 1958.

D

27-7080

(Continued from inside of front cover)

Vol. 8. 1. Life history and ecology of the five-lined skink, Eumeces fasciatus. By Henry
S. Fitch. Pp. 1-156, 26 figs, in text. September 1, 1954.

2. Myology and serology of the Avian Family Fringillidae, a taxonomic study.
By William B. Stallcup. Pp. 157-211, 23 figures in text, 4 tables. November
15, 1954. , „ . s ^ „

3. An ecological study of the collared lizard (Crotaphytus collaris). By Henry
S. Fitch. Pp. 213-274, 10 figiures in text. February 10, 1956.

4. A field study of the Kansas ant-eating frog, Gastrophryne olivacea. By Henry
S. Fitch. Pp. 275-306, 9 figures in text. February 10, 1956.

5. Check-list of the birds of Kansas. By Harrison B. Tordoff. Pp. 307-359, 1
figure in text. March 10, 1956.

6. A population study of the prairie vole (Microtus ochrogaster) in northeastern
Kansas. By Edwin P. Martin. Pp. 361-416, 19 figures in text. April 2,
1956.

7. Temperature responses in free-living amphibians and reptiles of northeastern
Kansas. By Henry S. Fitch. Pp. 417-476, 10 figures in text, 6 tables. June
1, 1956.

8. Food of the crow, Corvus brachyrhynchos Brehm, in south-central Kansas. By
Dwight Piatt. Pp. 477-498, 4 tables. June 8, 1956.

9. Ecological observations on the woodrat Neotoma floridana. By Henry S.
Fitch and Dennis G. Rainey. Pp. 499-533, 3 figures in text. June 12, 1956.

10. Eastern woodrat, Neotoma floridana; Life history and ecology. By Dennis
G. Rainey. Pp. 535-646, 12 plates, 13 figures in text. August 15, 1956.
Index. Pp. 647-675.
Vol. 9. 1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18
figtures in text. December 10, 1955.

2. Additional records and extensions of ranges of mammals from Utah. By
Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80.
December 10, 1955.

3. A new long-eared myotis (Mvotis evotis) from northeastern Mexico. By Rol-
lin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.

4. Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming.
By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956.

5. The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6
figiu-es in text. May 19, 1956.

6. Additional remains of the multituberculate genus Eucosmodon. By Robert
W. Wilson. Pp. 117-123. 10 Gguies in text. May 19, 1956.

7. Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures
in text. June 15, 1956.

8. Comments on the taxonomic status of Apodemus peninsulae, with description
of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 337-346,
1 figure in text, 1 table. August 15, 1956.

9. Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp.
347-351. August 15, 1956.

10. A new bat (Genus Leptonycteris ) from Coahuila. By Howard J. Stains.
Pp. 353-356. January 21, 1957.

11. A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico.
By Robert J. RusseU. Pp. 357-361. January 21, 1957.

12. Geographic variation in the pocket gopher, Thomomys bottae, in Colorado.
By Phillip M. Yoimgman. Pp. 363-387, 7 figures in text. February 21, 1958.

13. New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones,
Jr. Pp. 385-388. May 12, 1958.

14. Pleistocene bats from San Josecito Cave, Nuevo Leon, Mexico. By J. Knox
Jones. Jr. Pp. 389-396. December 19, 1958.

15 New subspecies of the rodent Baiomys from Central America. By Robert
L. Packard. Pp. 397-404. December 19, 1958.
More numbers will appear in voliune 9.
Vol 10. 1. Studies of birds killed in nocturnal migration. By Harrison B. Tordoff and
Robert M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 1956.

2. Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.

3. The forest habitat of the University of Kansas Natural History Reservation.
By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures
in text, 4 tables. December 31, 1956.

4. Aspects of reproduction and development in the prairie vole (Microtus ochro-
gaster). By Henry S. Fitch. Pp. 129-161, 8 figiures in text, 4 tables. Decem-
ber 19, 1957.

5 Birds found on the Arctic slope of northern Alaska. By James W. Bee.

Pp. 163-211, pis. 9-10, 1 figure in text, March 12, 1958.
6. The wood rats of Colorado: distribution and ecology. By Robert B. Finley,

Jr. Pp. 213-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958.

More numbers wiU appear in volume 10.

(Continued on outside of back cover)

(Continued from inside of back cover)

Vol. 11. 1. The systematic status of the colubrid snake, Leptodeira discolor Giinther.
By William E. Duelhnan. Pp. 1-9, 4 figs. July 14, 1958.

2. Natural historv of the six-lined racerunncr, Cnemidophorus sexlineatus. By
Henry S. Fitch. Pp. 11-62, 9 figs., 9 tables. September 19, 1958.

3. Home ranges, territories, and seasonal movements of vertebrates of the
Natural History Reservation. By Henry S. Fitch. Pp. 63-326, 6 plates, 24
figures in text, 3 tables. December 12, 1958.

4. A new snake of the genus Geophis from Chihuahua, Mexico. By John M,

Legler. Pp. 327-334. January 28, 1959.

5. A new tortoise, genus Gopherus, from north-central Mexico. By John M.
Legler. Pp. 335-343, April 24, 1959.

6. Fishes of Chautauqua, Cowley and Elk counties, Kansas. By Artie L.
Metcalf. Pp. 345-4^00, 2 plates, 2 figures in text, 10 tables. May 6, 1959.

7. Fishes of the Big Blue River Basin, Kansas. By W. L. Minckley. Pp. 401-
442, 2 plates, 4 figures in text, 5 tables. May 8, 1959.

More numbers will appear in volume 11.

4

4

f n~ L.

SEP "3 1959

mmmi

University of Kansas Publications "
Museum of Natural History

Volume 11, No. 8, pp. 443-516
August 1, 1959

Birds From Coahuila, Mexico

BY

EMIL K. URBAN

University of Kansas
Lavs^rence
^ 1959

UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY

Institutional libraries interested in publications exchange may obtain this
series by addressing the Exchange Librarian, University of Kansas Library,
Lawrence, Kansas. Copies for individuals, persons working in a particular
field of study, may be obtained by addressing instead the Museum of Natural
History, University of Kansas, Lawrence, Kansas. There is no provision for
sale of this series by the University Library which meets institutional requests,
or by the Museum of Natural History which meets the requests of individuals.
However, when individuals request copies from the Museum, 25 cents should
be included, for each separate number that is 100 pages or more in length, for
the purpose of defraying the costs of wrapping and mailing.

* An asterisk designates those numbers of which the Museum's supply (not the Library's
supply) is exhausted. Numbers published to date, in this series, are as follows:
Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950.

*Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140
figures in text. April 9, 1948.

Vol. 3. *1. The avifaima of Micronesia, its origin, evolution, and distribution. By Rol-
lin H. Baker. Pp. 1-359, 16 figvues in text. June 12, 1951.
*2. A quantitative study of the nocturnal migration of birds. By George H.
Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951.

3. Phvlogenv of the waxwings and allied birds. By M. Dale Arvey. Pp. 473-
530, 49 figures in text, 13 tables. October 10, 1951.

4. Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and
Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10,
1951.

Index. Pp. 651-681.

*Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31
figures in text. December 27, 1951.

Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953.
*Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D.
Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952.

Vol. 7. *1. Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, 73 figines in
text, 37 tables. August 25, 1952.

2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry
S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 figures in text. August
24, 1953.

3. The silky pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker.
Pp. 339-347, 1 figure in text. February 15, 1954.

4. North American jumping mice (Genus Zapus). By Philip H. Krutzsch. Pp.
349-472, 47 figures in text, 4 tables. April 21, 1954.

5. Mammals from Southeastern Alaska. By Bollin H. Baker and James S
Findley. Pp. 473-477. April 21, 1954.

6. Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479-
487. April 21. 1954.

7. Subspeciation in the montane meadow mouse. Microtus montanus, in Wyo-
ming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text.
July 23, 1954.

8. A new subspecies of bat (Myotis velifer) from southeastern California and
Arizona. By Terry A. Vaughan. Pp. 507-512. July 23, 1954.

9. Mammals of the San Gabriel mountains of California. By Terry A. Vaughan.
Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954.

10. A new bat (Genus Pipistrellus) from northeastern Mexico. By Rollin H.
Baker. Pp. 583-586. November 15, 1954.

11. A new subspecies of pocket mouse from Kansas. By E. Raymond Hall. Pp.
587-590. November 15, 1954.

12. Geographic variation in the pocket gopher, Cratogeomys castanops, in Coa-
huila, Mexico. By Robert J. RusseU and Rollin H. Baker. Pp. 591-608.
March 15, 1955.

13. A now cottontail (Sylvilagus floridnnus) from northeastern Mexico. By Rollin
H. Baker. Pp. 609-612. April 8, 1955.

14. Taxonomy and distribution of some American shrews. By James S. Findley.
Pp. 613-618. June 10, 1955.

15. The pigmy woodrat, Neotoma goldmani, its distribution and systematic posi-
tion. By Dennis G. Rainey and Rollin H. Baker. Pp. 619-624, 2 figures in
text. June 10, 1955.

Index. Pp. 625-651.

(Continued on inside of back cover)

University of Kansas Publications
Museum of Natural History

Volume 11, No. 8, pp. 443-516
August 1, 1959

Birds From Coahuila, Mexico

BY

EMIL K. URBAN

University of Kansas

Lawrence

1959

University of Kansas Publications, Musexjm of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 11, No. 8, pp. 443-516
Published August 1, 1959

University of Kansas
Lawrence, Kansas

MUS. COMP. ZOOL
LIBRARY

SEP -3 1959

HARVARD
UNIVERSITY

PRTNTED IN

THE STATE PRJNTING PLANT

TOPEKA. KANSAS

I 959

Birds From Coahuila, Mexico

BY

EMIL K. URBAN

INTRODUCTION

The following account is a summary of the present knowlege
of the birds of Coahuila. Some 500 specimens from Coahuila in
the Museum of Natural History at the University of Kansas are
the basis for this report; these are supplemented by records of
birds previously listed from the State.

In Coahuila, habitats vary from those characteristic near tree-
line to those of the floors of the low deserts. Because of the
variety of habitats, many kinds of birds are present in the State;
at least 312 living named kinds of 249 species have been recorded.
Possibly another 100 species will be reported after further studies
have been made there. At least 154 of the species listed in this
paper probably breed in Coahuila. The bird fauna in the State
includes species characteristic of eastern North America and of
western North America, species that range from the Atlantic to
the Pacific Ocean, and species found only, or mostly, in Mexico.

I thank Professor E. Raymond Hall, Doctor Richard F. Johnston
and Doctor Robert M. Mengel for their kind help, and Doctor
Harrison B. Tordoff for first suggesting this study to me. Unless
otherwise stated, the nomenclature in this paper is that of the
A. O. U. Check-list Committee (1957). Catalogue numbers are
those of the Museum of Natural History at the University of
Kansas. In so far as known to me, all birds recorded in the litera-
ture from Coahuila are listed below. In a few instances the only
support for occurrence is the ascription of a given kind to Coahuila
(without mention of date, catalogue number, or precise locality)
by Friedmann, Griscom, and Moore (1950), and/or the A. O. U.
Check-list Committee (1957); when this is so the entire entry is
inclosed within brackets. In the accounts beyond, an asterisk in-
dicates that the kind breeds in Coahuila; two asterisks indicate
probable breeding in the State.

(445)

446

Uni\trsity of Kansas Publs., Mus. Nat. Hist.

LIST OF COLLECTORS

Persons who have obtained specimens of birds from Coahuila
for the Museum of Natural History are as follows:

Albert A. Alcorn
Joseph Raymond Alcom
Sydney Anderson
Rollin Harold Baker
James Sheldon Carey
Peter Stanley Chrapliwy
W. Kim Clark
Robert William Dickerman
John R. Esther
James Smith Findley
John Keever Greer

John William Hardy
Gcrd H. Jleinrich
William McKee Lynn
Jack M. Mohler
Roger O. Olmstead
Robert Lewis Packard
Robert Julian Russell
William J. Schaldach, Jr.
Harrison Bruce Tordoff
South Van Hoose, Jr.
Olin Lawrence Webb

GAZETTEER OF LOCALITIES IN COAHUILA

The following place-names were used to record the localities
of Coahuilan birds now specimens in the University of Kansas
Museum of Natural History. Each place-name is followed by
its location in degrees and minutes of latitude and longitude,
respectively.

Ocampo.— -27°22', 102°26'.

Paila.— 25°38', 102°09'.

Parras.— 25°25', 102° 12'.

Piedras Blanca.— 29°02', 102°33'.

Piedras Negras.— 28M3', 100°32'.

Sabinas.— 27°52', 101°07'.

Saltillo.— 25°26', 101°00'.

San Antonio de las Alazanas. —
25° 16', 100°37'.

San Buenaventura.— 27°06', 101 °32'.

San Francisco.— 27°37', 102°37'.

San Geronimo.— 28°30', 101 MS'.

San Isidro.— 27°33', 102°27'.

San Juan de Sabinas. —
27°55', lOVlT.

San Lorenzo.— 25°28', 102° 12'.

San Marcos.— 26°41', 102''07'.

San Miguel.— 29° 14', 101 °22'.

San Pedro de las Colonias (San
Pedro).— 25°45', 102°58'.

Santa Teresa.— 26°27', 101 °21'.

Tanque Alvarez.— 27°56', 102°38'.

Torrc6n.— 25°33', 103°27'.

Villa Acuiia.— 29°19', 100°56'.

Acebuches.— 28°17', 102°56'.
Americanos.— 27° 12', 103° 14'.
Australia.— 26° 18', 102° 18'.
Bella Union.— 25°26', 100°51'.
Boquillas.— 29°11', 102°55'.
Castillon.— 28°21', 103°33'.
Cuatro Cienegas.— 26°58', 102°04'.
Diamante.— 25°22', 100°54'.
Don Martin.— 27°32', 100°37'.
Fortin.— 28°48', 10r41'.
General Cepeda.— 25°22', 101°28'.
Gomez Farias.— 24°58', 101 °02'.
Hermanas.— 27°13', 101°13'.
Iglesias.— 27°34', 101 °20'.
Jaco.— 27°50', 103°55'.
Jimenez.— 29°04', 100°42'.
La Babia.— 28°33', 102°03'.
La Gacha.— 28°09', 101°31'.
La Mariposa.— 28° 12', 101°49'.
La Ventura.— 24°48', 100°38'.
Las Delicias.— 26°10', 102°49'.
Las Margaritas.— 28°42', 101° 14'.
Mesa de Tablas.— 25°14', 100°24'.
Muzquiz.— 27°53', 101°32'.
Nava.— 28°25', 100°46'.

Birds From Coahuila, Mexico 447

For mountain ranges, the approximate center of the highland
of each range is used as the point of reference.

Pico de Jimulco.— 25°08', 103° 16'. Sierra de Guadalupe.—
Sierra del Carmen.— 25° 13', 10r32'.

29°00', 102°30'. Sierra del Pino.— 28°15', 103'03°.

Sierra de la Encantada. — Sierra de la Madera. —

28°25', 102°30'. 27°03', 102°30'.

DISTRIBUTION OF THE KNOWN BREEDING BIRDS

OF COAHUILA

Topography and Climate

Coahuila lies in the broad northern end of Mexico, immediately
east of the center of the continental mass. The mountains of Coa-
huila, which are part of the Rocky Mountain-Sierra Madre Ori-
ental Axis, extend in a north-south direction and divide the lower
lands into two areas, a larger one, a part of the Central Plateau,
to the westward and a smaller one, a part of the Gulf Coastal Plain,
to the northeastward. Most of the mountains of Coahuila do not
exceed 6000 feet in elevation. A few peaks such as in the Sierra
del Carmen, Sierra del Pino, Sierra de la Madera, Sierra Encar-
nacion, and Sierra de Guadalupe, are more than 9000 feet high,
and some more than 10,000 feet in elevation occur near the south-
eastern border of the State in the Sierra Madre Oriental. The Gulf
Coastal Plain of northeastern Coahuila ranges from 700 feet to
1800 feet. The desert plains of the Mesa del Norte to the west of
the Sierra Madre Oriental Axis are higher, more rugged, and more
dissected than those of the Coastal Plain and are marked by scat-
tered desert ranges, buttes, low hills, and knobs.

Most of Coahuila is arid. Rainfall is moderate on the Coastal
Plain and is low west of the central mountains. Baker (1956:128-
132) and Muller (1947:35-38) give good summary discussions of
the topography and climate of Coahuila, and the reader is referred
to these for further details.

Biotic Communities

Baker (1956:132) stated that "the biotic communities of Coahuila
might be divided in accordance with the three physiographic areas
of the State: the Gulf Coastal Plain, the mountains, and the desert
plains of the Mesa del Noite." Goldman and Moore (1945:348-
349) hsted three biotic provinces in Coahuila: the Chihuahua-
Zacatecas Biotic Province, in the western half of the State; the
Tamaulipas Biotic Province, in the northeastern part of the State;

448 University of Kansas Publs., Mus. Nat. Hist.

and the Sierra Madre Oriental Biotic Province, in the southeastern
part of the State. Merriam (1898) noted that definable portions
of the Lower Sonoran Life-zone, the Upper Sonoran Life-zone, the
Transition Life-zone, and the Canadian Life-zone can be distin-
guished in Coahuila. In my study of the distribution of the avi-
fauna of Coahuila, I found that the three biotic provinces listed by
Goldman and Moore (op. cit.) as major headings and Merriam's
life-zones as supplements are the most satisfactory divisions.

The Tamaulipas Biotic Province. — This province consists of low-
land plains and a few isolated ranges of low mountains. The aver-
age rainfall is 23 inches (Baker, 1956:130), considerably more than
the 10 inches falling in the western part of the State. In the north-
eastern section of the State, the moderate amount of rain, mesic
vegetation, and close proximity to the eastern migration pathway
importantly influence the types of birds found.

In Coahuila, the Coastal Plain and the Rio Grande Plain lie in
the path of the northernmost trade winds; they account for the
more humid eastern slopes of the mountains of the northeastern
part of the State (Muller, 1947:38). Nevertheless, the northeastern
section of the State is semi-arid and can be placed in the Lower
Sonoran Life-zone. The vegetation consists mainly of thorny shrubs
and small trees with a liberal admixture of yuccas, agaves, and
cacti, and closely resembles that of southern Texas, northern Nuevo
Leon, and northern Tamaulipas (Goldman and Moore, 1945:354).

Migrant birds from the eastern flyway and less commonly mi-
grants from western North America pass through northeastern Coa-
huila. The following breeding birds seem to be associated with
this province: Harris' Hawk, Bobwhite (C. v. texanus). Scaled
Quail (C. s. castanogastris). Yellow-billed Cuckoo, Groove-billed
Ani, Green Kingfisher, Golden-fronted Woodpecker, Hairy Wood-
pecker (D. V. intermedius) , Ladder-backed Woodpecker (D. s.
symplectus), Vermilion Flycatcher (P. r. mexicantis). Cave Swal-
low, Gray-breasted Martin, Black-crested Titmouse (P. a. atri-
cristatus), Carolina Wren, Long-billed Thrasher, Curve-billed
Thrasher (T. c. oberhoheri). Blue-gray Gnatcatcher (P. c. cae-
rulea), Hutton's Vireo (V. ^. carolinae). Bell's Vireo (V. b. medius).
Yellow-throated Vireo, Red-eyed Vireo, Summer Tanager (P. r.
rubra), Olive Sparrow, Cassin's Sparrow, and Black-throated Spar-
row (A. b. bilineata).

The Sierra Madre Oriental Biotic Province. — Southeastern Coa-
huila is in this province that includes mountains in southern Nuevo

Birds From Coahuila, Mexico 449

Leon, southwestern Tamaulipas, and eastern San Luis Potosi. Areas
classifiable as Canadian, Transition, Upper Sonoran, and Lower
Sonoran in life-zone are found in this province. This region of
Coahuila receives the highest rainfall; this is evidenced by the lux-
uriant growth of boreal plants living in the higher places there
(Baker, 1956:131). Spruce, pine, and aspen occur at higher ele-
vations and oaks, thorny shrubs, and grasslands are present lower
down.

Birds of central or southern Mexico reach the southern part of
Coahuila; the Thick-billed Parrot, Hooded Yellowthroat, and
Rufous-capped Atlapetes are examples. A boreal forest on the
higher slopes of the mountains of southeastern Coahuila is suitable
for certain northern birds such as Goshawks, Pine Siskins, and
Brown Creepers. Some species of birds ordinarily associated wdth
western North America are present in Coahuila only in its south-
eastern part; striking examples of disjunction in range thus occur.
Probably sometime in the past these birds were distributed through-
out most of Coahuila. When this area became arid, these species
disappeared from all of Coahuila except from the high mountains
in the southeastern part. For example, Steller's Jay and the Scrub
Jay are absent in the Sierra del Carmen of northwestern Coahuila
but do occur in southeastern Coahuila.

Migrants of the eastern flyway as well as migrants associated
with western North America pass through this section of Coahuila.
The following breeding birds are associated with this province:
Goshawk, Band-tailed Pigeon, Thick-billed Parrot, Golden-fronted
Woodpecker, Ladder-backed Woodpecker (D. s. giraudi). Pine
Flycatcher, BuflF-breasted Flycatcher, Vermilion Flycatcher (P. r.
mexicanus), Steller's Jay, Scrub Jay, Mexican Chickadee, Black-
crested Titmouse ( P. a. atricristatiis ) , Cactus Wren ( C. h. guttatus ) ,
Robin, Blue-gray Gnatcatcher (P. c. amoenissima) , Button's Vireo
(V. h. stephensi). Bell's Vireo (V. b. medius), Hartlaub's Warbler,
Summer Tanager (P. r. cooperi). Pine Siskin, Rufous-capped At-
laptes, and Black-throated Sparrow (A. b. grisea).

The Chihuahiia-Zacatecas Biotic Province. — This province in
Coahuila covers the arid, interior, western desert area; it consists
of rolling plains with mountains that rise islandlike above the gen-
eral surface. Some of the mountains, such as in the Sierra del
Carmen and the Sierra del Pino, are more than 9000 feet high.
The major part of this biotic area lies within the Lower Sonoran
Life-zone. Areas of the Transition and Canadian life-zones are

450 University of Kansas Publs., Mus. Nat. Hist.

present on some of the higher mountains; their discontinuity re-
sults in a discontinuous distribution of the conifer-dependent avi-
fauna.

The large desert restricts the movement of birds considerably.
Major results of this include isolation of certain populations and
absence of others in the boreal islands. For example, Miller ( 1955a:
157) noted that the "dispersal of conifer-belt birds to and from the
Sierra del Carmen, although not as difficult as to well separated
islands [such as off the coast of Baja California], is nevertheless a
formidable matter to accomplish across the great deserts of Texas,
Chihuahua, and Coahuila." Miller (loc. cit.) noted also that the
avifauna of the Sierra del Carmen, due to its insularity, is unbal-
anced and stated that "as a consequence of unbalance, species that
are present show ecologic extension and unusual numerical rela-
tions." At least in this type of environment, an extension or ex-
pansion of the ecologic habits of the related types takes place
when some species are absent.

This isolation influences local variation among some of the birds
found in Coahuila. Niches elsewhere usually occupied by certain
species, absent here, are occupied by other species. These other
species thus enjoy an ecologic freedom and can expand their
niches in the absence of related types of similar ecologic scope.
For example, Miller (1955a: 158-159) reported that Hairy Wood-
peckers occurred only casually in the Sierra del Carmen and that
the Ladder-backed Woodpecker has spread out and seems to
occupy the niche or niches usually characteristic of the Hairy
Woodpecker. Changes usually thought of as of subspecific char-
acter seem to be taking place between the Ladder-backed Wood-
peckers of the Sierra del Carmen and of other areas, possibly be-
cause the Ladder-backed Woodpecker in the Sierra del Carmen is
extending its ecologic sphere more than in areas where the Hairy
Woodpecker exists. Restriction in dispersal due to geographic
isolation has probably hindered gene flow, thus allowing rapid
local adaptation, recognizable in variation at the infraspecific level.
Miller {loc. cit.) listed other birds that have expanded their eco-
logic scope; his work should be referred to for further details.

The following birds are associated with this province: Black
Vulture, Scaled Quail (C. 5. pallida), Turkey, Elf Owl, Green
Kingfisher, Hairy Woodpecker (D. v. icastus). Ladder-backed
Woodpecker (D. s. cactopliilus) , Wied's Crested Flycatcher, Buff-

Birds From Coahuila, Mexico 451

breasted Flycatcher, Vermilion Flycatcher (P. r. flammeus). Black-
crested Titmouse (P. a. dysleptiis), Cactus Wren (C. b. couesi).
Curve-billed Thrasher (T. c. celsum). Blue-gray Gnatcatcher (P. c.
amoenissima) , Hutton's Vireo (V. h. carolinae). Summer Tanager
(P. r. cooperi), and Black-throated Sparrow (A. fo. opuntia). Sev-
eral kinds of birds, such as the Band-tailed Pigeon, occur in the
"pine islands" in this province rather than on the desert floor.

There remain several kinds of birds that are not especially asso-
ciated with any one or two of the above-named provinces. These
birds are widely distributed and vary geographically without cor-
responding to the Biotic Provinces. Examples of these species are:
Black Phoebe (S. n. semiatra in northern Coahuila; S. n. nigricans
in southern Coahuila), Violet-green Swallow (T. t. lepida in north-
western Coahuila; T. t. thalassina in southeastern Coahuila), Black-
eared Bushtit (P. m. llotjdi in northern Coahuila; P. m. iuhis in
southeastern Coahuila), White-breasted Nuthatch (S. c. nehoni in
northern Coahuila; S. c. mexicana in southern Coahuila), Brown-
throated Wren ( T. b. cahooni in northern Coahuila; T. b. composi-
tus in southern Coahuila ) , Crissal Thrasher ( T. d. dorsale in north-
ern Coahuila; T. d. dumomm in southern Coahuila), and Rufous-
crowned Sparrow (A. r. tenuirostris in northern Coahuila; A. r.
boucardi in southern Coahuila ) .

Some representatives of the avifauna of the central and southern
sections of the Central Plateau reach southwestern Coahuila. The
subspecies squamata of the Scaled Quail and eurhtjncha of the
Blue Grosbeak are examples. Each in Coahuila seems to be at
the northern limit of its range.

In summaiy, tliere are three associations of vegetation in Coahuila
and each has characteristic birds. Gross cHmate and topography,
through their influence on vegetation, are the prime factors in the
distribution and kinds of birds in the State. Some birds of central
and southern Mexico reach southeastern and southwestern Coahuila.
Representatives of the Gulf Coastal Plain in Tamaulipas and Nuevo
Leon as well as migrants of the eastern flyway occur in north-
eastern Coahuila. Most of the species that occur in Coahuila seem
to be associated with western North America. The aridity of
western Coahuila restricts, to a large extent, the diversity of the
breeding populations of its avifauna. Xeric conditions surround-
ing some of the higher mountains are barriers to movement of
some species.

452 Unrtrsity of Kansas Publs., Mus. Nat. Hist.

ORIGIN OF BREEDING BIRDS OF COAHUILA

Probably beginning in the late Pliocene and ending in the Ice
Age (Griscom, 1950:379) the refrigeration of climate in the North-
em Hemisphere initiated a period of southward withdrawal of
birds from the northern part of North America. Some members
of the avifauna of Coahuila probably reached the State in this
time. When the continental deserts were formed, or reformed,
many tropical and subtropical Middle American species were
forced to leave Coahuila. Species associated with arid conditions
found their way there. Many representatives of the Old World
element also seem to have found their way to the State during the
refrigeration of climate in the Northern Hemisphere. The separa-
tion of North and South America in the greater part of the Tertiary
(Mayr, 1946:9) that deterred mammals from intercontinental col-
onization seemingly did not hinder birds. Some South American
species moved northward into Mexico, all the way north to Coa-
huila.

The avifauna of Coahuila today is a mixture of the several
mentioned elements. Of the breeding populations, 43 per cent
breed in the western rather than the eastern United States, 6 per
cent breed in the eastern rather than the western United States,
30 per cent breed in both the eastern and western United States,
20 per cent are restricted to the Republic of Mexico, and the
southern parts of Arizona, New Mexico, and Texas, and 1 per cent
(Aztec Thrush and Rufous-capped Atlapetes) is endemic to the
Republic of Mexico.

It is instructive to consider also the origin of avifaunal elements
at the level of Family. According to Mayr (1946:11) most North
American families and subfamilies clearly originated in the Old
World, in South America, or from a North American element that
developed in the partial isolation of North America in the Tertiary.
Three other elements, the Panboreal, the Pan-American, and the
Pantropical are represented by some North American families and
subfamilies. Because of the obscurity of the place of origin of
certain groups, an additional unanalyzed element must be recog-
nized.

The Caprimulgidae and Picidae probably originated in North
America (Mayr, 1946:26). Although the Psittacidac are Pantropi-
cal in distribution, indications are that they probably originated in
the Old World (Mayr, 1946:17). The Phasianidae, Turdidae {My-
adestes-Hylocichla group), and Sylviidae ( Polioptilinae ) seem to

Birds From Coahuila, Mexico 453

have originated in the Old World (Mayr, 1946:27). However,
Mayr considered these groups to have had a secondary center of
proHferation in North America, and I thus consider these groups
to have a North American origin. Mayr (1946:27) considered
the Trochilidae, Tyrannidae, and Icteridae Pan-American in dis-
tribution; however, he suggested that they probably originated in
South America, and I here treat them as South American in origin.
No representatives of the Pan-American element that probably
originated in North America have been recorded from Coahuila
nor have members of the Panboreal element (Mayr, 1946:11) been
recorded in the State. According to my analysis, representatives
of families of birds known to breed in Coahuila and those that
probably breed there thus seem to have been derived historically
from the following sources:

Old World 24.7% South America 24.0%

North America 37.0% Unanalyzed 14.3%

Mayr (1946:28-29) gave examples of analysis by geographic
origin of the breeding species of several districts of North America.
For instance, at Yakutat Bay in southeastern Alaska the South
American element of breeding passerine species was 3 per cent,
the North American element 39 per cent, and the Old World ele-
ment 58 per cent whereas at Sonora, Mexico, the South American
element of breeding passerine species was 27 per cent, the North
American element 52 per cent, and the Old World element 21 per
cent. The breeding avifauna of Coahuila is thus in line with
Mayr's analysis, resembhng that of Sonora to a considerable degree
at the taxonomic level of Family.

ACCOUNTS OF SPECIES

**Podiceps caspicus (Hablizl). — On March 31, 1952, Olmstead saw "many
Eared Grebes" on a pond 10 mi. E Hacienda La Mariposa. This is the first
record of the Eared Grebe in Coahuila.

[Pelecanus erythrorhynchos Gmelin. — The White Pelican is uncommon, if
not rare; Friedmann, Griscom, and Moore ( 1950:21) list it.]

Anhinga anhinga (Linnaeus). — On March 31, 1952, Olmstead noted an
Anhinga perched on a submerged fence post in a lake 10 mi. E Hacienda La
Mariposa. This is the first record of the Anhinga in Coahuila.

**Ardea herodias Linnaeus. — Two subspecies of the Great Blue Heron,
treganzai and toardi, have been recorded from Coahuila. Friedmann, Griscom,
and Moore (1950:27) listed A. h. treganzai from the State; presumably this
subspecies occurs widely in low density. They (loc. cit.) remarked also that a
record of A. h. toardi from Coahuila "cannot be allocated subspecifically."

454 University of Kansas Publs., Mus. Nat. Hist,

Dickerman saw two Great Blue Herons in a marshy area at San Marcos
(=20 mi. S Cuatro Cienegas) on May 4, 1954. Van Tyne and Sutton
(1937:12) noted the Great Blue Heron "near Boquillas [Texas], along the
Rio Grande, on May 10 and 15. . . ."

**Butorides virescens (Linnaeus). — Ohnstead saw a Green Heron at Bo-
quillas, 700 feet, on March 10, 1952. Findley reported seeing Green Herons
2 mi. W Jimenez, 850 feet, on June 19, 1952, and 2 mi. S and 3 mi. E San
Juan de Sabinas on June 22, 1952.

'^'*Casmcrodius albus egretta (Gmelin). — The Common Egret is an uncom-
mon migrant in Coahuila. Friedmann, Griscom, and Moore (1950:30)
recorded C. a. egretta from the "extreme northern part" of Coahuila. Van
Tyne and Sutton (1937:12) saw the Common Egret "along the Rio Grande
on the Graham ranch just west of Boquillas," Texas, on May 16, which might
well be the locality to which Friedmann, Griscom, and Moore referred.
Olmstead saw a Common Egret at Don Martin on March 22, 1952.

**Ntjcticorax nycticorax hoactli (Gmelin). — This subspecies of the Black-
crowned Night Heron was listed from the "extreme north" section of Coahuila
by Friedmann, Griscom, and Moore (1950:32). Van Tyne and Sutton
(1937:14) saw three Black-crowned Night Herons along the Rio Grande
about two miles west of Boquillas, Texas, on May 16. This record probably
represents the locality to which Friedmann, Griscom, and Moore (op. cit.)
referred.

Nyctanassa violacea violacea (Linnaeus). — A Yellow-crowned Night Heron
in immediate post-juvenal plumage, No. 36413, was obtained on September
7, 1958, 16 km. south of Cuatro Cienegas, by W. L. Minckley. According
to him the bird was accompanied by "several" other herons seemingly of
the same species and condition of plumage. The species seems not to have
been recorded previously from Coahuila [Eds.].

Botaurus lentiginosus (Rackett). — Friedmann, Griscom, and Moore (1950:
34) listed the American Bittern from the "extreme northern part" of Coahuila.
Van Tyne and Sutton (1937:14) saw two representatives of this bittern
"along the Rio Grande not far from Hot Springs," Texas, on May 15. I
suspect that this is the locality to which Friedmann, Griscom, and Moore
(loc. cit.) referred.

[Branta canadensis leucopareia ( Brandt ) . — Friedmann, Griscom, and Moore
(1950:38) listed B. c. leucopareia from "northern Coahuila."]

**Dendrocygna autumnalis (Linnaeus). — Evendcn (1952:112) reported
a Black-bellied Tree Duck standing beside a reservoir in southern Coahuila
along the railroad between Saltillo, Coahuila, and Avalos, Zacatecas.

**Anas platyrhynchos Linnaeus. — On March 30, 1952, Olmstead recorded a
Mallard from 10 mi. E Hacienda La Mariposa, 2000 feet.

Anas strepera Linnaeus. — Specimen examined: one, S 31016, from 10 mi.
E Hacienda La Mariposa, 2000 ft., March 30, 1952.

The Gadwcll is not an uncommon spring migrant; Olmstead saw it 10 mi.
E Hacienda La Mariposa on March 30, 1952, and Baker observed it 8 mi. N

Birds From Coahuila, Mexico 455

and 4 mi. W Muzquiz on March 30, 1952. Friedmann, Griscom, and Moore
( 1950:42) recorded the Gadwell from Coahuila.

Anas acuta Linnaeus. — Miller (1955a: 161) took a Pintail on September 10
in the Sierra del Carmen.

Anas carolinensis Gmelin. — The Green-winged Teal has been recorded from
northern Coahuila. Van Tyne and Sutton (1937:15) recorded two mated
pairs along the Rio Grande at Lajitas, Texas, on May 10. Miller (1955a: 161)
remarked that a male of the year was taken in the Sierra del Carmen on
September 4.

Anas discors discors Linnaeus. — Specimens examined: total 2: sex ?
31646 and sex ? 31647 from .5 mi. S Las Margaritas, 2800 ft., September 28,
1953.

The Blue-winged Teal is a fairly common spring and fall migrant in Coa-
huila. Van Tyne and Sutton (1937:15) noted the Blue-winged Teal at several
different localities along the Rio Grande: "on May 8, four males and several
females resting on a mud bar along the Rio Grande near Hot Springs [Texas];
. . . on May 7, three pairs in a flock, along the Rio Grande, Castalon
[Texas]; . . . and on May 20, three pairs, along the Rio Grande, San
Vicente [Texas]." Miller (1955a: 161) reported that Marsh took a male of
the year in the Sierra del Carmen on September 10. Dickerman observed
Blue-winged Teal 8 mi. E and 2 mi. S Americanos on May 18, 1954. Olm-
stead listed Blue-winged Teal from 10 mi. E Hacienda La Mariposa on March
30, 1952. Nos. 31646-31647, which are probably females, represent the sub-
species discors because the light edgings of their crowns are definitely present;
the areas of their backs are brownish, not more intensively black, and their
underparts are brownish, less blackish.

**Anas cyanoptera septentrionalium Snyder and Lumsden. — Van Tyne and
Sutton (1937:15) listed several locahties along the Rio Grande in Brewster
County, Texas, where Cinnamon Teal were seen. I suspect that Friedmann,
Griscom, and Moore (1950:41) referred to those locahties. Dickerman saw
four pairs of Cinnamon Teal 14 mi. E and 16 mi. N Ocampo on May 9, 1954,
and also saw Cinnamon Teal 8 mi. E and 2 mi. S Americanos on May 18, 1954.

Mareca americana (Gmelin). — The American Widgeon is a fairly common
spring migrant in Coahuila. Olmstead observed this duck 10 mi. E Hacienda
La Mariposa on March 30, 1952. Dickerman saw five to seven American
Widgeons 8 mi. E and 2 mi. S Americanos on May 18, 1954.

Spatula cltjpeata (Linnaeus). — The Shoveler is a spring and probably fall
migrant in Coahuila, and has been observed at several localities. Van Tyne
and Sutton (1937:16) saw two pairs along the Rio Grande at Castalon, Texas,
on May 7 and saw "a fair-sized flock along the Rio Grande on the Johnson
ranch [in Texas] on May 13 and 14." Dickerman saw 12 pairs of Shovelers
on two ponds 14 mi. E and 16 mi. N Ocampo on May 9, 1954, and 10 more
8 mi. E and 2 mi. S Americanos on May 18.

Aythya affinis (Eyton). — Olmstead observed Lesser Scaup 10 mi. E Haci-
enda La Mariposa on March 30, 1952.

456 Univ'ERSity of Kansas Publs., Mus. Nat. Hist.

[Bucephala albeola (Linnaeus). — Friedmann, Griscom, and Moore (1950:
44) listed the Bufflehead from the State.]

*Cathartes aura aura (Linnaeus). — Specimen examined: one, $ 31017
(skeleton only), from 4 mi. W Hacienda La Mariposa, 2300 ft., March
26, 1952.

Miller (1955a: 161) took a female Turkey Vulture, which was in breeding
condition, in the Sierra del Carmen on April 17 and stated that "until more
statistics are available on breeding birds of northern Coahuila, they must be
considered C. a. aura. . . ." Amadon and Phillips (1947:577) took a Turkey
Vulture at Las Delicias which represented C. a. aura. Burleigh and Lowery
(1942:188) stated that this species was not uncommon, and was noted each
day soaring overhead both in the valleys and over the tops of the ridges
of southeastern Coahuila.

Friedmann, Griscom, and Moore (1950:47) listed C. a. teter from Coahuila.
Miller (1955a: 161) remarked that the subspecies aura and teter might inter-
grade in the Sierra del Carmen. At the present time it is possible to say
only that teter is present in Coahuila in migrant and wintering populations,
but the extent to which teter remains in northeastern Me.Yico is undetermined.
However, all indications point to this area as being the region where aura
and teter intergrade.

**Coragtjps atratus (Bechstein). — The Black Vulture is locally common
throughout most of eastern Coahuila but is uncommon in the western part
of the State. Sutton and Burleigh (1939a:25) noted the Black Vulture
"regularly east of Saltillo in low country," but did not see Black Vultures at
San Pedro or elsewhere in southwestern Coahuila. Burleigh and Lowery
(1942:188) stated that "the Black Vulture apparently avoids to a large
extent the higher altitudes, and only rarely was it observed at all, even about
Saltillo." Olmstead saw Black Vultures 8 mi. N and 4 mi. W Muzquiz, 1800
feet, on March 31, 1952, and Dickerman observed a flock at La Gacha
(=Rancho La Coucha), 1600 feet, on December 2, 1953.

**Accipiter gentilis (Linnaeus). — On July 6, 1955, Hardy saw a Goshawk
13 mi. E San Antonio de las Alazanas; this is the first record of occurrence
of this species from northeastern Mexico.

*Accipiter striatus velox (Wilson). — Specimen examined: one, $ 31018,
from along the Rio Grande (z=Boquillas), 700 ft., March 10, 1952, measure-
ments: wing, 207 mm.; tail, 171 mm.; tarsus, 53 mm.; culmen, 12 mm.

Our specimen of the Sharp-shinned Hawk is referred to velox on the basis
of the reddish, maculated breast, sides, and thighs. The collector's field
notes recorded the iris as blood-red. Marsh and Stevenson (1938:286)
thought that this subspecies was resident in the pine and Douglas-fir forest
of upper Vivoras Canyon of the Sierra del Carmen at 8500 feet, where
Marsh observed a family group including three immature birds. Friedmann
(1950:196) indicated that the immature male obtained by Marsh and Steven-
son is A. s. suttoni; Miller (1955a: 161), nevertheless, remarked that this
male has well barred feathers and thus is velox. Miller (loc. cit.) obtained
also an adult male of A. s. velox in the Sierra del Carmen at 7000 feet on
April 18.

Birds From Coahuila, Mexico 457

**Accipiter striatus suttoni van Rossem. — Specimen examined: one, $
32626, from 13 mi. E San Antonio, 9950 ft., July 6, 1955, measurements:
wing, 186 mm.; tail, 144 mm.; tarsus, 49 mm.; culmen, 11 mm.; weight,
103 gms.

The recording of A. s. suttoni in Coahuila by Friedmann, Griscom, and
Moore (1950:52), seems to have been based on their knowledge of the
specimen earlier mentioned by Friedmann (1950:196) and later identified by
Miller (1955a: 161) as A. s. velox. Therefore the KU specimen seems to be
the first record of A. s. suttoni in Coahuila. The size of its testes (right,
2.5x4 mm.; left, 3x4 mm.) does not indicate breeding; however, the time of
the year in which it was obtained suggests that it may have been a resident.

*Accipiter cooperii (Bonaparte). — Miller (1955a: 161) found Cooper's Hawk
breeding in the Sierra del Carmen on April 26.

Buteo jamaicensis borealis (Gmelin). — The Red-tailed Hawk is common in
Coahuila. Burleigh and Lowery (1942:188) noted the Red-tailed Hawk
on the higher ridges above an elevation of 6000 feet in southeastern Coahuila.
On April 17, Burleigh and Lowery (loc. cit.) saw two Red-tailed Hawks "in
tlie open valley south of Diamante Pass" and on April 20, "just outside of
Saltillo," these workers obtained an immature male that was referred to
B. j. borealis.

**Buteo jamaicensis fuertesi Sutton and Van Tyne. — Miller (1955a: 161)
took a male Red-tailed Hawk, on April 14 at 7000 feet in the Sierra del
Carmen, that was referred to as B. j. fuertesi. To my knowledge, there are
no other records of this subspecies from Coahuila, but this must be the
resident form over the bulk of western Coahuila.

There are several sight records of the Red-tailed Hawk. Olmstead saw one
16 mi. S Boquillas, 1600 feet, on March 6, 1952; Dickerman saw a Red-tailed
Hawk 16 mi. E and 18 mi. N Ocampo on May 7, 1954, one 20 mi. S Ocampo
on April 4, 1954; and an immature at Saltillo on January 17, 1954.

Buteo platypterus platypterus (Vieillot). — Specimen examined: one, $
32628, from 13 mi. E San Antonio de las Alazanas, 9950 ft., July 6, 1955.

The Broad-winged Hawk is rare in Coahuila. No. 32628, if a migrant,
was retarded from moving northward by the loss of its right foot and distal
one-third of its tarsus. Packard (1957:371) reported this specimen as the
first record of the species in Coahuila.

*Buteo swainsoni Bonaparte. — Specimens examined: total 2: $ 32022 from
2 mi. W Jimenez, 850 ft., June 20, 1952; and $ 29555 from Iglesias (=15 mi.
SW Sabinas), 1000 ft., August 22, 1949.

Swainson's Hawk is not common in Coahuila. The size of the testes (6x4
mm.) of No. 32022, the adult plumage, and the date (June 20) on which it
was obtained suggest that it was a breeding bird. Friedmann, Griscom, and
Moore (1950:55) reported that this species breeds as far east as Durango and
Chihuahua. Findley saw a Swainson's Hawk 2 mi. S and 3 mi. E San Juan
de Sabinas on June 22, 1952.

**Buteo albonotatus Kaup. — The Zone-tailed Hawk is uncommon in Coa-
huila. Sutton and Burleigh (1939a:26) noted the species "a few miles west
of Saltillo ... on January 30."

458 Unr'ersity of Kansas Publs., Mus. Nat. Hist.

Buteo regalis (Gray). — The Ferruginous Hawk is uncommon in Coahuila.
Sutton and Burleigh (1939a:26) noted a single bird "not far from Parras, on
January 30."

Buteo nitidus (Latham). — Evenden (1952:112) saw a Gray Hawk one
mile northeast of Saltillo, at Ramos Arizpe on March 4. Although there are
no other records of the Gray Hawk from Coahuila, its occurrence in the State
would be expected because this species has been recorded from Nuevo Leon
and Tamaulipas in northeastern Mexico (Friedmann, Griscom, and Moore,
1950:57).

**Parabuteo unicinctus harrisi (Audubon). — Harris' Hawk is fairly common
in southern Coahuila. Sutton and Burleigh (1939a:26) stated that Harris'
Hawk "was one of the few birds noted repeatedly in . . . southern Coa-
huila. . . ."

Circus cyaneus (Linnaeus). — The Marsh Hawk is a common migrant and
winter visitant in Coahuila. Miller (1955a: 161) remarked that the Marsh
Hawk was "seen in northward migratory flight across the desert east of the
Sierra del Carmen on March 31 and on April 11 along the west face of Loomis
Peak at 8800 feet." Olmstead saw a female Marsh Hawk 1 mi. W San
Buenaventura on April 2, 1952. Burleigh and Lowery (1942:188) noted the
Marsh Hawk "about Saltillo" and "above the summit of Diamante Pass at
about 8,000 feet" on April 14. Sutton and Burleigh (1939a: 26) noted winter-
ing Marsh Hawks "near San Pedro."

Pandion haliaettis (Linnaeus). — Miller (1955a: 161) reported seeing an Os-
prey on April 9 in Corte Madera Canyon, "apparently in migration;" this is
the only record from Coahuila.

'**Caracara cheriicatj (Jacquin). — From the few records in the literature, I
judge that the Caracara is uncommon in Coahuila. Evcndcn (1952:113)
saw three Caracaras "south of Saltillo" on March 5. Baker saw a Caracara in
the Sierra del Pino (=6 mi. NW Tanque Alvarez), 3400 feet, on July 6, 1953.
No specimens of the Caracara have been taken from Coahuila.

Falco mexicanus Schlegel. — Specimen examined: one, $ 31596, from Sal-
tillo, January 10, 1954.

The Prairie Falcon is an uncommon winter visitant in Coahuila. Friedmann,
Griscom, and Moore (1950:65) indicated that Falco mexicanus winters south
to Sonora, Oaxaca, Chiluiahua, Durango, Zacatecas, Auguascalientes, Hidalgo,
Nuevo Leon, and Tamaulipas.

No. 31596 is the first recorded specimen of the Prairie Falcon from Coahuila.
The bird was heavily parasitized by worms in the mesenteries and seems to be
an adult. Although its nuchal collar, as in immatures, is washed with pale
cinnamon-buff, its thighs are not heavily marked with dark brown spots. The
supercihary lines have blackish rather than brownish streaks, and the scapulars
do not have four or five dark bars (Friedmann, 1950:624).

Falco columharitis hendirei Swann. — Specimen examined: one, $ 31634,
from Don Martin Dam (=:Rio Salado), November 27, 1953, measurements:
wing, 191 mm.; tail, 111 mm.; tarsus, 37 mm.; culmen, 12 mm.; testes, 3x1
mm.

Birds From Coahuila, Mexico 459

The Pigeon Hawk seems to be uncommon in Coahuila. No. 31634, the
first record of this species in Coahuila, has pale gray interspaces on the
rectrices of its tail that are definitely wider than the three black bands, indi-
cating affinity with bendirei (Friedmann, 1950:702). Our bird was obtained
near the base of the Don Martin Dam of the Rio Salado, and was observed
hunting dragonflies over the water.

Falco sparverius sparverius Linnaeus. — Specimen examined: one, $ 31648,
from the north foot of Sierra Guadalupe (=10 mi. S, 5 mi. W General
Cepeda), 6400 ft., April 17, 1953, weight, 104 gms.

The Sparrow Hawk is locally common in Coahuila. Miller (1955a: 162)
noted the species occasionally in the lower canyon areas of the Sierra del
Carmen at 5000 feet from April 20 to 28. Dickerman saw two Sparrow
Hawks in the Sierra del Pino on May 12, 1954. Sutton and Burleigh
(1939a: 27) took a male at La Rosa on January 30 that was typical for
F. s. sparverius. Burleigh and Lowery (1942:188) noted Sparrow Hawks
almost daily about Saltillo and infrequently in the open valley south of
Diamante Pass; they took a female at Saltillo on April 22 that was assigned
to F. s. sparverius.

*Colinus virginianus texanus (Lawrence). — Specimens examined: total 9:
S 29408 and 9 29409 from 1 mi. S, 9 mi. W Villa Acuiia, April 2, 1950,
weights, 169.5 and 174.7 gms., S 29410 and 5 9 29411-29413 from 3 mi.
W, 1 mi. S San Geronimo, April 9, 1950, weights, 152.6, 158.6, 158.2, 159.0,
and 152.8 gms.; and $ $ 32032-32034 from 9 mi. S, 11 mi. E Sabinas, 1050
ft., June 13 and 14, 1952.

The Bobwhite is locally common in northeastern Coahuila. Aldrich and
Duvall (1955:18) indicated that C. v. texanus has been recorded from two
separate localities in northeastern Coahuila and from several localities in Texas
along the Rio Grande. Findley saw Bobwhites 2 mi. S and 3 mi. E San
Juan de Sabinas on June 22, 1952. The specimens of Bobwhite from 3 mi.
W and 1 mi. S San Geronimo seem to extend the known range of texanus
nearly 100 miles west. The sizes of the testes (11, 16, 15, 15 mm.) of
Nos. 29408, 32032-32034, respectively, indicates breeding by the Bobwhite
in Coahuila.

*Callipepla squamata pallida Brewster. — Specimens examined: total 16:
$ 29414 from 1 mi. S, 13 mi. W Villa Acuiia, April 4, 1950; $ $ 29415-
29417 from 11 mi. W Hacienda San Miguel, 2200 ft., April 4, 1950, weights,
205.3, 198.6, and 182.7 gms.; 5 5 31019-31021 from 10 mi. S, 5 mi. E
Boquillas, 1500 ft., March 2 and 4, 1952, weights, 184.4, 180 and 154.2
gms.; $ 34454 from 2 mi. SSE Castillon, 4050 ft., June 29, 1953, weight,
169 gms.; $ 29418 and 5 29419 from 8 mi. N, 2 mi. E La Babia, April 8,
1950; 5 $ 32023-32024, S 32026, 5 32025, and 5 32027 from 5 mi. N,
19 mi. W Cuatro Cienegas, 3250 ft., July 6, 1952; and $ 32640 from 2 mi.
W Paila, July 3, 1955.

The Scaled Quail is common in Coahuila. The subspecies pallida occurs
in northwestern Coahuila south to Sierra de los Alamitos. Intergrades of
pallida, castanogastris, and squamata are present in the central part of the
State. No. 32640, obtained 2 mi. W Paila, has some resemblance to
squamata. Five specimens of pallida from the central part of Coahuila (5
mi. N and 19 mi. W Cuatro Cienegas), show little or no approach toward
squamata. Miller (1955a: 162) stated that two of the Scaled Quail collected

2—7905

460 University of Kansas Publs., Mus. Nat. Hist.

in the Sierra del Carmen show no approach to the race castanogastris of
eastern Coahuila nor to C. s. squamata of southern Coahuila. From the
specimens that I have examined, I judge that the range of pallida extends
as far south as the Sierra de los Alamitos rather than only to the northwestern
part of Coahuila as reported by Aldrich and Duvall (1955:17). In north-
eastern Coahuila pallida seems to intergrade with castanogastris; No. 29414
has an indistinct rusty chestnut patch on its abdomen, thus resembhng castan-
ogastris.

The sizes of the testes (9-12 mm.) and of the largest ova (14 mm. in
diameter and an egg 23 mm. long) of birds labeled with reference to Cuatro
Cienegas indicate breeding activity.

**Callipepla squamata castanogastris Brewster. — Specimen examined: one,
S 32028, from 9 mi. S, 11 mi. E Sabinas, June 14, 1952.

Typical representatives of C. s. castanogastris in Coahuila occur only in
tlie extreme northeastern section of the State, and most specimens of the
Scaled Quail from northeastern Coahuila are intergrades between pallida and
castanogastris.

No. 32028 is identified as C. s. castanogastris because there is a distinct
rusty chestnut patch on its abdomen. This patch, nevertheless, is not so
large as in a more nearly typical male C. s. castanogastris from 15 mi. NNW
Anahuac, Nuevo Leon.

**CaUipepla squamata squamata (Vigors). — Specimens examined: total 2:
$ 30231 and 2 30232 from 10 mi. NW San Lorenzo, 4200 ft., Februar>' 3,
1951.

The subspecies squamata occurs in southern Coahuila. Amadon and
Phillips (1947:577) took a Scaled Quail at Las Delicias on August 18
that "was only two-thirds grown, though well able to fly" and obtained an
adult 19 mi. W Saltillo that was t>'pical squamata. Burleigh and Lowery
(1942:188) stated that C. s. squamata was one of the characteristic birds of
the open desert country of soutlieastern Coahuila. Scaled Quail were seen
by Burleigh and Lowery {loc. cit.) "about Saltillo and in the open valley
south of Diamante Pass."

The breast and upper back of both specimens from 10 mi. NW San
Lorenzo, are plumbeous-gray rather than pale dull gray. Also the lower
back, rump, abdomen, forehead, and crown more closely resemble the sub-
species squamata rather than C. s. pallida. However, the upper backs of
both specimens are not so plumbeous-gray as on a male (32030) and a female
(32031) of the subspecies squamata from 1 mi. N Chorro, Durango, 6450 ft.,
July 11, 1952. The two birds from Durango appear to be slightly darker
than the Coahuilan specimens that approach the subspecies pallida.

Burleigh and Lowery (1942:188-189) stated that one of their specimens
of C. s. squamata obtained at Saltillo seems to be "very close to castanogas-
tris, suggesting that southeastern Coahuila is in the region of intergradation
between the two races." Aldrich and Duvall (1955:17) indicated that squa-
rmita and casianogastris intergrade near Sabinas. The two specimens that I
have examined show no sign of approach toward castanogastris. More spec-
imens of Scaled Quail from Coahuila are needed to permit accurate defini-
tion of the distribution of the subspecies.

Birds From Coahuila, Mexico 461

*Cyrtonyx montezumae mearnsi Nelson. — The Harlequin Quail is locally
common in Coahuila; C. 7n. mearnsi is present in northwestern Coahuila (Al-
drich and Duvall, 1955:20). Miller (1955a: 162) stated that an area in the
head of Corte Madera Canyon of the Sierra del Carmen at 7500 feet was
the principal location for C m. mearnsi. He further suggested that the Harle-
quin Quail breeds in the Sierra del Camien and remarked that Marsh took
a male on September 7 at Jardin del Sur. He added that the occurrence of
C m. montezumae in northern Coahuila as reported by Friedmann, Griscom,
and Moore (1950:79) is "possibly an error or was based on individual dark
variants. . . ." Baker noted the Harlequin Quail (unidentified to sub-
species) 9 mi. W and 1 mi. S Villa Acuna, 1120 feet, on April 4, 1950.

^Cyrtonyx montezumae montezumae (Vigors). — This subspecies of the Har-
lequin Quail has been recorded from southeastern Coahuila. Ridgway and
Friedmann (1946:396) listed C. m. montezumae from Saltillo. Baker saw
a pair of Harlequin Quail (unidentified to subspecies) at San Antonio de las
Alazanas on March 25, 1950. More collecting is necessary for an under-
standing of the distribution and intergradation of these subspecies in Coa-
huila.

*Meleagris gallopavo intermedia Sennett. — Specimens examined: total 4:
9 31022 from Fortin (=33 mi. N, 8 mi. W San Geronimo), 3300 ft., March
28, 1952, weight, 9 lbs.; 9 29420 from 3 mi. W, 1 mi. S San Geronimo, April
9, 1950, weight, 7 lbs.; and $ 29421 and 9 29422 from 3 mi. W, 2 mi. S San
Geronimo, April 9, 1950, weights, 11.5 and 8.5 lbs.

The Turkey in Coahuila is restricted to the northern section of the State.
Miller (1955a: 162) remarked that the population of Turkeys in the Sierra
del Carmen was sparse and did not range above 7500 feet into the highest
pine-oak and Douglas fir areas. Baker saw Turkeys 4 mi. W Hacienda La
Mariposa, 2300 feet, on March 23, 1952. William Schaldach, Jr., noted 30
Turkeys "just west of Rancho San Geronimo" on April 9, 1950. Aldrich and
Duvall (1955:22) indicated several localities in northern Coahuila where
the Turkey occurs or occurred.

Grus canadensis (Linnaeus). — Sutton and Burleigh (1939a: 28) remarked
that a flock of Sandliill Cranes was heard "near Mayran ... on Janu-
ary 30."

Porzana Carolina (Linnaeus). — The Sora is an uncommon spring and faU
migrant in Coahuila. Amadon and Phillips (1947:577) obtained an adult
male Sora at Las Dehcias on August 15.

Fulica americana GmeUn. — The American Coot probably occurs in the
State as a spring and fall migrant. Dickerman saw two coots 8 mi. E and 2
mi. S Americanos on May 18, 1954.

*Charadrius vocifcrus vociferus Linnaeus. — Specimen examined: one, $
31023, from the Rio Grande (=17 mi. S Dryden, Terrell Co., Texas in Coa-
huila), 600 ft, March 19, 1952, weight, 82.7 gms.

In Coahuila the Killdeer seems to be uncommon. Van Tyne and Sutton
(1937:28) remarked that it nested in Brewster County, Texas. The species
probably nests in northern Coahuila as well; the lengths of the testes (left,
9 mm.; right, 4 mm.) of No. 31023 support this view.

462 Uni\trsity of Kansas Publs., Mus. Nat. Hist.

Eupoda montana (Townsend). — Pitelka (1948:118) recorded one female
Mountain Plover from Hipolito on February 23. Van Tyne and Sutton (1937:
28) reported that the Mountain Plover nested in Brewster County, Texas.
Possibly Eupoda montana nests in northern Coahuila as well.

Numenius americanus parvus Bishop. — Specimens examined: total 3: $
31434 and 9 35406 (skeleton only) from 8 mi. E, 2 mi. S Americanos, May
19, 1954, measurements: wing, 250, 258 mm.; tail, 96, 100 mm.; tarsus, 74,
81.5 mm.; culmen, 11.2, 16.5 mm.; $ gonad, 6x2 mm.; and 9 35400 (skeleton
only) from 7 mi. W San Antonio de las Alazanas, January 11, 1954.

The Long-billed Curlew is not common in Coahuila. Dickerman obtained
both the specimens from 8 mi. E and 2 mi. S Americanos out of a flock of
35. Sutton and Burleigh (1939a: 28) noted the Long-billed Curlew "near
San Pedro . . . [on] January 29."

Actitis macularia ( Linnaeus ) . — The Spotted Sandpiper has been obtained
from two localities in Coahuila. Miller (1955a: 162) stated that an immature
in fall migration was taken "at the tank in the western hills" of the Sierra del
Carmen on September 4. Burleigh and Lowery ( 1942:189) found the Spotted
Sandpiper "at the Chorro del Agua near Arteaga" on April 17 and 19.

Totanus melanoleucus (Gmelin). — Specimen examined: one, 9 31024, from
the Rio Grande (=17 mi. S Dryden, Terrell Co., Texas, in Coahuila), 600 ft.,
March 18, 1952, weight, 224 gms.

The Greater YeUowlegs is an uncommon spring and probably fall migrant
in Coahuila. No. 31024 is the first record of this species in Coahuila.

Totanus flavipes (Gmelin). — Miller (1955a: 162) reported that Marsh took
a Lesser YeUowlegs "at the tank in the western hills" of the Sierra del Carmen
on September 4.

Erolia melanotos (Vieillot). — Miller (1955a:162) reported taking a Pectoral
Sandpiper on September 4 "at the tank in the western hills" of the Sierra del
Carmen.

[Erolia minutilla (Vieillot). — Friedmann, Griscom, and Moore (1950:99)
listed the Least Sandpiper from Coahuila.]

*Recurvirostra americana Gmelin. — Specimens examined: total 2: $ 31433
and $ 31432 from 8 mi. E, 2 mi. S Americanos, May 19, 1954.

Van Hoose (1955:302) reported a small breeding colony of American Avo-
cets 8 mi. E and 2 mi. S Americanos "on a large grassy playa traversed by rows
of creosote (Larrea tridentato)." No. 31432 was taken from a nest con-
taining four partly-incubated eggs. Van Hoose (loc. cit.) also reported that
four eggs in a second nest contained well-developed, dowTiy young.

Steganopus tricolor Vieillot. — Wilson's Phalarope occurs in Coahuila as a
spring and probably fall migrant. Van Tyne and Sutton (1937:31) saw the
species several times along the Rio Grande. Friedmann, Griscom, and
Moore (1950:102) listed Steganopus tricolor from the Rio Grande along the
borders of Chihauhua and Coahuila on May 10-16.

[Larus argentatus smithsonianus Coues. — Friedmann, Griscom, and Moore
(1950:104) listed this subspecies of the Herring Gull from the State.]

Birds From Coahuila, Mexico 463

[Larus delawarensis Ord. — Friedmann, Griscom, and Moore (1950:103)
stated that the Ring-billed Gull occurs in Coahuila.]

*Columba fasciata fasciata Say. — Specimens examined: total 3: $ 35401
(skeleton only) from 22 mi. S and 5 mi. W Ocampo, 7000 ft., April 6, 1954;
and $ 32035 and 2 32036 from 4 mi. N, 21 mi. W Cuatro Cienegas, 6200 ft.,
July 2, 1952.

The Band-tailed Pigeon is locally common in Coahuila. Miller (1955a: 162)
remarked that the Band-tailed Pigeon "was unaccountably rare in 1953" in
the Sierra del Carmen, and said that a specimen was taken on August 7 in
Vivoras Canyon. Burleigh and Lowery (1942:189) wrote that "small flocks
were seen each day ... on the summit of Diamante Pass, but only on
one occasion was a pair noted." Dickerman saw 10 Band-tailed Pigeons 20
mi. S Ocampo, 6000 feet, on April 4, 1954. The enlarged testes (17 mm.)
of No. 32035, and an egg (38 mm. long) in No. 32036 show that the species
breeds in central Coahuila.

*Zencida asiatica asiatica (Linnaeus). — Specim.en exam,ined: one, 9 31025
(skeleton only), from 10 mi. S and 5 mi. E Boquillas, 1500 ft., March 4, 1952.

Miller (1955a: 163) took a male White-winged Dove on April 23 "at the
mouth of Boquillas Canyon at about 4900 feet" and remarked that this bird
might either be a straggler or a representative of a normal breeding popu-
lation (Z. a. asiatica). On January 29 and 30, Sutton and Burleigh (1939a:
29) saw White-winged Doves several times in southern Coahuila along the
highway from Saltillo to San Pedro.

**Zenaida asiatica mearnsi (Ridgway). — Amadon and Phillips (1947:577)
obtained an adult White-winged Dove of this subspecies at Delicias on
August 14.

Zenaidura macroura carolinensis (Linnaeus). — Friedmann, Griscom, and
Moore (1950:117) Hsted this subspecies of the Mourning Dove as a spring
and fall migrant on the "Central Plateau" (Coahuila is part of the "Central
Plateau,"), "as proven by banding records not indicated in literature."

*Zenaidura macroura marginella (Woodhouse). — Specimen examined: one,
$ 34455, from 1 mi. W Jaco, June 27, 1953.

Miller (1955a: 163) reported Z. m. marginella as occurring commonly in the
desert border and lower canyons at the base of the mountains of the Sierra
del Carmen, "but it occasionally ranged up to 6000 feet in openings in the
pine-oak belt." Burleigh and Lowery (1942:189) remarked that this dove was
fairly plentiful in the open desert country "about Saltillo, and was . . ,
noted in small numbers in Diamente Valley on April 17 and 19;" they also
said that the Mourning Dove was not seen above an elevation of about 7500
feet. Sutton and Burleigh (1939a: 29) observed Mourning Doves along
the highway across southern Coahuila. Mourning Doves were seen by Find-
ley 2 mi. W Jimenez, 850 feet, on June 19, 1952, and 2 mi. S and 11 mi.
E Nava, 810 feet, on June 15, 1952. Dickerman saw one in the Sierra del
Pino on May 12, 1954. Findley saw more than one 2 mi, S and 3 mi. E
San Juan de Sabinas on June 22, 1952, as did Dickerman at San Marcos
(=20 mi. S Cuatro Cienegas) on May 4, 1954.

464 University of Kansas Publs., Mus. Nat. Hist.

**Columbigalli7m passerina (Linnaeus). — The Ground Dove seems to be un-
common in Coahuila. Van Tyne and Sutton (1937:34) saw a single Ground
Dove fly across the Rio Grande into Coahuila at Lajitas, Texas, on May 10.
Findley saw one 2 mi. S and 3 mi. E San Juan de Sabinas on June 22, 1952.

"'* Scar da fella inca (Lesson). — The Inca Dove has been recorded from two
locahties in Coahuila. Hellmayr and Conover (1942:510) listed it from
Sabinas. Burleigh and Lowery (1942:189) collected a male and female on
April 16 and 19, respectively, "outside the city limits of Saltillo."

Leptotila verreauxi angelica Bangs and Penard. — Specimens examined: total
2: $ i 31026-31027 (skeletons only) from 4 mi. W Hacienda La Mariposa,
2300 ft., March 25, 1952.

The White-fronted Dove seems to be uncommon in Coahuila. Hellmayr and
Conover (1942:570) listed L. v. angelica from Sabinas.

**Rhynchopsitta pachyrhyncha terrisi Moore. — Specimens examined: total
4: S S 31531-31532 from 13 mi. E San Antonio de las Alazanas, 9345 ft.,
April 10, 1954, weights, 391.5 and 467.5 gms.; $ 31533 from 13 mi. E San
Antonio de las Alazanas, 10,000 ft., April 10, 1954, weight, 466 gms.; and
sex ? 31534 from Mesa de las Tablas, June, 1951.

The Thick-billed Parrot occurs in the southeastern section of the State,
where it is fairly common. Moore (1947:27-28) described this parrot as
Rhtjnchopsitta terrisi: he thought it differed decidedly from Rhijnchopsitta
pachyrhyncha. However, Hardy and Dickemian (1955:305-306) decided
that uniting the two forms as a single species better expresses their relationship.

Burleigh and Lowery (1942:189) reported seeing a small flock of Thick-
billed Parrots on the summit of Diamante Pass. Dickerman, in his field
notes, wrote that at a place 13 mi. E San Antonio de las Alazanas, 9345
feet, a large flock of about 300 birds was in a spruce-fir-pine-aspen association.

*Coccyztis americanus americanus (Linnaeus). — Specimens examined: total
2: $ $ 32037-32038 from 12 mi. N, 12 mi. W Jimenez, 850 ft, June 19, 1952,
measurements: vdng, 141, 146 mm.; tail, 142, 149 mm.; tarsus, 27, 27 mm.;
culmen, 25, 24 mm.

In Coahuila, the Yellow-billed Cuckoo seems to be uncommon. It occurs
in the northeastern section of the State, in the Gulf Coastal Plain (Baker,
1956:128), and probably breeds there. One subspecies, americanus, has been
recorded from Coahuila.

According to Ridgway (1916:13-17) the difference between C. a. ameri-
canus and C. a. occidentalis is size. His (loc. cit.) average measurements of
males of occidentalis are: wing, 149.6 mm.; tail, 147.1 mm.; tarsus, 26.7 mm.;
and culmen, 27.7 mm. whereas average measurements given by him of males
of americanus are: wing, 143.6 mm.; tail, 140.7 mm.; tarsus, 25.2 mm.; and
culmen, 26.4 mm. Van Tyne and Sutton (1937:35) question the value of
maintaining the subspecies occidentalis, because individuals of americanus
and occidentalis are almost impossible to tell apart. Friedmann, Griscom,
and Moore (1950:132) stated that americanus occurs in eastern North America
whereas occidentalis occurs in western North America. If the subspecies
occidentalis exists, then Nos. 32037 and 32038 are, by size, americanus and

Birds From Coahuila, Mexico 465

No. 32038 is an intergrade between the two subspecies ( or a large individual of
americanus ) .

The Yellow-billed Cuckoo was seen also by Findley 2 mi. S and 3 mi. E
San Juan de Sabinas on June 22, 1952, and by Dickerson at Torreon on July
2, 1955. The sizes of the testes of the birds from 12 mi. N and 12 mi. W
Jimenez (9, 10 mm. long) and the date (June 19) on which they occurred
there indicate that the birds possibly were breeding.

Coccyzus ertjthropthalmus (Wilson). — Miller (1955a: 163) reported a mi-
grant Black-billed Cuckoo taken in the maples and basswood near a water
hole in the bottom of Boquillas Canyon in the Sierra del Carmen, 5200 feet,
on April 22. Friedmann, Griscom, and Moore (1950:132) reported that this
cuckoo is presumably a regular transient in Mexico, but generally overlooked.

"^Geococcyx californianus (Lesson). — Specimen examined: one, 2 32049,
from 8 mi. N, 2 mi. W Piedras Negras, June 18, 1952.

Miller (1955a: 163) heard several Roadrunners calling at Boquillas Canyon
in the Sierra del Carmen, where he obtained two females. Burleigh and
Lowery (1942:190) stated that the species proved to be "unexpectedly scarce"
and was noted but once by them on April 22 when a single bird was observed
in "the open desert west of Saltillo." Sutton and Burleigh (1939a: 30) noted
that the Roadrunner was not common anywhere in southern Coahuila; they
obtained one female at San Pedro on January 29. The size of the largest ovum
(15 mm. in diameter) of No. 32040 indicates that this species breeds in
Coahuila.

*Crotophaga sulcirostris sulcirostris Swainson. — Specimen examined: one,
9 32039, from 2 mi. S, 3 mi. E San Juan de Sabinas, June 22, 1952.

No. 32039, obtained by Harrison B. Tordoff in a cypress woods along the
shore of a lagoon, provides the first record of the Groove-billed Ani in Coa-
huila. The size of its largest ovum (10 mm. in diameter) and the date indi-
cate breeding by this species in Coahuila.

**Tyto alba pratincola (Bonaparte). — The Bam Owl seems to be uncommon
in Coahuila. To my knowledge, there are two records of the Barn Owl in
Coahuila. Ridgway (1914:607) recorded this owl at the "head of Las Vacas
Creek." Miller (1955a: 163) heard the Barn Owl at 5000 feet in the oak belt
on April 25 in the Sierra del Carmen.

*Otus asio suttoni Moore. — O. a. suttoni is found in the higher country of
the Sierra del Carmen and western Coahuila. Miller (1955a: 163) stated that
Screech Owls were common in groves of oaks both at 7000 feet and 5000 feet
in the Sierra del Carmen, and remarked that his series from the Sierra del
Carmen matched well a series of suttoni from Chihuahua and Durango. Miller
(1955a: 163-164) also stated that Marsh took an adult at Jardi'n del Sur on
August 28 and said that the specimen from the Sierra del Carmen referred to
as cineraceus by Marsh and Stevenson (1938:286) agreed well with his series
of suttoni. O. a. suttoni probably occurs no farther east than the Sierra del
Carmen.

466 University of Kansas Publs., Mus. Nat. Hist.

*'''Otus asio mccalUi (Cassin). — Specimens examined: total 2: $ 32041
from 2 mi. W Jimenez, 850 ft., June 20, 1952; and sex ? 31645 from La Gacha,
1600 ft., December 9, 1953.

Three subspecies of the Screech Owl, Otus asio, occur in northeastern
Mexico; two of these, suttoni and mccallii, occur in Coahuila, the latter in the
eastern part. Tlie third subspecies, O. a. semplei, occurs still farther east, for
example in the state of Nuevo Leon on the Mesa del Chipinque 6 miles south
of Monterrey (Sutton and Burleigh, 1939b: 174).

Nos. 31645 and 32041 differ from semplei in that the tops of their heads do
not appear to be solid blackish brown at a distance of four to five feet and
in that the dark streakings of their backs and scapulars are not so heavy as in
semplei. The mentioned specimens are brownish, not more black and white
throughout as in suttoni nor are their toes heavily feathered (see Moore,
1941:154).

Findley observed a Screech Owl 2 mi. S and 3 mi. E San Juan de Sabinas
on June 22, 1952. La Gacha would seem to represent the western extent of
mccallii in Coahuila. O. a. mccallii and suttoni probably intergrade along the
eastern slope of the Sierra del Carmen. Tordoff took No. 32041 near a tree
that contained three young Screech Owls.

"^Otus flammeolus flammeolus (Kaup). — Specimens examined: total 2:
$ 31600 from 20 mi. S Ocampo, 6000 ft., April 4, 1954; and S 31581 from
13 mi. E San Antonio de las Alazanas, 9345 ft., April 9, 1954.

Miller (1955a: 163) collected seven Flammulated Owls in the pines and
oaks at 7000 feet in Carboneras Canyon and said that these owls were common
there. Nos. 31600 and 31581 are suffused with cinnamoneous pigmentation,
but represent the grayish phase, as described by Ridgway (1914:729). Van
Hoose (1955:302) previously recorded Nos. 31600 and 31581 from Coahuila.

*Buho virginianus pallescens Stone. — Specimens examined: total 2: 9 32042
from 2 mi. S, 12 mi. E Nava, June 15, 1952; measurements: wing, 367 mm.;
tail, 233 mm.; culmen, 29 mm.; and S 31677 from 1.5 mi. NE Las Mar-
garitas, 3100 ft., May 31, 1954; measurements: wing, 345 mm.; tail, 213 mm.;
culmen, 26 mm.; testes, 8 mm. long.

Miller (1955a: 164) took a male Great Homed Owl in the Sierra del Car-
men on April 22. Ridgway (1914:742) listed B. v. pallescens from Sabinas.
Sutton and Burleigh (1939a:30) obtained a female "near San Pedro" on Jan-
uary 29. Friedmann, Griscom, and Moore (1950:143) recorded B. v. pal-
lescens from Coahuila on February 24, May 26, and June 10. The record of
B. V. mayensis from Las Delicias (Amadon and Phillips, 1947:578) has been
reidentified by Webster and Orr (1958:141) as B. v. pallescens. Dickerman
saw a Great Horned Owl in the Sierra del Pino on May 12, 1954.

*Glaucidium gnoma californicum Sclater. — Specimen examined: one, $
31582, from 20 mi. S Ocampo, 6500 ft., April 5, 1954, weight, 55 gms.

Concerning forms of Pygmy Owls, Miller (1955a: 164) remarked that the
best distinguishing characters of G. g. gnoma and californicum are tail length
and wing length. The characters of No. 31582 (wing, 94 mm.; tail, 69 mm.)
are clearly those of californicum and not those of gnoma. Miller (loc. cit.)
remarked that he "encountered at least five different individuals, chiefly in
the pine-oak at 7000 feet" and one in oaks and piiions at 5000 feet in Bo-

Birds From Coahuila, Mexico 467

quillas Canyon of the Sierra del Carmen. The size of the testes (left, 9x5
mm.; right, 7x4 mm.) of No. 31582 indicates good probability of breeding
by the Pygmy Owl in the State.

**Micrathene whitneyi (Cooper). — Miller (1955a: 164) heard the Elf Owl
at close range in oaks at 5000 feet in Boquillas Canyon of the Sierra del
Carmen on April 24.

**Speottjto cunicularia hypugaea (Bonaparte). — Specimens examined: total
3: 9 32043 from 3 mi. S, 9 mi. E Cuatro Cienegas, 2250 ft., June 30, 1952;
$ 32653 from 6 mi. W San Antonio de las Alazanas, July 5, 1955; and $
31602 from 14 mi. W San Antonio de las Alazanas, 6500 ft., January 7, 1954.

Although there are no records in Coahuila of specimens of Burrowing Owls
north of 3 mi. S and 9 mi. E Cuatro Cienegas, this owl probably occurs in
the northern section of the State. The records of Burrowing Owls from the
southern part of Brewster County, Texas (Van Tyne and Sutton, 1937:38),
suggest such occurrence. No. 31602 was shot in a prairie dog colony; No.
32043 was captured in a steel trap. Baker saw Bvurowing Owls 7 mi. S and
4 mi. E Bella Union, 7200 feet, on June 25, 1952.

*Caprimulgus vociferus arizonae (Brewster). — Specimens examined: total
4: $ 31449 and 5 31450 from Sierra del Pino (=5 mi. S, 3 mi. W Ace-
buches), May 12 and 15, 1954, weights, 48 and 42 gms.; and $ $ 31028-31029
from 2 mi. N, 18 mi. W Santa Teresa, 7250 ft., AprU 3, 1952.

The Whip-poor-will occurs between 5000 feet and 9000 feet in Coahuila.
Miller (1955a: 164) reported C. v. arizonae in the Sierra del Carmen. Ridg-
way (1914:521) stated that Antrostomus vociferus macromystax i=C. v.
arizonae) occurs in the southeastern sector of Coahuila at Sierra Guadalupe.
Burleigh and Lowery (1942:190) reported the Whip-poor-will "near the
summit of Diamante Pass"; because of its size (wing, 170 mm.; tail, 135 mm.)
this Whip-poor-will from Diamante Pass seems to represent the subspecies
arizonae. Dickerman saw four Whip-poor-wills 20 mi. S Ocampo, 6000 feet,
on April 4, 1954. The sizes of the testes of Nos. 31449, 31028, and 31029
(13, 12, and 13 mm. long) and an egg taken from No. 31450 indicate
breeding by this species in the State.

*Phalaenoptilus nuttallii nuttallii (Audubon). — Specimens examined: total
6: $ 31032 from 37 mi. S, 21 mi. E Boquillas, 4100 ft., March 12, 1952;
$ 31446 and ? 31447 from Sierra del Pino (=5 mi. S, 3 mi. W Acebuches),
6200 ft., May 13, 1954; $ $ 32048-32049 from 2 mi. S, 11 mi. E Nava, June
16, 1952; and $ 31033 from 4 mi. W Hacienda La Mariposa, 2300 ft.,
March 26, 1952.

Miller (1955a: 164) found the Poor- will common along the rocky can-
yon walls up to 5000 feet in the Sierra del Carmen. Ridgway (1914:550)
recorded the Poor-will at Sabinas on May 21 and at Saltillo on May 6.
Van Tyne and Sutton (1937:39) stated that P. n. nuttallii was not com-
mon in the Big Bend Country of Texas; this probably is true for north-
western Coahuila as well. The specimens from the Sierra del Pino, col-
lected by Dickerman in a pine-oak association at 6200 feet, were taken near
the upper limit of their range. Findley saw Poor-wills 2 mi. W Jimenez, 850
feet, on June 19, 1952; 2 mi. S and 11 mi. E Nava, 810 feet, on June 15,
1952; and 2 mi. S and 3 mi. E San Juan de Sabinas on June 22, 1952.

468 University of Kansas Publs., Mus. Nat. Hist.

Dickerman noted Poor-wills 16 mi. E and 18 mi. N Ocampo on May 7, 1954,
and 20 mi. S Ocampo, 6000 feet, on April 4, 1954.

The size of the testes (9x5 mm.) of No. 31446 and an enlarged oviduct
and an ovum (4 mm. in diameter) of No, 31447 indicate breeding by the
Poor-will in Coahuila.

**Chordeiles minor howelli Oberholser. — Specimen examined: one, $
31443, from 3 mi. N, 4 mi, E San Francisco (=25 mi, N Ocampo), 4850
ft., May 16, 1954.

Van Hoose (1955:302) wrote that nighthawks were heard and seen
frequently 3 mi. N and 4 mi, E San Francisco. Blake (1953:227) said
that the Common Nighthawk breeds in Sonora, Chihuahua, Tamaulipas, and
Durango; the size of the testes (7x4.5 mm.) of No. 31443 indicates the
possibility of breeding by this species in the State.

*Chordeiles acutipennis texensis Lawrence. — Specimens examined: total 4:
S 32044 and 9 32045 from 2 mi. W Jimenez, 850 ft., June 20, 1952; $ 31442
from 5 mi. N, 13 mi. E Ocampo, May 6, 1954; and $ 32046 from 2 mi. N,
1 mi. W Ocampo, 4050 ft., July 6, 1952.

Specimens of the Lesser Nighthawk, subspecies C. a. texensis, have been
recorded in the literature from representative localities throughout most of
Coahuila. Burleigh and Lowery (1942:190-191) obtained, on April 18, at
"a small pond at the edge of Saltillo," one male that was exceedingly fat;
they (loc. cit.) suggested that their specimen was a migrant. Goldman
(1951:377, 389) stated that C. a. texensis occupied the Lower and Upper
Sonoran and Upper Austral life-zones of Coahuila. Dickerman saw Lesser
Nighthawks at San Marcos (=20 mi. S Cuatro Cienegas) on May 4, 1954.
Van Tyne and Sutton (1937:41) reported that the Lesser Nighthawk was
common throughout the lower parts of the Big Bend in Texas. This is
probably true for northwestern Coahuila as well.

The presence of an egg in the oviduct of No. 32045 and the dates (May
6, June 20, and July 6) on which Nos. 31442, and 32044-32046 were ob-
tained indicate breeding by this species in Coahuila.

**Aeronautes saxatalis saxatalis ( Woodhouse ) . — Specimen examined: one,
S 31672, from Pico de Jimulco, 5600 ft., April 5, 1953, weight, 35 gms.

Burleigh and Lowery (1942:191) found the White-throated Swift to be
common at "the summit of Diamante Pass and on the nearby ridges." Miller
(1955a: 164) saw the species from 4800 feet up to the crest of the Sierra
del Carmen. Several White-throated Swifts were seen flying overhead at
Pico de Jimulco on April 5.

No specimens of A. s. sclateri from Mexico are known. Miller (1955a: 165)
listed one specimen with dimensions (wing, 145 mm.) that approaches
sclateri. The mea.surements of No. 31672 (wing, 143 mm.; tail, 58 mm.)
also approach the dimensions of specimens of sclateri but are best referred to
A. s. saxatalis.

*'^Calothorax lucifer (Swainson). — Burleigh and Lowery (1942:191) ob-
tained a male Lucifer Hummingbird at the Chorro del Agua on April 19.
Van Tyne and Sutton (1937:43) reported a male from the Rio Grande (=3
mi. W Boquillas, Texas),

Birds From Coahuila, Mexico 469

Archilochus colubris (Linnaeus). — Friedmann, Griscom, and Moore (1950:
180) remarked that the Ruby-throated Hummingbird is a moderately com-
mon migrant, wintering from sea level to 9350 feet throughout Mexico,
except in a few states. The only published record of a specimen of this
hummer in the State is of a male taken on April 22 in a small arroyo twenty
miles west of Saltillo (Burleigh and Lowery, 1942:191).

* Archilochus alexandri (Bourcier and Mulsant). — Specimens examined:
total 2: $ 31035 from the Rio Grande (=17 mi. S Dryden, Terrell County,
Texas, in Coahuila), 600 ft., March 18, 1952; and $ 32052 from 2 mi. S,
11 mi. E Nava, 810 ft., June 16, 1952.

Miller (1955a: 165) stated the Black-chinned Hummingbird was common
in the desert area at the base of the mountains of the Sierra del Carmen,
and that Marsh, on July 25, obtained this hummingbird "near Piedra Blanca
(Conejo)." Burleigh and Lowery (1942:191) obtained a female in an
arroyo about twenty miles west of Saltillo on April 22.

**Selasphorus phtycercus platycercus (Swainson). — Specimens examined:
total 2: $ 31583 from 20 mi. S Ocampo, 6500 ft., April 4, 1954; and ? 32673
from 13 mi. E San Antonio de las Alazanas, 9950 ft., July 6, 1955.

Miller (1955a:165) indicated that the Broad-tailed Hummingbird was
fairly common in the Sierra del Carmen. According to Burleigh and Lowery
(1942:191) this species is not uncommon in the pine woods bordering the
summit of Diamante Pass; they saw the bird between 4900 and 10,000 feet.

*Eugenes fulgens fulgens Boucard. — Miller (1955a: 165) obtained a male
Rivoli's Hummingbird on Loomis Peak, 8800 feet, on April 11 in the Sierra
del Carmen.

**Lampornis clemenciae clemenciae (Lesson). — Specimens examined: total
2: S 31036 from 26 mi. W Santa Teresa, 7050 ft., April 5, 1952; and $
32668 from 13 mi. E San Antonio de las Alazanas, 9950 ft., July 6, 1955.

The Blue-throated Hummingbird occurs between 5000 and 9950 feet in
Coahuila. Miller (1955a: 165) remarked that it was found in canyon bottoms,
"whether at 7500 feet among the rocky slopes, oaks, and white pines . . .
or at 5000 feet in the madrone, maples, elms, and basswoods. . . ."
No. 32668, was obtained in a Douglas fir-pine-aspen association at 9950 feet.

[Amazilia yucatanensis chalconota Oberholser. — The Buff-bellied Hiun-
mingbird seems to be uncommon in Coahuila. Friedmann, Griscom, and Moore
(1950:172) and the AOU Check-Hst Committee (1957:306) stated that this
hummingbird occurs in Coahuila.]

Megaceryle alcyon alcyon (Linnaeus). — Miller (1955a: 165) saw a Belted
Kingfisher at Carboneras Canyon in the Sierra del Carmen. Miller, Fried-
mann, Griscom, and Moore (1957:15) stated tliat one specimen of M. a.
alcyon was obtained in Coahuila on November 14.

**Chloroceryle americana hachisukai (Laubmann). — Specimens examined:
total 2: 9 32053 from 12 mi. N, 12 mi, W Jimenez, June 19, 1952; and
2 31038 from 8 mi. N, 4 mi. W Muzquiz, 1800 ft., March 31, 1952.

The Green Kingfisher has been recorded in Coahuila as far south as 8 mi.
N and 4 mi. W Muzquiz. The forehead of No. 31038, when compared with

470 University of Kansas Publs., Mus. Nat. Hist.

typical representatives of C. a. hachisukai, is not extensively streaked with
white, nor are all the coverts conspicuously spotted with white, yet it clearly
has more extensive white markings than typical representatives of C. a. sep-
tentrionalis. This specimen from 8 mi. N and 4 mi. W Muzquiz probably is
intermediate between hachisukai and septentrionalis. Miller (1955a: 165)
stated that Marsh took a specimen at Tanque de los Melones on La Bavia
Ranch east of Fresno Mesa in the Sierra del Carmen that is a t>'pical C. a.
hachisukai.

Colaptes cafer coUaris Vigors. — Ridgway (1914:34) Usted this subspecies
of the Red-shafted Flicker from Saltillo and Agua Nueva.

*Colaptes cafer nanus Griscom. — Specimens examined: total 3: 9 31463
from Sierra del Pino (=5 mi. W, 3 mi. S Acebuches), May 13, 1954, weight,
108 gms.; and $ 31651 and $ 31652 from the north foot of Sierra Guadalupe
(=10 mi. S, 5 mi. W General Ccpeda), 6500 ft., April 21, 1953.

In suitable habitats in Coahuila the Red-shafted FUcker is common.
Miller (1955a: 165-166) stated that C. c. nanus was common at Corte Madera
Canyon, Boquillas Canyon, and Carboneras Canyon in the Sierra del Cannen
and recorded a specimen also from Sierra de Jardin on August 7. Burleigh
and Lowery (1942:192) recorded C. c. nanus from Diamante Pass and
Saltillo. Sutton and Burleigh (1939a:31) stated that the Red-shafted Flickers,
obtained 5 mi. E La Rosa and at Diamante Pass, may be Intermediate between
C. c. collaris (then called C. c. chihuahuae) and C. c. nanus. Nesting of the
Red-shafted FHcker in Coahuila was reported by Miller (1955a: 165) and Bur-
leigh and Lowery (1942:192). Hardy saw Red-shafted FHckers 13 mi. E San
Antonio de las Alazanas on July 6, 1955.

*Centurus aurifrons aurifrons (Wagler). — Specimens examined: total 6:
$ 32054 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 19, 1952; 9 9
32055-32057 from 2 mi. W Jimenez, 850 ft., June 20, 1952; 9 31039 from
8 mi. N, 4 mi. W Muzquiz, 1800 ft., April 1, 1952; and $ 33150 (skeleton
only) from Parras, July 4, 1955.

The Golden-fronted Woodpecker occurs throughout Coahuila, but uncom-
monly in the western part. Wetmore (1948:185-186) examined a series of
Golden-fronted Woodpeckers that showed a definite cline in dorsal coloration
from north-central Texas to Jalisco, Michoacan, and Hidalgo in south-central
Mexico. He (loc. cit.) stated that "the extremes arc easily separable, but in
southern and southwestern Texas and in northeastern Mexico the two kinds
[C. a. aurifrons and C. a. incanescens] . . , merge so gradually that
over a broad area the whole population is intermediate, making decisions as
to any sharply drawn dividing line difficult and in part arbitrary." C. a.
incanescens, according to Wetmore, occurs in western and central Texas south
to northeastern Chihuahua and northern Coahuila whereas C. a. aurifrons
occurs in north-central Coahuila (Monclova) and southern Texas south to
Jalisco, Michoacan, Hidalgo, and central Tamaulipas.

The specimens that I have examined from Coahuila are variably inter-
mediate between the subspecies aurifrons and incanescens. The dark and
white cross-bars on the back of No. 31039 are nearly equal (dark bands
wider in aurifrons; white bands wider in incanescens); the nunp and upper
tail coverts are more or less mixed with black as in aurifrons. The dark

Birds From Coahuila, Mexico 471

cross-bars on the backs of Nos. 32054-32057 are slightly larger than the
wliite cross-bars; the rump and upper tail coverts of these specimens are
somewhat mixed with black.

*Centurus aurifrons incanescens Todd. — This subspecies of the Golden-
fronted Woodpecker is listed by Miller, Friedmann, Griscom, and Moore
(1957:34) from ". , . northern Coahuila (upper Rio Grande valley)."
Burleigh and Lowery (1942:192) stated that the Golden-fronted Woodpecker
"apparently avoids the mountain slopes, but was found to be not uncommon on
the arid plateau about Saltillo." Findley saw Golden-fronted Woodpeckers
2 mi. W Jimenez, 850 feet, on June 19, 1952; and 2 mi. S and 3 mi. E San
Juan de Sabinas on June 22, 1952. Cory (1919:424) Hsted the Golden-
fronted Woodpecker from Sabinas. Nos. 32055-32057 are juveniles, and
thus document breeding by this woodpecker in Coahuila.

*Melanerpes formicivorus formicivorus (Swainson). — Specimens examined:
total 5: $ 31040 (skeleton only) from Fortin (=33 mi. N, 8 mi. W San
Geronimo), 3300 ft., March 28, 1952; $ 29423 and $ 29424 from Club
Sierra del Carmen (=2 mi. N, 6 mi. W Piedra Blanca), 4950 ft., April 8,
1950; $ 31041 (skeleton only) from 26 mi. W Santa Teresa, 7050 ft., April 5,
1952; and $ 31668 from Sierra Guadalupe, Caiion d. Meco (=10 mi. S
General Cepeda), 6500 ft., April 23, 1953.

The Acorn Woodpecker in Coahuila is common in the oak and pine-oak
belts, from 4950 to 8000 feet. Miller (1955a:166) stated that the Acorn
Woodpecker in the Sierra del Carmen was an abundant and conspicuous bird
throughout the oak and pine-oak belts, from 5000 to 8000 feet. Dickerman
saw two Acorn Woodpeckers in the Sierra de la Madera on December 13,
1953, and four 20 mi. S Ocampo, 6000 feet, on April 4, 1954. Breeding of
M. f. formicivorus was reported by Miller (loc. cit.) who took females nearly
ready to begin laying; one of our specimens (No. 29423) had enlarged testes
(11 mm.).

Sphyrapicus varius varius (Linnaeus). — Specimen examined: one, 9 31649,
from tlie north foot of Sierra Guadalupe (=10 mi. S, 5 mi. W General
Cepeda), 6400 ft., April 19, 1953.

Miller (1955a: 166) reported this Yellow-bellied Sapsucker as a winter
visitant or migrant in the Sierra del Carmen. Burleigh and Lowery (1942:
192) took a female S. v. varius at the summit of Diamante Pass on April 14.
Ridgway (1914:275) listed S. v. varius from Sierra de Guadalupe.

Sphyrapicus varius nuchalis Baird. — Miller (1955a: 166) reported this Yel-
low-bellied Sapsucker as "indeed common" in the Sierra del Carmen, and
indicated that both S. v. nuchalis and S. v. varius were "found only at the
upper levels in the pine-oak formation and usually in relatively dense clumps
of trees in the canyon bottoms." Ridgway (1914:280) listed C. v. nuchalis
from Rio Sabinas.

**Dendrocopos villosus icasius ( Oberholser ) . — Miller, Friedmann, Griscom,
and Moore (1957:39) remarked that this subspecies of the Hairy Woodpecker
occurs, in northwestern Mexico, from eastern Sonora, Jahsco, Chihuahua, Du-
rango, Zacatecas to southern Coahuila. Ridgway (1914:222) stated that D. v.
icastus occurs in Coahuila at Cameros and Sierra de Guadalupe, the former
being the easternmost known locahty for the subspecies.

472 University of Kansas Publs., Mus. Nat. Hist.

**Dendrocopos villosus intermedius (Nelson). — Specimens examined: total
2: $ 32701 from 13 mi. E San Antonio de las Alazanas, 9950 ft., July 6,
1955; and $ 31604 from 2 mi. E Mesa de Tablas, 8500 ft., January 15, 1954.

In northeastern Mexico this subspecies, according to Miller, Friedmann,
Griscom, and Moore (1957:39), occurs in Guanajuato, San Luis Potosi, Hi-
dalgo, Nuevo Leon, and Tamaulipas. Nos. 31604 and 32701 represent the
first records of D. v. intermedius from Coahuila.

*Dendrocopos scalaris cactophilus (Oberholser). — Specimens examined:
total 2: $ 31042 from 7 mi. S, 2 mi. E Boquillas, 800 ft., March 1, 1952;
and 9 31043 from 10 mi. S, 5 mi. E Boquillas, 1500 ft., March 3, 1952.

These specimens of the Ladder-backed Woodpecker show signs of intergra-
dation with D. s. sijmplectus. Both specimens are pale enough above for D. s.
syrnplcctiis, but the wing and the tail of each (wing, 102, 103 mm.; tail, 60,
65 mm. ) are too long for sijmplectus. This suggestion of intergradation is
not unexpected because symplectus occurs in northeastern Coahuila and
cactophilus in the Chisos Mountains of Texas. Miller (1955a:166) also ob-
tained one female D. s. cactophilus from the Sierra del Carmen that sug-
gested intergradation with symplectus.

Miller {lac. cit.) wrote that D. c. cactophilus was found chiefly in the
oaks and was common in the lower oak belt at 5000 feet. Tlie upper limit
of the range of the Ladder-backed Woodpecker, according to Miller, is 6800
feet. He reported the species as breeding in the Sierra del Carmen.

Miller (1955b:317) took a hybrid woodpecker representing a cross between
Dendwcopos villosus and Dendrocopos scalaris in the Sierra del Camien,
where, although Ladder-backed Woodpeckers were common, he found no
Hairy Woodpeckers.

^-'Dendrocopos scalaris symplectus (Oberholser). — Specimens examined:
total 2: $ 32058 from 2 mi. W Jimenez, June 20, 1952; and $ 31667 from
Sierra Guadalupe, Domingo Canyon (=10 mi. S, 14 mi. W General Cepeda),
6700 ft, April 18, 1953.

This Ladder-backed Woodpecker, according to the AOU Check-list Com-
mittee (1957:327) and Oberholser (1912:156), occurs in Texas (east of
Pecos), northeastern Coahuila, Nuevo Leon, and Tamaulipas. The area of
intergradation of D. s. symplectus and giraudi is in southeastern Coahuila.
The dark smoky underparts and the eqvial size of the white and black bars
of the upper parts of No. 31667 suggest intergradation with D. s. giraudi.
Yet, the size of the wing indicates that tliis specimen is closer to D. s. sym-
plectus. No. 32058 has characters of typical representatives of D. s. sym-
plectus. Burleigh and Lowery (1942:193) reported D. s. symplectus "in the
desert country west of Saltillo." Ridgway (1914:257) listed the Ladder-
backed Woodpecker from Sabinas. Cory (1919:494) listed D. s- symplectus
from Pabinas (=:Sabinas?).

**Dcndrocopos scalaris giraudi (Stone). — Specimen examined: one, S
34623, from 12 mi. N, 10 mi. E Parras, 3850 ft., July 12, 1953, weight, 35
gms.

Ridgway (1914:259) reported that D. s. giraudi (then called Dryohates
.scalaris bairdi) has been recorded from La Ventura, Carneros, and Jaral.
Oberholser (1912:159) indicated that the subspecies giraudi occurs north
to Jaral in southern Coahuila, east to La Ventura in southeastern Coahuila,
and south through central Mexico. The present specimen is darker vcntrally

BiEDS From Coahuila, Mexico 473

than specimens of either D. s. cactophilus or D. s. stjmplectus, and on its
upper surface the black bars are wider than the white. The specimen is in
worn plumage, but nevertheless suggests intergradation with D. s. stjmplectus.

Platypsaris aglaiae albiventris (Lawrence). — The Rose-throated Becard, if
it occurs in Coahuila at all, is rare in the State. Without giving any spe-
cific locality, Sclater (1857:74) described Pachtjrhamphus aglaiae from Coa-
huila. Hellmayr (1929:202) considered Sclater's record as representing
Platypsaris aglaiae albiventris ( Lawrence ) .

*Tyrannus vociferans vociferans Swainson. — Specimens examined: total 2:
$ 32064 from 7 mi. S, 4 mi. E Bella Union, 7200 ft., June 24, 1952; and
9 31650 from the north foot of Sierra Guadalupe (=10 mi. S, 5 mi. W
General Cepeda), 6400 ft., April 15, 1953.

The small number of records of Cassin's Kingbird in Coahuila is surprising,
for I would expect the species in most areas of the State between 6000 and
7000 feet. All Coahuilan records are from the southeastern part of the State.
Burleigh and Lowery (1942:193) found it "on infrequent occasions in the
arid country near Saltillo." Miller, Friedmann, Giiscom, and Moore (1957:71)
reported that T. v. vociferans nested in Coahuila on July 7. The size of the
testes ( 15x7 mm. ) of our specimen from near Bella Union also indicates
breeding.

**Muscivora forficata (Gmelin). — Specimen examined: one, 9 32063, from
12 mi. N, 12 mi. W Jimenez, 850 ft., June 19, 1952.

The status of the Scissor-tailed Flycatcher in Coahuila is uncertain. Al-
though the condition of the gonads of No. 32063 was not recorded by the
collector, the late date (June 19) on which is was obtained suggests that this
female was a resident in northeastern Coahuila. Findley saw a Scissor-tailed
Flycatcher 2 mi. S and 11 mi. E Nava, 810 feet, on June 15, 1952. Dicker-
man saw one 4 mi. N San Isidro on May 10, 1954. Miller, Friedmann, Gris-
com, and Moore (1957:69) recorded this flycatcher from Sabinas on April 12.
No other records of Muscivora forficata in Coahuila have come to my atten-
tion, but the species probably occurs in the State in migration.

*Myiarchus crinitus crinitus (Linnaeus). — Specimens examined: total 3:

9 32065 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 19, 1952; and $

32066 and 9 32067 from 2 mi. W Jimenez, 850 ft., June 20, 1952.

The Great Crested Flycatcher seems to be rare in Coahuila. Nos. 32065-

32067 are the first specimens that I know of from Coahuila. Bangs (1898:
179-180) said that the subspecies crinitus has an "enormous swollen bill"
(exposed culmen of male, 20 mm.; breadth of bill at middle of nostril, 10.3
mm.) whereas M. c. horeus has a "small slender bill" (exposed culmen of
male, 18.6 mm.; breadth of bill at middle of nostril, 8.6 mm.). No. 32066
has a large bill (exposed culmen, 21 mm.; breadth of bill at middle of nostril,

10 mm.). Bangs {loc. cit.) did not give any measurements for females of
M. c. crinitus or horeus. Nevertheless, I suspect that Nos. 32065 and 32067
represent crinitus (No. 32065: exposed culmen, 19 mm.; breadth of bill at
middle of nostril, 9 mm.; No. 32067: exposed culmen, 20 mm.; breadth of
bill at middle of nostril, 9.5 mm.). The size of the testes (9 mm. long) of
No. 32066 and the dates (June 19 and 20) on which the specimens were
collected indicate breeding by this species in the State,

474 University of Kansas Publs., Mus. Nat. Hist.

*\Iyiarchus tyrannulus cooperi Baird. — Specimens examined: total 2: $
32068 and $ 32069 from 2 mi. S, 3 mi. E San Juan de Sabinas, 1160 ft.,
June 22, 1952, measurements: wing, 102, 97 mm.; tail, 93, 90 mm.

Wied's Crested Flycatcher is not common in Coahuila. Eastern Coahuila
represents the eastern limit of the range of M. t. cooperi. Measurements of
our specimens agree well with the average measurements of typical M. t.
cooperi. According to Ridgway (1907:621), M. t. cooperi (then called
Mtjiarchus mexicanu.i mexicanus) has been recorded from Sabinas. The size
of the testes (14x7 mm.) of No. 32068 and the date (June 22) on which the
specimens were collected indicate breeding by this species in the State.

*Myiarchus cinerascens cinerascens (Lawrence). — Specimens examined:
total 7: 9 31045 from 15 mi. SE Boquillas, 1500 ft., March 16, 1952; $
32070 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 19, 1952; 9 32071
from 2 mi. W Jimenez, 850 ft., June 20, 1952; $ 35413 from 6 mi. N, 2 mi.
W Castillon, 3750 ft., June 30, 1953; S 32072 from 9 mi. S, 11 mi. E Sabinas,
June 14, 1952; 9 31584 from 17 mi. S Ocampo, 5300 ft., April 7, 1954; and
6 31673 from the west foot of Pico de Jimulco, 5000 ft., April 4, 1953.

The Ash-throated Flycatcher is common in Coahuila. All specimens
examined by me from there are typical of M. c. cinerascens. Ridgway
(1907:626) hsted M. c. cinerascens from Monclovia ( r^Monclova? ) and
Sabinas. Miller (1955a:166) found the species "only in the oak belt at
5000 feet, where it was common." Burleigh and Lowery (1942:193) re-
corded M. c. cinerascens from "near Saltillo." Dickerman obtained No.
31584 in a mesquite-grassland-shrubby area. Miller, Friedmann, Griscom,
and Moore (1957:78) stated that M. c. cinerascens breeds in Coahuila.

Sayornis phoebe (Latham). — Miller, Friedmann, Griscom, and Moore ( 1957:
66) recorded the Eastern Phoebe from Coahuila on November 4-14.

*Sayornis nigricans semiatra (Vigors). — Specimens examined: total 2: $
31046 from 1 mi. N Boquillas, 700 ft., March 7, 1952; and 9 31047 from
Fortin (=33 mi. N, 8 mi. W San Geronimo), 3300 ft., March 27, 1952.

Of the Black Phoebe, the two subspecies semiatra and nigricans intergrade
in Coahuila. Typical representatives of S. n. semiatra are present in northern
Coahuila. The under tail coverts of Nos. 31046-31047 are immaculate and
white. Miller (1955:167) noted the two specimens collected from the Sierra
del Carmen to have narrow dark shaft streaks on the under tail coverts. He
{loc. cit.) remarked also that "the marking of the under tail coverts may
indicate a beginning of a gradient in increased darkening of these feathers
toward S. n. nigricans in southern Coahuila."

*Sayornis nigricans nigricans (Swainson). — Burleigh and Lowery (1942:
193) collected several specimens of S. n. nigricans "on the outskirts of
Saltillo" and saw a pair at the Chorro del Agua on April 19.

*Sayornis saya saya (Bonaparte). — Specimens examined: total 4: 9 31049
from the Rio Grande (=17 mi. S Dryden, Terrell Co., Texas, in Coahuila),
600 ft., March 18, 1952; sex ? 31048 from 10 mi. S, 5 mi. E Boquillas, 1500
ft, March 4, 1952; $ 31050 from Fortin (=33 mi. N, 8 mi. W San Gcr6nimo),
3300 ft., March 29, 1952; and 9 32059 from 7 mi. S, 4 mi. E Bella Union,
7200 ft., June 25, 1952.

Say's Phoebe occurs commonly in Coahuila. Miller (1955a:167) ob-
tained a female with an active brood patch in Boquillas Canyon of the Sierra

Birds From Coahuila, Mexico 475

del Carmen, and remarked that Marsh took a juvenile on September 2 at
El Rincon. Sutton and Burleigh (1939a: 33) saw this phoebe several times
in southern Coahuila and obtained a male "near San Pedro" on January 29.
Burleigh and Lowery (1942:193) recorded this species as "a common breed-
ing bird both on Diamante Pass and on the arid plateau around Saltillo."
No. 32059 was a juvenile.

Empidonax traillii brewsteri Oberholser. — Amadou and Phillips (1947:578)
took a Traill Flycatcher of the subspecies brewsteri at Las Delicias on August
11.

Empidonax minimus (Baird and Baird). — Specimen examined: one, $
31470, from Sierra del Pino (=5 mi. S, 3 mi. W Acebuches), 6200 ft., May

13, 1954.

Amadou and Phillips (1947:578) obtained two Least Flycatchers at Las
Delicias on August 12. Dickennan took No. 31470 in pine-oak vegetation.

Empidonax hammondii (Xantus). — Specimen examined: one, sex ? 31657,
from the north slope of Sierra Guadalupe (=11 mi. S, 7 mi. W General
Cepeda), 7800 ft., April 20, 1953.

No. 31657 is similar to E. wrightii (Wright's Flycatcher); however,
the outmost (tenth) primary is equal to or shghtly larger than the fifth
primary. Yet, the underparts of No. 31657 are darker and more uniform
in coloration than those of typical representatives of E. wrightii. Miller,
Friedmann, Griscom, and Moore (1957:88) stated that Hammond's Flycatcher
is "transient" in Coahuila. Burleigh and Lowery (1942:193-194) reported
that E. ham.mondii was the most prevalent of the small flycatchers in south-
eastern Coahuila. They (lac. cit.) obtained specimens of this flycatcher from
the Chorro del Agua and Diamante Pass. Miller (1955a: 167) characterized
E. hammondii as a common migrant, "chiefly in stands of low oaks in the
pine-oak belt but also occasionally in the desert scrub" of the Sierra del
Carmen.

Empidonax wrightii Baird. — ^Amadon and Phillips (1947:578) reported one
Wright's Flycatcher from Las Delicias. Miller (1955a: 167) stated that this
species was a common migrant and occurred chiefly in the lower oak belt
and in the desert scrub. Sutton and Burleigh (1939a: 34) obtained speci-
mens of E. wrightii from San Pedro on January 29.

Empidonax griseus Brewster. — Sutton and Burleigh (1939a: 34) noted that
the Gray Flycatcher was common "in the San Pedro region" and collected
two at San Pedro on January 29. Burleigh and Lowery (1942:194) col-
lected specimens "near the Chorro del Agua, at Saltillo, and ... in the
open desert some twenty miles west of Saltillo."

**Empidonax affinis trepidus Nelson. — Specimen examined: one, $ 32750,
from 13 mi. E San Antonio de las Alazanas, 9950 ft., July 6, 1955.

Ridgway (1907:576) previously reported the httle known and poorly de-
fined Pine Flycatcher from Sierra Guadalupe. Because of its small size (wing,
75 mm.; tail, 65 mm.). No. 32750 is referable to E. a. trepidus. No indica-
tion of breeding of the subspecies trepidus exists for Coahuila. Nevertheless,
the date (July 6) on which No. 32750 was obtained suggests that this fly-
catcher may breed in southeastern Coahuila.

3—7905

476 Unimehsity of Kansas Publs., Mus. Nat. Hist.

*Empidonax difficilis hellmatfri Brodkorb. — Specimen examined: one, $
31469, from Sierra del Pino (=5 mi. S, 3 mi. W Acebuches), May 13, 1954,
measurements: wing, 71 mm.; tail, 65 mm.; culmen, 11.5 mm.; tarsus, 17.0
mm.

Miller (1955a: 167) reported that the Western Flycatcher breeds in the
Sierra del Carmen, from 6800 to 7500 feet. According to Miller, Friedmann,
Griscom, and Moore (1957:91), this subspecies has been recorded from Sierra
Guadalupe. No. 31469 closely resembles, especially in measurements, the
specimens of the Western Flycatcher from the Chisos Mountains of Texas
as reported by Brodkorb (1935:2).

Empidonax difficilis subsp. — Burleigh and Lowery (1942:194) obtained a
specimen of £. difficilis "near the summit of Diamante Pass" that they tenta-
tively identified as E. d. sahini. However, they considered the specimen as
possibly E. d. immcmoratus or E. d. occidentalis.

Localities listed by Miller, Friedmann, Griscom, and Moore (1957:92) of
E. d. salvini and E. d. immemoratus are south of twenty-three degrees north
latitude whereas the range of E. d. occidentalis includes parts of Nuevo Leon
north of tvvcnty-three degrees north latitude (occidentalis intergrades with
hellmayri at Cerro Potosi, Nuevo Leon, [Miller, Friedmann, Griscom, and
Moore, loc. cit.]). Thus, the specimen from Diamante Pass probably is either
E. d. hellmaijri or E. d. occidentalis.

**Empidonax fulvifrons pygmaeus Coues. — Miller, Friedmann, Griscom, and
Moore (1957:95) listed this subspecies of the Buff-breasted Flycatcher from
the Sierra Guadalupe on April 21. This record represents the eastern limit
ol the range of E. f. pygmaeus in northeastern Me.xico.

'^■*Contopus pertinax pertinax Cabanis and Heine. — Burleigh and Lowery
(1942:194) obtained one female Coues' Flycatcher "in a small gully just below
the summit of Diamante Pass."

Contopus virens (Linnaeus). — I judge from the paucity of records in the
literature that the Eastern Wood Pewee is uncommon in Coahuila. Burleigh
and Lowery (1942:194) obtained two males at tlie Chorro del Agua, and
remarked also that "it is possible that as far west as Saltillo, this species is
a rather uncommon transient." Ridgway (1907:519) listed Contopus virens
from Sabinas.

*Contopus sordidulus veliei Coues. — Specimens examined: total 2: S
31467 from Sierra del Pino (=5 mi. S, 3 mi. W Acebuches), May 14 1954;
and $ 31653 from the north foot of Sierra Guadalupe (=10 mi S 5 mi W
General Cepeda), 6400 ft., April 19, 1953.

Specimen No. 31467 of the Western Wood Pewee was obtained in pine
and oak vegetation by Dickerman. Ridgway (1907:523) reported Contopus
sordidulus veliei, under the name Myiochancs richardsonii rcchardsonii, from
Sierra Encamacion. Amadon and Phillips (1947:578) obtained a specimen
of C. sordidulus from Las Delicias. Miller, Friedmann, Griscom, and Moore
(1957:83) remarked that the Western Wood Pewee breeds in the State.

Nuttnllnrnis hnrealis (Swainson). — Several records of the Olive-sided Fly-
catcher from Coahuila are present in the literature. Miller (1955a: 167) re-
ported it as a migrant in the desert at the base of the Sierra del Carmen on

Birds From Coahuila, Mexico 477

April 24. Miller, Friedmann, Griscom, and Moore (1957:82) reported this
species from Ocampo on June 16. Amadon and Phillips (1947:578) obtained
one at Las Delicias on August 16. Burleigh and Lowery (1942:194) collected
two males "near the summit of Diamante Pass" on April 14.

*Pyrocephalus rubinus mexicanus Sclater. — Specimens examined: total 3:
$ 32060 and 9 32061 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 19,
1952; and sex ? 32062 from 2 mi. W Jimenez, 850 ft., June 20, 1952.

The Vermilion Flycatcher is common in Coahuila. Except in the north-
western part of tlie State, the subspecies mexicanus is present throughout Coa-
huila. The size of No. 32060 (wing, 80 mm.; tail, 62 mm.) suggests that
the specimen is an intergrade between P. r. flammeus and mexicanus. P. r.
flammeus and mexicanus seem to intergrade in northern Coahuila.

Burleigh and Lowery (1942:195) found mexicanus to be "quite plentiful on
the plains surrounding Saltillo." Sutton and Burleigh (1939a:33) noted the
Vermilion Flycatcher "near San Pedro." Cory and Hellmayr (1927:92) listed
P. r. mexicanus from Sabinas. The size of the testes (6x4 mm. long) of No.
32060 and the dates (June 19 and 20) on which our specimens were collected
indicate breeding.

Ptjrocephalus rubinus flammeus van Rossem. — This subspecies of Vermilion
Flycatcher occupies the northwestern section of Coahuila. Marsh and Steven-
son (1938:287) took a specimen of P. r. flammeus "near Santo Domingo, east
of the Del Carmens. . . ." Miller (1955a: 167) re-examined this specimen
and stated that "its affinity with the northwest race of the species seems correct
in tenns of the characters outlined by van Rossem (1934:353)."

Eremophila alpestris enthymia ( Oberholser ) . — Miller, Friedmann, Griscom,
and Moore (1957:105) recorded this subspecies of Homed Lark wintering 4
mi. S Hipolito (November 2 to February 24). They reported also that E. a.
enthymia breeds in Coahuila (4 mi. S Hipolito ?).

*Eremophilia alpestris diaphora (Oberholser). — Specimens examined: total
9: $ $ 32073-32078 and 9 32079 from 7 mi. S, 4 mi. E Bella Union, 7200
ft., June 25, 1952; and $ $ 31605-31606 from 14 mi. W San Antonio de las
Alazanas, 6500 ft., January 9, 1954.

Burleigh and Lowery (1942:195) wrote that this subspecies of Horned
Lark was uncommon "about Saltillo," but "fairly common" in Diamante Val-
ley, at about 7000 feet. Oberholser (1902:863) recorded the subspecies
aphrasta from La Ventura. Ridgway (1907:326) listed Otocoris alpestris
aphrasta Oberholser from Saltillo and La Ventura. However, Miller, Fried-
mann, Griscom, and Moore (1957:105) indicated that E. a. diaphora rather
than aphrasta was recorded from Saltillo and La Ventura. Sutton and Bur-
leigh (1939a: 35) remarked that "specimens collected near Ramos Arizpe [in
southeastern Coahuila]" proved to be E. a. aphrasta. However, this record
might be questioned, as Miller, Friedmann, Griscom, and Moore (op. cit.)
indicated, because diaphora, rather than aphrasta, seems to occupy the south-
eastern sector of CoahuHa. Possibly typical representatives of aphrasta and/or
intergrades between aphrasta and diaphora are present in western Coahuila;
however, no records of E. a. aphrasta exist from western Coahuila.

The sizes of Nos. 32073-32077 and 31605-31606 (wing, 98.0-101.5 mm.,
averaging 99.7 mm.), the bright yellow throat, and the vinaceous color of the

478 Unixtrsity of Kansas Publs., Mus. Nat. Hist.

hindneck characterize clearly the subspecies diaphora. The sizes of the testes
(9x6 mm.; 8x5 mm.; 10.\6 mm.; 8.x4 mm.; 11x6 mm.) of Nos. 32073-32077,
the size of the largest ovum (6.5 mm.) of No. 32079, and the juvenile (32078)
are evidence of breeding of E. a. diapJwra in Coahuila.

*Tachtjcineta thalassina thalassina (Swainson). — Specimen* examined: total
3: S 31471, $ 31473, and 9 31472 from Sierra del Pino (=5 mi. S, 3 mi.
W Acebuchcs), May 15, 1954, measurements: wings, 125, 118, 108 mm.;
tails, 56, 54, 46 mm.

The two subspecies of the Violet-green Swallow, thahs^na and lepida,
intergrade in Coahuila. Specimens from diflFerent localities in the State rep-
resent various stages of intergradation between the two subspecies; generally
those from northern Coahuila seem to be closer to T. t. lepida, and those from
southern Coahuila are closer to T. t. thalassina. Nos. 31471-31473 are inter-
grades between T. t. thalassina and T. t. lepida; in size the three resemble
T. t. thalassina, but in green, rather than purple, backs and scapulars resemble
T. t. lepida. The nunps of Nos. 31471-31473 show some purple with the
green, but are nearer thalassina in this character.

Gonadal sizes ( testes 10x7, 10x8 mm., one ovum 6 mm. ) indicate that
the Violet-green Swallow breeds in the Sierra del Pino.

*Tachycineta thalassina lepida Mearns. — Miller (1955a: 167) r<:>ported that
Violet-green Swallows taken in Boquillas Canyon of the Sierra del Carmen
are intermediate between T. t. lepida and thalassina; however, he referred
his sample to lepida on the basis of short wing.

Iridoprocne bicolor (Vieillot). — Miller, Fricdmann, Griscom, and Moore
(1957:114) reported the Tree Swallow from Ilipohto on February 22,

Stelgidopteryx ruficoUis psammochroiis Griscom. — Miller, Friedmann, Gris-
com, and Moore (1957:111) reported this subspecies of the Rough-winged
Swallow from Saltillo.

*Hirundo rustica erythrogaster Boddaert. — Burleigh and Lowery (1942:195)
stated that the Bam Swallow was the most abundant swallow "about Sal-
tillo." Because these swallows occupied houses about Saltillo and neighboring
villages, Burleigh and Lowery (loc. cit.) concluded that the species nests
commonly in the Saltillo area. Findley saw Barn Swallows 2 mi. S and 11
mi. E Nava, 810 feet, on June 15, 1952. Dickerman observed them at San
Marcos ( =20 mi. S Cuatro Cienegas ) on May 4, 1954.

*Pctrochelidon pyrrhonota minima van Rossem and Hachisuka. — Specimens
examined: total 2: $ $ 31585-31586 from 14 mi. W San Antonio de las
Alazanas, April 9, 1954.

Burleigh and Lowery (1942:196) noted the Cliff Swallow "in small num-
bers about Saltillo and the nearby villages" where the species "showed evidence
of beginning to nest in many of the liouses. . . ." They ( loc. cit. ) found
Cliff Swallows also at the Chorro del Agua and Diamante Valley. According
to Dickennan, Nos. 31585-31586 were from a colony, members of which were
collecting mud at a stock tank.

**Petrochelidon fulva pallida Nelson. — The Cave Swallow seems to be un-
common in eastern Coahuila. Selander and Baker (1957:345) hst Saltillo,

Birds From Coahuila, Mexico 479

Sabinas, and Monclova as the three known locahties for this swallow in the
State.

*Progne subis subis (Linnaeus). — Miller, Friedmann, Griscom, and Moore
(1957:107) reported, presumably on the basis of a pubHshed record not found
by me, that the Purple Martin breeds in Coahuila.

**Progne chalybea chaltjbea (Gmelin). — The only record of the Gray-
breasted Martin in Coahuila was given by Ridgway (1904:42) when he hsted
P. c. chalybea from Sabinas.

**Cyanocitta stelleri tnacrolopha Baird. — Specimens examined: total 2:
S 32788 and 5 32787 from 13 mi. E San Antonio de las Alazanas, 9950 ft.,
July 6, 1955.

From the paucity of records in the literature, I judge that Steller's
Jay is uncommon in Coahuila. Nos. 32787-32788 seemingly represent the
first records of this species in the State, and are referred to the subspecies
macrolopha on the basis of relatively long (150, 151 mm.) wing, near the
maximum for stelleri from Mexico. The date (July 6) of collection suggests
that these birds were resident in southeastern Coahuila.

'**Aphelocoma coerulescens cyanotis Ridgway. — The Scrub Jay is common
in southeastern Coahuila. Burleigh and Lowery (1942:196) noted this
species in small numbers at the summit of Diamante Pass, daily on the lower
slopes of the mountains, and a few birds "on the outskirts of Saltillo, where
they were probably nesting. . . ." Sutton and Burleigh (1939a: 35) also
noted several flocks at Diamante Pass. Miller, Friedmann, Griscom, and
Moore (1957:123) reported A. c. cyanotis from El Diamante. Ridgway
(1904:335) hsted this subspecies of the Scrub Jay (then called Aphelocoma
cyanotis) from Carneros, Sierra Encamacion, and Sierra Guadalupe.

* Aphelocoma ultramarina couchii (Baird). — Specimens examined: total
14: 5 31051 from Sierra de la Encantada (=38 mi. S, 23 mi. E Boquillas),
4400 ft., March 15, 1952; $ $ 29425-29426 and $ 9 29427-29428 from
Club Sierra del Carmen {=2 mi. N, 6 mi. W Piedra Blanca), 4950 ft., April
8, 1950; sex ? 31052 (skeleton only) from 4 mi. W Hacienda La Mariposa,
2300 ft., March 25, 1952; 5 9 31635-31636 from Canyon de Parazos in
the Sierra de la Parazos Azula (=9 mi. E Hermanas), 2100 ft., December
7, 1953; 9 32082 from 4 mi. N, 21 mi. W Cuatro Cienegas, 6200 ft., July 3,
1952; 9 31053 (skeleton only) from 26 mi. W Santa Teresa, 7050 ft., April
5, 1952; $ 32081 from 7 mi. S, 4 mi. E Bella Union, 7200 ft., June 25, 1952;
9 33173 (skeleton only) from 13 mi. E San Antonio de las Alazanas, 9950
ft., July 6, 1955; S 31607 from 13 mi. E, 3 mi. S San Antonio de las Alazanas,
8900 ft., January 11, 1954; and $ 31654 from Sierra Guadalupe (=10 mi. S,
5 mi. W General Cepeda), 7000 ft., April 13, 1953.

The Mexican Jay is common in Coahuila. This species occupies various
habitats in the State and has been collected at stations ranging from 2100
to 9950 feet.

Miller (1955a: 167) stated that the Mexican Jay was the most abundant
species of bird in the Sierra del Carmen. Burleigh and Lowery (1942:196)
remarked that specimens of A. u. couchii were obtained "in the area surround-
ing the summit of Diamante Pass." At no time did Burleigh and Lowery
(loc. cit.) see this species below 6500 feet. The Mexican Jay is restricted to
the higher altitudes in southern Coahuila but is more widespread in northern

480 Unh'ersity of Kansas Publs., Mus. Nat. Hist.

Coahuila. Dickerman saw Mexican Jays in the Sierra del Pino on May 12,
1954; 16 mi. E and 18 mi. N Ocampo on May 7, 1954; in the Sierra de la
Madera on December 13, 1953; and 20 mi. S Ocampo on April 4, 1954.

Ridgway (1904:340) remarked tliat Aphelocoma sieberii potosina Nelson
[=A. u. couchii (Baird)] has been recorded in southern Coahuila, at Cameros.
Miller, Friedmann, Griscom, and Moore (1957:124) recorded A. u. couchii
from Cameros, Nuevo Leon. I suspect (Ridgway, 1904:340 and Goldman,
1951: map opposite p. 34) that the locality given by Miller, Friedmann,
Griscom, and Moore (op. cit.) should be Cameros, Coahuila, rather than
Cameros, Nuevo Leon.

Specimens (31051, 29425-29428, 31635-31636, 32081-32082, 31607, and
31654) of the Mexican Jay increase in size from northern Coahuila to southern
Coahuila. The average length of the wings of Nos. 29425-29426 from Club
Sierra del Carmen is 152 mm. whereas the average length of the wings of
No. 32081 from 7 mi. S and 4 mi. E Bella Union, of No. 31607 from 13 mi.
E and 3 mi. S San Antonio de las Alazanas, and of No. 31654 from Sierra
Guadalupe is 164 mm.

Miller (1955a: 169) indicated that the Mexican Jay breeds in the Sierra
del Carmen. Burleigh and Lowery (1942:196) remarked that the species
breeds at Diamante Pass. The sizes of the testes (12, 11 mm.) of Nos.
29425-29426, the size of the ovum (8 mm.) of No. 29428, and the short
tail (126 mm.) of the immature female from 4 mi. N and 21 mi. W Cuatro
Cienegas are also evidence of breeding by this species in the State.

*Corvus corax sinuatus Wagler. — The Common Raven seems to occur in
low density in Coahuila. Miller (1955a: 168) saw ravens in the pine-oak
and cliff areas of the Sierra del Carmen, and took a breeding female at the
head of Corte Madera Canyon, 7500 feet. Burleigh and Lowery (1942:196)
found this species to be a bird of the higher slopes of the mountains although
not uncommon in the broad open valley south of Diamante Pass. Sutton
and Burleigh (1939a:35) took a male at Santa Rosa.

*Corvus cryptoleucus Couch. — Specimens examined: total 4: $ 32080
from 2 mi. W Jimenez, 850 ft., June 20, 1952; 9 35404 (skeleton only) from
4 mi. N San Isidro (=16 mi. N Ocampo), May 10, 1954; and S 6 31474-
31475 from R. de Almendarez (=53 mi. NW Ocampo), May 11, 1954.

The White-necked Raven occurs throughout Coahuila. Sutton and Bur-
leigh (1939a:36) observed this species "in the vicinity of Saltillo," but not
farther west. Burleigh and Lowery (1942:197) noted C. cryptoleucus fre-
quently "on the arid plateau around Saltillo" and obtained a specimen "in
the high fertile valley south of Diamante Pass." Burleigh and Lowery (loc.
cit.) remarked that the White-necked Raven avoids the mountain slopes;
7000 feet was the uppermost limit of occurrence. The sizes of the testes ( 32080,
20 mm.; 31474: left, 12x20 mm., right, 10x16 mm.; and 31475: left, 10x16
mm., right, 10x14 mm.) of the specimens that I have examined and the dates
(May 11, June 20) on which they were collected indicate breeding by the
White-necked Raven in Coahuila.

*Farus sclateri eidoa (Peters). — Specimens examined: total 4: 9 32083
from 7 mi. S, 4 mi. E Bella Uni6n, 7200 ft., June 25, 1952; $ 31609 from 2
mi. E Mesa de Tablas, 9000 ft., January 15, 1954; $ 31656 and 9 31655
from the north slope of Sierra Guadalupe (^H rni. S, 7 mi. W General
Cepeda), 7800 ft., April 20, 1953.

Birds From Coahuila, Mexico 481

The Mexican Chickadee is common in southeastern Coahuila. Miller,
Friedmann, Griscom, and Moore (1957:133) stated that P. s. eidos and P. s.
sdateri intergrade in southern Coahuila. The specimens that I have exam-
ined also show signs of intergradation, but are closer to P. s. eidos.

Sutton and Burleigh (1939a: 36) reported taking specimens of P. s. eidos
at Diamante Pass where they saw several individuals. Burleigh and Lowery
(1942:197) also observed these chickadees "in the pine woods about the
summit of Diamante Pass. . . ." The size of the testes (6x5 mm.) of
No. 31656 and the fact that No. 31655 was incubating eggs at the time it
was obtained are evidence of breeding by this chickadee in the State.

**Parus sdateri sdateri Kleinschmidt. — Miller, Friedmann, Griscom, and
Moore (1957:133) stated that a specimen (or specimens ?) of P. s. sdateri
which showed evidence of intergradation with P. s. eidos was obtained at El
Diamante.

**Parus atricristatus dysleptus Van Tyne. — Spedmen examined: one, $
31054 from 37 mi. S, 21 mi. E Boquillas, 4100 ft., March 13, 1952.

Two subspecies of the Black-crested Titmouse are present in Coahuila.
P. a. dysleptus occurs in northwestern Coahuila. Miller (1955a: 168) stated
that the Black-crested Titmouse, identified as dysleptus, was the only repre-
sentative of the genus Parus in the Sierra del Carmen. The weak extension
of black onto the nape in No. 31054 suggests intergradation between P. a.
dysleptus and P. a. atricristatus; the latter lacks the black nape of dysleptus.

*Parus atricristatus atricristatus Cassin. — Spedmens examined: total 4:
9 32084 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 19, 1952; sex ? 32085
from 2 mi. W Jimenez, 850 ft., June 20, 1952; 9 31055 (skeleton only) from
Fortin (=33 mi. N, 1 mi. E San Geronimo), 3300 ft., on March 29, 1952;
and 9 35399 (skeleton only) from 3.5 mi. W, 22 mi. S Ocampo, December
15, 1953, weight, 16 gms.

Typical P. a. atricristatus occurs in Coahuila in the southeastern sector
(Dixon, 1955:184). Black-crested Titmice intermediate between atricristatus
and dysleptus were hsted from Cuidad Miizquiz and Sabinas by Dixon (loc.
cit. -.189), as dysleptus but were shown on his map (loc. cit. :184) as
atricristatus. Our Nos, 32084 and 32085 (wdng, 71, 71, tail, 63, 64 mm.) are
small and fall in the upper range of size for atricristatus to which the specimens
are here referred.

^Auriparus flaviceps ornatus (Lawrence). — Specimen examined: one, $
31056, from Sierra de la Encantada (=38 mi. S, 23 mi. E Boquillas), 4400
ft., March 15, 1952.

The Verdin occurs up to about 5000 feet in Coahuila. Miller (1955a: 168)
reported that "this desert species followed the catclaw scrub up the washes
to about 4800 feet, the Umit of such habitat" in the Sierra del Carmen and
also gave evidence of breeding by the Verdin in Coahuila. Amadon and Phillips
(1947:578) reported a nest of Auriparus flaviceps at Las Delicias. Burleigh
and Lowery (1942:197) obtained a single specimen "in an arroyo a few miles
east of Saltillo" and a single specimen "about twenty miles west" of Saltillo.
Ridgway (1904:421) reported A. f. ornatus [he referred to it as A. f. flaviceps]
at Monclova. Hellmayr (1934:88) listed A. f. ornatus from Jaral. Findley
saw a Verdin 2 mi. S and 3 mi. E San Juan de Sabinas on June 22, 1952.

The size (wing, 50 mm.; tail, 43 mm.) of No. 31056 is small for typical

482 Uixn^RsiTY of Kansas Publs., Mus. Nat. Hist.

representatives of A. /. ornatus. The yellow of the head of No. 31056 is
darker than that of the other subspecies of the Verdin, and I have accordingly
allocated the specimen to A. f. ornatus.

*Psahriporus melanotis lloydi Sennett. — Specimens examined: total 5:
$ 3105S and 9 31057 from 37 mi. S, 21 mi. E Boquillas, 4100 ft., March 13,
1952; $ 31060 and 9 31059 from Sierra de la Encantada (=38 mi. S, 23 mi.
E Boquillas), 4400 ft., March 15, 1952; and $ 35407 (skeleton only) from
Sierra del Pino (=5 mi. W, 3 mi. S Acebuches), 6200 ft., May 14, 1954.

The Black-eared Bushtit is common in Coahuila. Typical representatives
of P. m. Uotjdi in the northern part of the State range from 4100 feet to
8000 feet. Miller ( 1955a:168-169) reported P. m. Uotjdi breeding in the
Sierra del Cannen. Marsh and Stevenson (1938:287) obtained a male in
the Sierra del Carmen at El Jardin. The size of the testes (3.5 mm.) of
No. 31058 suggests breeding by the Black-eared Bushtit 37 mi. S and 21 mi.
E Boquillas.

**Psaltriparus melanotis iulus Jouy. — Specimens examined: total 2: $
31659 from the north slope of Sierra Guadalupe (=10 mi. S, 7 mi. W General
Cepeda), 7000 ft., April 20, 1953; and 9 31658 from tlie north slope of
Sierra Guadalupe (=10 mi. S, 5 mi. W General Cepeda), 6700 ft., April
15, 1953.

Typical representatives of this Black-eared Bushtit are present in the
southeastern sector of Coahuila. The backs of Nos. 31658-31659 differ
slightly in color from the backs of typical representatives of lloydi. I suspect
that the specimens from the Sierra Guadalupe are intergrades between lloydi
and iulus.

Burleigh and Lowery (1942:197) took two males and two females of
Psaltriparus melanotis iulus (they called their specimens Psaltriparus minimus
iulus) at Saltillo and stated that their specimens tended to approach lloydi
rather than being typical iulus. Sutton and Burleigh (1939a:36) recorded
P. m. iulus only at Diamante Pass where they took two specimens.

*Sitta carolinensis nelsoni Meams. — Miller (1955a: 169) reported that this
subspecies of the White-breasted Nuthatch breeds and was common in the
oaks and open conifers from 6500 to 8000 feet in the Sierra del Cannen and
stated also that the populations of the White-breasted Nuthatch in "the Chisos
Mountains [of Te.xas] and the Sierra del Carmen seem best regarded as a stage
in the clinc of which nelsoni and mexicana arc end points, although falling
closer to nelsoni."

*Sitta carolinensis mexicana Nelson and Palmer. — Specimens examined:
total 2: $ 31669 and 9 31670 from the Canon d. Meco, Sierra Guadalupe
(=10 mi. S General Cepeda), 6500 ft., April 23, 1953, weights, 17, 18 gms.

Typical populations of this White-breasted Nuthatch occur in southern
Coahuila. Ridgway (1904:449) listed S. c. mexicana from Sierra Guadalupe
in southern Coahuila. The underparts of our specimens are darker than in
nelsoni, and their bills (culmen, 15, 13 mm.) are shorter than the average
( $, 19.8 mm.; 9, 18.6 mm. [Ridgway, 1904:447]) in nelsoni. The large size
of the testes (5.\3 mm.) of No. 31669 suggests breeding by S. c. mexicana in
Sierra Guadalupe.

*Sitta pygmaea melanotis van Rossem. — In Coahuila the Pigmy Nuthatch
seems to be locally common. Miller (1955a:169) reported it so between 7500

Birds From Coahuila, Mexico 483

and 8000 feet in pine-oak on the mesa tops and in the heads of canyons of
the Sierra del Carmen and noted that it breeds there. Hardy saw the Pigmy
Nuthatch 13 mi. E San Antonio de las Alazanas on July 6, 1955.

Certhia familiaris americana Bonaparte. — Specimens examined: total 2:
9 31612 from the base of Don Martin Dam, November 27, 1953, skull par-
tially unossified; and ? 31587 from 20 mi. S Ocampo, 6500 ft., April 5, 1954,
weight, 7 gms.

This subspecies of the Brown Creeper can be considered a sparse winter
visitant to Coahuila. Van Hoose (1955:302) reported that Nos. 31612 and
31587 constitute the southernmost records of C. /. americana and represent
the first records of occurrence of americana in Mexico.

Certhia familiaris montana Ridgway. — Miller (1955a: 169) reported this
subspecies of the Brown Creeper, which he assumed to be a winter visitant
or a migrant, in the Sierra del Carmen. He (loc. cit.) remarked also that the
higher conifers would seem to constitute favorable habitat for nesting by the
Brown Creeper, but did not find any evidence of a breeding population of
creepers in the Sierra del Carmen. Miller, Friedmann, Griscom, and Moore
(1957:143) reported C. /. montana from San Lazaro Mountain on November 9.

**Certhia familiaris albescens Berlepsch. — Specimens examined: total 3:
sex ? 32805 from 13 mi. E San Antonio de las Alazanas, July 7, 1955; $ $
31610-31611 from 3 mi. S, 13 mi. E San Antonio de las Alazanas, 8900 ft.,
January 12, 1954.

Miller, Friedmann, Griscom, and Moore (1957:143) reported C. /. albe-
scens from "southern Coahuila." Nos. 31610-31611 and 32805 represent the
only other records of this subspecies from the State. The date (July 7) on
which No. 32805 was obtained suggests that this bird was a resident 13 mi. E
San Antonio de las Alazanas.

Troglodytes aedon parkmanii Audubon. — Specimen examined: one, sex ?
29556, from 1.5 mi, N Parras, 5500 ft., November 10, 1949, weight, 9.8 gms.

Burleigh and Lowery (1942:197) recorded the House Wren "in small
numbers about Saltillo where occasional birds, presumably migrants, were noted
in thickets or stretches of underbrush fringing cultivated fields." They ob-
tained a single male "on the outskirts of Saltillo." Hellmayr (1934:218)
listed T. a. parkmanii from Sabinas.

*Troglodytes brunneicollis cahooni Brewster. — Typical representatives of
this subspecies of the Brown-throated Wren occur in northern Coahuila. In
the Sierra del Carmen, Miller (1955a: 170) found T. h. cahooni that in no
way suggested compositus of the Sierra Madre Oriental. Burleigh and Lowery
(1942:198) recorded a cc^ooni-like specimen from Diamante Pass in southern
Coahuila.

**Troglodytes brunneicollis compositus Griscom. — Specimen examined: one,
$ 32819, from 13 mi. E San Antonio de las Alazanas, July 6, 1955.

The subspecies cahooni and compositus of the Brown-throated Wren seem
to intergrade in the southern part of the State. Although No. 32819 represents
the subspecies compositus, the somewhat whitish abdomen and the fairly
large spots of the lesser wing coverts suggest some relationship with cahooni.
In addition to the present record. Miller, Friedmann, Griscom, and Moore

484 University of Kansas Publs., Mus. Nat. Hist.

(1957:163) recorded T. b. compositus from southern Coahuila at Sierra
Guadalupe. The record of T. h. cahooni from Sierra Guadalupe (Ridgway,
1904:588) I suspect probably represents T. h. compositus or an intergrade
between compositus and cahooni. The date (July 6) on which No. 32819 was
obtained 13 mi. E San Antonio de las Alazanas suggests that this bird was
resident there.

*Thryomanes bewickii eremophilus Oberholser. — Specimens examined:
total 3: S 32088 from 2 mi. W Jimenez, 850 ft., June 20, 1952; 9 31061
from 4 mi. W Hacienda La Mariposa, 2300 ft., March 24 1952, weight, 10.8
gms.; and $ 31660 from the north foot of Sierra Guadalupe (=10 mi. S,
5 mi. W General Cepeda), 6500 ft., April 21, 1953, weight, 13 gms.

Bewick's Wren occurs commonly in Coahuila. Miller, Friedmann, Gris-
com, and Moore (1957:160) reported that, in Coahuila, T. b. eremophilus
"intergrades in the eastern and southern sections with T. b. cryptus and T. b.
murinus, respectively." The slightly darker coloration of No. 31660, sug-
gesting a resemblance to T. b. murinus, is the only evidence of intergradation
of T. b. murinus and eremophilus that I have found.

Miller (1955a: 170) stated that T. b. eremophilus was "common in the
piedmont area on yucca-dotted slopes and along the lower canyon walls in
growth of piiion, yucca, and cactus" in the Sierra del Carmen, and reported
breeding there. Burleigh and Lowery (1942:198) remarked that T. b. ere-
mophilus "proved without question to be the most widely distributed and
abundant wren" in the Saltillo region. The series that Burleigh and Lowery
{loc. cit.) assembled "proved to be uniform and clearly referable to" T. b.
eremophilus. Sutton and Burleigh (1939a: 36) noted Thryomanes bewickii
at San Pedro on January 29 and 30. Hardy saw a male T. b. eremophilus at
Parras on July 4, 1955. Ridgway (1904:557) listed T. b. eremophilus from
Saltillo in April and as breeding at Sabinas.

The sizes of the testes (8 mm.; 6x4 mm.) of Nos. 32088 and 31660,
respectively, suggest breeding 2 mi. W Jimenez and Sierra Guadalupe.

**Thryomanes bewickii cryptus Oberholser. — Miller, Friedmann, Griscom,
and Moore (1957:161) recorded T. b. cryptus from Saltillo.

*Thryothorus ludovicianus berlandieri Baird. — Specimens examined: total
3: $ 32086 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 19, 1952; sex ?
32087 from 2 mi. W Jimenez, June 20, 1952; and 9 31063 from 8 mi. N,
4 mi. W Muzquiz, 1800 ft., April 1, 1952, weight, 18.3 gms.

One subspecies of the Carolina Wren, berlandieri, occurs in Coahuila in
the northeastern section of the State. Ridgway (1904:547) recorded T. I.
berlandieri from Sabinas. The fact that No. 32086 was a juvenile suggests
that the Carolina Wren breeds 12 mi. N and 12 mi. W Jimenez.

*Catnpylorhynchus brunneicapillum couesi Sharpe. — Specimens examined:
total 6: $ 29429 from Caiion del Cochino (=16 mi. N, 21 mi. E Piedra
Blanca), 3200 ft., April 6, 1950; $ 31064 from 7 mi. S, 2 mi. E Boquillas, 800
ft, February 29, 1952, weight, 38.1 gms.; 9 31066 (skeleton only) from 10
mi. S, 5 mi. E Boquillas, 1500 ft., March 5, 1952; $ 31637 from La Gacha
(=La Concha), December 1, 1953, weight, 40 gms.; $ 31638 from 18 mi. S
Ocampo, December 16, 1953; and sex ? 29557 from 7 mi. S, 1 mi. E Gomez
Farias, 6500 ft., November 18, 1949, weight, 41 gms.

This subspecies of the Cactus Wren occurs throughout Coahuila except
in the extreme southeastern section of the State, where the subspecies C. b.

Birds From Coahuila, Mexico 485

guttatus occurs. Miller (1955a: 169) found C b. couesi breeding and occupy-
ing the open swales and mesas at the base of the mountains of the Sierra
del Carmen. Burleigh and Lowery (1942:198) found C. brunneicapillum "to
be rather scarce and decidedly local in its distribution" and observed an oc-
casional bird "in the open desert country west of Saltillo." Miller, Friedmann,
Griscom, and Moore (1957:151) recorded C b. couesi "south to [the] vicinity
of Monclova."

No. 29557 does not have any characters of C. b. guttatus; its under tail
coverts and flanks have roundish black spots, rather than black bars as in
C. b. guttatus.

**Campylorhynchus brunneicapillum guttatus (Gould). — This subspecies
of Cactus Wren seems to occupy the extreme southeastern section of Coahuila.
Miller, Friedmann, Griscom, and Moore (1957:152) recorded C b. guttatus
from Hipolito.

Telmatodytes palustris plesius ( Oberholser ) . — Miller, Friedmann, Griscom,
and Moore (1957:148) reported this subspecies of the Long-biUed Marsh
Wren from 8 mi. S Cuatro Cienegas.

*Catherpes mexicanus albifrons (Giraud). — Miller (1955a: 170) found this
subspecies of Canon Wren "in shaded rocky canyons and on larger cliff
slopes at the base of the mountains from 4700 to 5300 feet" in the Sierra del
Carmen where it nested. Burleigh and Lowery (1942:198) noted that the
Caiion Wren was "decidedly uncommon" at Saltillo and obtained a male at the
Chorro de Agua on April 19. Ridgway (1904:657) listed C. m. albifrons from
Patos.

*Salpinctes obsoletus obsoletus (Say). — Specimens examined: total 4: $

31067 from 1 mi. N Boquillas, 700 ft., March 6, 1952, weight, 16.1 gms.; 9

31068 from 7 mi. S, 2 mi. E Boquillas, 800 ft., March 1, 1952, weight, 18.2
gms.; sex ? 29558 from 12 mi. N, 10 mi. E Parras, 5000 ft., November 11, 1949,
weight, 16.9 gms.; and $ 32089 from 7 mi. S, 4 mi. E Bella Union, 7200 ft.,
June 24, 1952.

The Rock Wren is common in Coahuila. Miller (1955a: 170) found S. o.
obsoletus "only in the rocky piedmont and on lower bare canyon faces" and
stated that Marsh took a bird in fresh fall plumage on September 6 at El
Jardin. Sutton and Burleigh (1939a:37) found the Rock Wren "near San
Pedro." Burleigh and Lowery (1942:198) wrote that the subspecies obsoletus
was "characteristically a bird of the arroyos of the arid plateau about Saltillo,
where it was fairly common. . . ." The large size of the testes ( 5x3 mm. )
of No. 32089 and the date (June 24) on which it was obtained suggest breed-
ing by the Rock Wren 7 mi. S and 4 mi. E Bella Union.

*Mimus polyglottos leucopterus (Vigors). — Specimens examined: total 5:
9 31070 from 10 mi. S, 5 mi. E Boquillas, 1500 ft., March 5, 1952, weight,

55.1 gms.; $ $ 32094-32095 from 2 mi. W Jimenez, 850 ft., June 20, 1952;
$ 32096 from 5 mi. N, 19 mi. W Cuatro Cienegas, 3250 ft, July 5, 1952;

and $ 33186 (skeleton only) from Parras, July 5, 1955, testes, 6x3 mm.

The Mockingbird is sparsely distributed throughout Coahuila. Miller
(1955a: 170) found M. p. leucopterus in the mesquite and catclaw at the
base of the mountains in the Sierra del Carmen. Sutton and Burleigh
(1939a:37) reported M. p. leucopterus from Diamante Pass. Amadon and

486 Unh^ersity of Kansas Publs., Mus. Nat. Hist.

Phillips (1947:578) found a young Mockingbird out of the nest begging
for food from an adult on August 18 at Las Delicias. Burleigh and Lowery
(1942:199) found the Mockingbird on the arid plateau "about Saltillo."
Hellmayr (1934:308) hsted M. p. leucopterus from Jaral. Findley saw
Mockingbirds 2 mi. S and 3 mi. E San Juan de Sabinas on June 22, 1952.
Dickerman saw Mockingbirds in the Sierra del Pino on May 12, 1954, and 8
mi. E and 2 mi. S Americanos on May 18, 1954. The sizes of the testes
(8, 7 mm.) of Nos. 32094 and 32096, respectively, suggest breeding 2 mi. W
Jimenez and 5 mi. N and 19 mi. W Cuatro Cienegas, as does No. 32095, a
juvenile.

*Toxostoma longirostre sennetti (Ridgway). — Specimen examined: one, $
32090, from 2 mi. S, 3 mi. E San Juan de Sabinas, June 22, 1952.

In Coahuila the Long-billed Thrasher seems to be uncommon. It has been
recorded in Coahuila as far west as San Juan de Sabinas. Ridgway ( 1907:192)
recorded T. I. sennetti from Sabinas, the only other record of the Long-billed
Thrasher, to my knowledge, from the State. The large size of the testes
(11x6 mm.) of No. 32090 and the date (June 22) on which it was obtained
suggest that T. I. sennetti breeds 2 mi. S and 3 mi. E San Juan de Sabinas.

*Toxostoma curvirostre celsum Moore. — Specimens examined: total 2: 9
31071 from 7 mi. S, 2 mi. E Boquillas, March 1, 1952, wing, 111 mm., tail,
114 mm., weight, 97.2 gms.; and 9 31072 (skeleton only) from 10 mi. S,
5 mi. E Boquillas, 1500 ft., March 5, 1952.

This subspecies of the Curve-billed Thrasher occurs in northwestern Coa-
huila. Specimens of T. c. celsum and oberholseri from Coahuila are too few
to show clearly the distribution and intergradation in Coahuila.

No. 31071 is referred to T. c. celsum because of large size; the spots on its
upper abdomen, which are large and pronounced, suggest a relationship with
T. c. oberholseri. Miller (1955a: 170) remarked that T. c. celsum was a scarce
resident of the desert scrub at the mouth of Boquillas Canyon of the Sierra
del Carmen.

*Toxnstoma curvirostre oberholseri Law. — Specimens examined: total 5:
? 35405 (skelton only) from 4 mi. N San Isidro, May 11, 1954; 9 32091
from 5 mi. N, 19 mi. W Cuatro Cit'negas, 3250 ft., July 5, 1952; $ 32833 fnim
Parras, July 4, 1955, weight, 76.5 gms.; 9 32092 from 7 mi. S, 4 mi. E Bella
Union, 7200 ft., June 25, 1952; and $ 31614 from 16 mi. W San Antonio
de las Alazanas, 6500 ft., January 7, 1954, weight, 90 gms.

This subspecies of the Curve-billed Thrasher occurs in eastern and south-
ern Coahuila. Amadon and Phillips (1947:578) took a Curve-billed Thrasher
twenty miles west of Saltillo that had an enlarged ovary and a brood patch
still somewhat evident on August 27. Burleigh and Lowery (1942:199) stated
that T. c. oberholseri "was rather widely and commonly distributed, being
noted from the area about the summit of Diamante Pass at 7,800 feet down
to the desert country about Saltillo." Miller, Friodmann, Griscom, and Moore
(1957:177) recorded T. c. oberholseri from Sabinas, from 8 mi. S Cuatro
Cic'nc'gas, and from El Diamante. Hellmayr (1934:298) also recorded T. c.
oberholseri from Sabinas, as did Ridgway (1907:199) under the name T. c.
curvirostre before the subspecies oberholseri was named. The female from 4
mi. N San Isidro had an egg in its oviduct. The immature male (32833), the
large size of ovum (8 mm.) of No. 32092, and the presence of a brood patch

Birds From Coahuila, Mexico 487

on No. 32091 also are evidences of breeding by the Curve-billed Thrasher in
Coahuila.

*Toxostoma dorsale dorsale Henry. — The Crissal Thrasher is uncommon in
Coahuila. The subspecies dorsale occvirs in northern Coahuila. Miller ( 1955a:
170-171) found the subspecies dorsale, at about 4700 feet, only in the mes-
quite, desert willow, and walnut scrub along the wash of Boquillas Canyon
of the Sierra del Carmen and remarked also that the bird nested there.

*'*Toxostoma dorsale dumosum Moore. — Specimen examined: one, sex ?
29559, from 8 mi. N La Ventura, 6000 ft., November 17, 1949, weight, 57.0
gms.

The subspecies dumosum of the Crissal Thrasher in Coahuila has been re-
ported only from the southeastern section of the State. Burleigh and Lowery
(1942:199-200) found T. d. dumosum "not uncommon in the lower foot-hills
outside of Saltillo as well as on the summit of Diamante Pass." The speci-
men of T. d. dorsale from Diamante Pass reported by Sutton and Burleigh
(1939a: 37) is closer, according to Burleigh and Lowery (1942:199), to T. d.
dumosum. No. 29559 is darker above and below than typical specimens of
T. d. dorsale and represents T. d. dumosum.

Oreoscoptes montanus (Townsend). — Specimen examined: one, sex ?
30237, from 1 mi. SW San Pedro de las Colonias, 3700 ft., February 8, 1951.

The Sage Tlrrasher seems to be a winter visitant to Coahuila. Miller, Fried-
mann, Griscom, and Moore (1957:173) recorded the species in November
from 8 mi. S Cuatro Cienegas.

*Turdus migratorius propinquus Ridgway. — Specimen examined: one, 9
31073 (skeleton only) from 4 mi. W Hacienda La Mariposa, 2300 ft., March
24, 1952.

Burleigh and Lowery (1942:200) stated that "the Robin apparently breeds
rather sparingly on the higher ridges" in southeastern Coahuila. They col-
lected a pair "in the open pine woods just below the summit of Diamante
Pass" on April 15 and noted another at the Chorro del Agua on April 19.

**Ridgwayia pinicola (Sclater). — Specimen examined: one, $ 31619, from
5 mi. W, 22 mi. S Ocampo, 6000 ft., December 15, 1953, weight, 88 gms.

The Aztec Thrush is rare in Coahuila. Van Hoose (1955:302) remarked
that No. 31619, the skull of which was incompletely ossified, "represents the
northernmost record for this species, which was previously unknown in Coa-
huila."

Hylocichla guttata guttata (Pallas). — Specimens examined: total 2: S $
31074-31075 from 4 mi. W Hacienda La Mariposa, 2300 ft., March 25 and
26, 1952, weights, 25, 21 gms.

The Hermit Thrush is a common migrant or winter visitant in Coahuila.
H. g. guttata has been reported from northern Coahuila. Miller (1955a: 171)
observed H. g. guttata (and H. g. auduboni) in the Douglas fir and pine-oak
belts and in the lower levels in the oaks at the foot of the Sierra del Carmen.
Bvurleigh and Lowery (1942:200) noted H. g. guttata "in small numbers in
the open woods surrounding the summit of Diamante Pass, and at infrequent
intervals in the arroyos on the arid plateau near Saltillo."

488 Univ^ersity of Kansas Publs., Mus. Nat. Hist.

Hylocichla guttata sequoiensis (Belding). — Ridgway (1907:45) recorded
this subspecies of Hermit Thrush from Sierra Guadalupe in April. However,
Miller, Friedmann, Griscom, and Moore (1957:188) suggest that the ma-
terial on which this identification was based needs redetermination.

Hylocichla guttata auduboni (Baird). — Specimens examined: total 3: 9
31488 from Sierra del Pino (=5 mi. S, 3 mi. W Acebuches), May 12, 1954,
weight, 25 gms.; 9 31076 (skeleton only) from Fortin (=:33 mi. N, 1 mi.
E San Geronimo), 3300 ft., March 28, 1952; and $ 31077 (skeleton only)
from 26 mi. W Santa Teresa, 7050 ft., April 4, 1952.

Miller (1955a: 171) found this subspecies of Hermit Thrush wintering with
H. g. guttata in the Sierra del Carmen. Hellmayr (1934:456) hsted H. g.
auduboni from Sabinas.

**SiaZia sialis fulva Brewster. — Hellmayr (1934:479) listed this subspecies
of the Eastern Bluebird from Sabinas.

*Sialia mexicana mexicana Swainson. — Miller, Friedinann, Griscom, and
Moore (1957:199) remarked that the subspecies mexicana of the Western
Bluebird breeds in the southern mountains of Coahuila where, at El Diamante,
on July 7, a specimen (or specimens ?) in breeding condition was obtained.
Burleigh and Lowery (1942:200) found S. m. mexicana "well distributed
in the open woods about" Diamante Pass, but at no time below an eleva-
tion of approximately 6500 feet. Sutton and Burleigh (1939a:38) also saw
"several brown-backed" Western Bluebirds at Diamante Pass on March 6.
Ridgway (1907:150) recorded S. m. mexicana from Saltillo, Carneros, and
Sierra Guadalupe.

Sialia currucoides (Bechstein). — Specimen examined: one, 9 31078, from
Sierra de la Encantada (=38 mi. S, 23 mi. E Boquillas), 4400 ft., March 14,
1952, weight, 23.7 gms.

The Mountain Bluebird is a winter visitant to Coahuila. Miller, Fried-
mann, Griscom, and Moore (1957:200) recorded Sialia currucoides from
Hipolito on February 24.

**Myadestes townsendi townsendi (Audubon). — Specimen examined: one,
9 3107"9, from 26 mi. W Santa Teresa, 7050 ft., April 4, 1952.

Miller (1955a: 171) detected Townsend's Solitaire in clumps of large pines
in two difiFerent locations at 7000 and 7500 feet on April 4, 6, and 8 in
the Sierra del Carmen. He did not find a breeding population of M. t.
townsendi, but did note favorable habitat for breeding. Ridgway (1907:164)
recorded M. townsendi from the Sierra Guadalupe on April 21. Dickerman
saw a Townsend's Solitaire in the Sierra de la Madera on December 13, 1953.

The underparts of No. 31079 are not uniformly dark, being paler on the
chin, throat, and abdomen than elsewhere as is true of typical representatives
of M. t. townsendi.

** Polioptila caerulea caerulea (Linnaeus). — Specimen examined: one, $
32097, from 2 mi. S, 11 mi. E Nava, June 15, 1952.

No. 32097 is the first record of occurrence of this su]:)spocics of Blue-gray
Gnatcatcher in the State. The white of the undcri^arts of No. 32097 is
less grayish than the underparts of typical representatives of F. c. amoenissima,
and the black at the base of the inner webs of the outermost rcctri.x docs not

Birds From Coahuila, Mexico 489

extend beyond the tip of the under tail coverts. Representatives of P. c.
amoenisdma have black at the base of the inner web of the outermost rectrix
more extended, usually showing beyond the tip of the under tail coverts.
The size of the testes (3x2 mm.) of No. 32097 does not suggest breeding,
but the date (June 15) indicates that it was a resident.

*Polioptila caerulea amoenissima Grinnell. — Specimen examined: one, $

31080, from 7 mi. S, 2 mi. E Boquillas, 800 ft., February 29, 1952, weight,
5.4 gms.

This subspecies of the Blue-gray Gnatcatcher breeds in Coahuila and occurs
tliroughout all of the State except the northeastern section. Miller (1955a:
171 ) remarked that "this gnatcatcher was apparently established on summer
territories in the oaks and walnuts of the wash of Boquillas Canyon at tlie
foot of the mountains" of the Sierra del Carmen. Burleigh and Lowery
(1942:200) noted "this species only in the open woods at the summit of Dia-
mante Pass, where, however, it was not uncommon." Miller, Friedmann,
Griscom, and Moore (1957:201) recorded P. c. amoenissima breeding at El
Diamante on July 8.

* Polioptila m^lanura melanura Lawrence. — Specimen examined: one, $

31081, from 7 mi. S, 2 mi. E Boquillas, 800 ft., March 1, 1952, weight, 5.6 gms.

In Coahuila this subspecies of the Black-tailed Gnatcatcher has been re-
corded from several localities. Burleigh and Lowery (1942:200) found that
it "was hmited in its distribution to the lower altitudes and was noted only in
the open desert country west of Saltillo." Sutton and Burleigh (1939a: 38)
noted it "several times near San Pedro" where on January 29 one female was
taken. Miller, Friedmann, Griscom, and Moore (1957:205) recorded P. m.
melanura at HipoHto on June 30 to July 2 in breeding condition.

Regulus satrapa satrapa Lichtenstein. — Miller (1955a: 171) found a small
wintering flock of Golden-crowned Kinglets in the Sierra del Carmen; this is
the only record of the species in Coahuila.

Regulus calendula calendula (Linnaeus). — Specimens examined: total 5:
9 31085 from the Rio Grande (= 17 mi. S Dryden, Terrell Co., Texas, in
Coahuila), 600 ft., March 19, 1952, weight, 6.3 gms.; S S 31082-31083 from
1 mi. N Boquillas, 700 ft., March 8, 1952, weight, 7.2, 6.5 gms.; S 31084
from Sierra de la Encantada (=38 mi. S, 23 mi. E Boquillas), 4400 ft.,
March 15, 1954, weight, 5.4 gms.; and $ 31661 from the north slope of Sierra
Guadalupe (=11 mi. S, 7 mi. W General Cepeda), 7800 ft., April 20, 1953,
weight, 5 gms.

In Coahuila this subspecies of the Ruby-crowned Kinglet is a common
migrant. Miller (1955a: 171) found it (and R. c. cinetaceus) "common in
the conifers and oaks of the upper levels of the mountains [Sierra del Carmen],
at 6500 to 7000 feet, as winter visitants or migrants." Burleigh and Lowery
(1942:201) took specimens of R. c. calendula at Diamante Pass on April 15,
at the Chorro del Agua on April 19, and at 20 mi. W Saltillo on April 22.
Sutton and Burleigh (1939a:38) noted the Ruby-crowned Kinglet "in the
arid parts of southern Coahuila." Hellmayr (1934:513) listed JR. c. calendula
from Sabinas. Dickerman saw Ruby-crowned Kinglets in the Sierra de la
Madera on December 13, 1953, 20 mi. S Ocampo on April 4, 1954, and 3 mi.
S and 13 mi. E San Antonio de las Alazanas on January 12, 1954.

490 University of Kansas Publs., Mus. Nat. Hist.

Regulus calendula cineraceus Grinnell. — Miller (1955a: 171) found R. c.
cineraceus common in the Sierra del Carmen; on April 3, 5, and 10 the birds
were "abundant, as though a wave of migrants were passing through."

Anthus spinoletta rubescens (Tunstall). — Specimens examined: Total 3:
S $ 31086-31087 and sex ? 31088 from 1 mi. N Boquillas, 700 ft., March 6,
7, and 8, 1952, weights, 19.3, 19.9, and 16.6 gms.

This subspecies of the Water Pipit is an uncommon winter visitant or mi-
grant in Coahuila. Burleigh and Lowery (1942:201) found "a flock of ten
birds ... on the outskirts of Saltillo" on April 18. Miller, Friedmann,
Griscom, and Moore (1957:210) recorded A. s. rubescens from Cuatro Cicne-
gas in November and from Hipolito in February.

Anthus spinoletta pacificus Todd. — Miller, Friedmann, Griscom, and Moore
(1957:209) recorded this Water Pipit from Cuatro Cienegas in February and
from Hipohto in November.

Bombijcilla cedrorum Vieillot. — The Cedar Waxwing is an uncommon
winter visitant to Coahuila. Miller (1955a: 171) recorded a flock in the
Sierra del Carmen on April 5, and another flock on April 21. Burleigh and
Lowery (1942:201) saw two small flocks on April 15 "in the open woods just
below the summit of Diamante Pass."

*Phainopepla nitens nitens Swainson. — Specimen examined: one, $ 31674,
from the west foot of Pico de Jimulco, 5000 ft., April 3, 1953, weight, 35 gms.

The Phainopepla occurs throughout most of Coahuila. Sutton and Bur-
leigh (1939a:39) reported P. n. nitens from Diamante Pass on March 6. On
April 15 and 17 Burleigh and Lowery (1942:201) saw scattered pairs of the
Phainopepla "only in the open woods surrounding Diamante Pass." Miller
(1955a: 171) noted P. nitens "on April 20 and 28 in large clumps of mcsquite
near Piedra Blanca, at about 4500 feet, on the foothills" of the Sierra del
Carmen. Dickerman saw a Phainopepla 20 mi. S Ocampo on April 4, 1954.
Baird (1858:320) listed a male P. n. nitens from Coahuila, Mexico. Hell-
mayr (1935:107) remarked that Phainopepla nitens was listed from Coahuila
by "Salvin and Godman, Biol. Centr. — Amer., Aves, 1, p. 220, 1883. . . ."
Miller, Friedmann, Griscom, and Moore (1957:213) stated that P. n. nitens
breeds at El Diamante. The long wing (100 mm.) and long tail (96 mm.)
of No. 31674 is typical for P. n. nitens.

Phainopepla nitens lepida Van Tyne. — Sutton and Burleigh (1939a: 39)
recorded P. n. lepida from Diamante Pass on March 6; Miller, Friedmann,
Griscom, and Moore (1957:213) suggest that this individual was a vagrant.

Lanius ludovicianus migrans Palmer. — Burleigh and Lowery (1942:202)
obtained this subspecies of Loggerhead Shrike "on the outskirts of Saltillo
on April 20." The specimen shows evidence of intergradation with excu-
bitorides.

** Lanius ludovicianus excubitorides Swainson. — Miller, Friedmann, Gris-
com, and Moore (1957:216) recorded L. I. excubitorides from Sabinas and
from Hipolito (November 2 to February 24).

Birds From CoAHxnLA, Mexico 491

*Lanius ludovicianus mexicanua Brehm. — Specimens examined: total 4;
sex ? 31089 from 7 mi. S, 2 mi. E Boquillas, 800 ft., February 29, 1952,
weight, 45.1 gms.; $ 31090 from 36 mi. S, 15 mi. E Boquillas, 2550 ft.,
March 12, 1952; sex ? 30233 from 1 mi. N San Lorenzo, 4200 ft., February
5, 1951; and 9 32098 from 7 mi. S, 4 mi. E Bella Uni6n, 7200 ft., June 25,
1952.

Miller (1931:66) suggested that L. /. mexicanus and L. I. excubitorides
intergrade in Coahuila; all of the specimens of Loggerhead Shrike from
Coahuila that I have examined are intergrades between mexicanus and ex-
cubitorides. Our four specimens have a superciliary line that is indistinct
and the black mask of each extends somewhat posterior to the auricular
region. The anterior part of their forehead is somewhat lighter than the
remaining part of their head and back.

Miller (1955a: 171) detected L. I. mexicanus only once in catclaw scrub
in the lower part of Boquillas wash at about 4600 feet in the Sierra del
Carmen. He (loc. cit.) remarked that his individual may best be considered
an intergrade between mexicanus and excubitorides, being "somewhat closer
to the fonner." Burleigh and Lowery (1942:201) obtained a male L. I.
mexicanus "in the open valley just below the summit of Diamante Pass on
April 23" that was typical of this subspecies. Burleigh and Lowery ( 1942:
201-202) suggested that L. I. mexicanus breeds in southeastern Coahuila.
Sutton and Burleigh (1939a:39) took a typical representative L. I. mexicanus
at Mayran on January 30.

Lanius ludovicianus gambeli Ridgway. — Miller, Friedmann, Griscom, and
Moore (1957:214) recorded this subspecies from Hipolito on November 6.

* Vireo atricapilla Woodhouse. — Specimens examined: total 4: sex ?
32099-32100 from Sierra del Pino (= 6 mi. N, 6 mi. W Acebuches). 5250 ft.,
July 3, 1952; and $ $ 31493-31494 from 16 mi. E, 18 mi. N Ocampo, May 8
and 9, 1954, enlarged testes.

The Black-capped Vireo seems to breed as far south as central Coahuila.
Miller ( 1955a: 171-172) reported this vireo as a summer resident in "the low
catclaw-dominated scrub in the lower washes of Boquillas Canyon and its
side valleys, at 4600 to 4800 feet" in the Sierra del Carmen. Dickerman
found the Black-capped Vireo common on the dry scrub and oak hillside
habitat 16 mi. E and 18 mi. N Ocampo.

Vireo griseus noveboracensis (Gmelin). — Ridgway (1904:184) recorded
this subspecies of the White-eyed Vireo "west to Sabinas." Miller, Fried-
mann, Griscom, and Moore (1957:220) also recorded V. g. noveboracensis
from Sabinas on May 25. I suspect that the specimen of V. g. noveboracensis
from Sabinas that was taken on May 25 was a vagrant. V. g. noveboracensis
may occur in Coahuila as a migrant or winter visitant; however, I do not be-
heve that representatives of noveboracensis normally are resident in Coahuila.

*Vireo griseus micrus Nelson. — Specimen examined: one, $ 32101, from
2 mi. W Jimenez, 850 ft., June 20, 1952, measiurements: wing, 58 mm.; tail,
43.5 mm.; culmen, 10 mm.; tarsus, 19 mm.

This subspecies of the White-eyed Vireo breeds in Coahuila. Miller, Fried-
mann, Griscom, and Moore (1957:221) recorded V. g. micrus from Sabinas

4—7905

492 University of Kansas Publs., Mus. Nat. Hist.

on March 9 and May 14. The enlarged testes (5x3 mm.) of No. 32101 and
the date (June 20) on which it was obtained suggest breeding in Coahuila,

Vireo huttoni stephensi Brewster. — This subspecies of Hutton's Vireo occurs
in southeastern Coahuila as a migrant. Sutton and Burleigh (1939a: 39) found
V. h. stepliensi fairly common at Diamante Pass on March 6. Ridgway (1904:
198) recorded V. li. stephensi from Sierra Guadalupe in April.

* Vireo huttoni carolinae Brandt. — Specimens examined: total 3: $ 31588
from 20 mi. S Ocampo, 6500 ft., April 5, 1954, weight, 11 gms.; 9 32851
from 13 mi. E San Antonio de las Alazanas, July 6, 1955, weight, 14.2 gms.;
and $ 32102 from 7 mi. S, 4 mi. E Bella Union, 7200 ft., June 25, 1952.

To my knowledge, V. h. carolinae is the only resident subspecies of Hut-
ton's Vireo in Coahuila. Burleigh and Lowcry (1942:202) found V. h.
carolinae in rather limited numbers in the woods bordering the summit of
Diamante Pass. Miller (1955a: 172) remarked that the subspecies carolinae
was a common bird from 6500 feet to 8000 feet in the Sierra del Carmen.
Miller {loc. cit.) took a female on April 12 that was nearly ready to lay and
said that his specimens of carolinae from the Sierra del Carmen seem to be
separate from V. h. stephensi and V. h. mexicanus. Our specimens showed
no overlapping of characters with V. h. stephensi and V. h. mexicanus. The
size of the largest ovum (2 mm.) of No. 32102 and the dates (June 25, and
July 6) on which Nos. 32102 and 32851 were obtained suggest tliat V. h.
carolinae is a resident in Coahuila.

* Vireo bellii medius Oberholser. — Specimen examined: one, S 31495, from
San Marcos, May 5, 1954, measurements: wing, 56 mm.; tail, 48 mm.; culmen,
9.5 mm.; tarsus, 18 mm.

Although Bell's Vireo seems to have been observed uncommonly in Coa-
huila, this species does breed in the State. Burleigh and Lowery (1942:202)
obtained a single female V. b. medius on April 20 "in an arroyo east of Saltillo"
and found this subspecies "not uncommon in the open desert twenty miles
west of Saltillo, where three singing males were secured." Miller, Friedmann,
Griscom, and Moore (1957:226) recorded V. b. medius in breeding condition
at Hipolito from June 30 to July 3. Ridgway (1904:207) recorded the sub-
species from Monclova. Hardy saw a Bell's Vireo at Parras on July 4, 1955.
The size of No. 31495 is typical for V. b. medius.

*Vireo flavifrons Vieillot. — Specimen examined: one, $ 32103, from 2 mi.
W Jimenez, 850 ft., June 20, 1952.

Van Hoose (1955:302-303) suggested that the occurrence of No. 32103
in Coahuila is evidence of a southward extension of the range of the Yellow-
throated Vireo within the last generation. The size of the testes (5x3 mm.)
of No. 32103 and the date (June 20) on which it was obtained suggest breed-
ing by Vireo flavifrons in Coahuila.

Vireo solitarius solitarius (Wilson). — Specimen examined: one, 9 31640,
from 9 mi. E Hermanas (i=Canyon de Parajos in the Sierra de Parajos Azule),
2100 ft, December 7, 1953.

This subspecies of the Solitary Vireo is an uncommon migrant or winter
visitant to Coahuila. Dickerman obtained No. 31540, whose skull was un-
ossified and whose ovary was small, in an oak and palm habitat. The bright

Birds From Coahuila, Mexico 493

yellow flanks, large and yellow wing bars, and the uniform olive green back
indicate that this specimen is a typical representative of V. s. solitarius. This
subspecies was previously unrecorded in Coahuila.

Vireo solitarius cassinii Xantus. — Specimens examined: total 2: $ 35408
(skeleton only) from 5 mi. W, 3 mi. S Acebuches, 6200 ft., May 12, 1954;
and $ 31589 from 20 mi. S Ocampo, 6200 ft., April 4, 1954, measurements:
wing, 75 mm.; tail, 57 mm.; cuhnen, 10.5 mm.; weight, 14 gms.

This subspecies of the Solitary Vireo seems to be uncommon in Coahuila.
No. 31589 provides the first record of V. s. cassinii in Coahuila. Van Hoose
has (1955:303) erroneously reported that Dickerman obtained No. 31589 on
July 4, 1954; the correct date is April 4, 1954.

Although the size of No. 31589 is large for V. s. cassinii, the color (sides
and flanks with less yellow, more olive; narrow white wing bars) resembles
that of typical representatives of cassinii. The testes of No. 31589 were not
enlarged. Dickennan suggested that the female from 5 mi. W and 3 mi.
S Acebuches showed some resemblance to V. s. plumbeus. This is not to be
unexpected since the subspecies plumbeus has been reported from the Chisos
Mountains of Texas (Van Tyne and Sutton, 1937:82) and from northern
Chihuahua (Miller, Friedmann, Griscom, and Moore, 1957:227).

*Vireo olivaceus (Linnaeus). — Specimen examined: one, $ 32104, from
12 mi. N, 12 mi. W Jimenez, 850 ft., June 19, 1952.

The Red-eyed Vireo has been uncommonly reported from eastern Coahuila.
Miller, Friedmann, Griscom, and Moore (1957:229) recorded V. olivaceus
from Sabinas on May 22 and from Las Vacas Creek on June 7 as late spring
migrants. Hellmayr (1935:131) listed the Red-eyed Vireo from northern Coa-
huila. Findley saw a Red-eyed Vireo 2 mi. W Jimenez on June 19, 1952. The
size of the testes (5x3 mm.) of No. 32104 and the date (June 19) on which
it was obtained indicate that the Red-eyed Vireo possibly breeds in north-
eastern Coahuila; if so, this is the first breeding record of the Red-eyed
Vireo in Coahuila.

Vireo gilvus gilvus (Vieillot). — This subspecies of the Warbling Vireo is
an uncommon migrant in Coahuila. Miller, Friedmann, Griscom, and Moore
(1957:232) recorded V. g. gilvus from 12 mi. W Saltillo on September 28.

Mniotilta varia (Linnaeus). — Specimen examined: one, $ 31662, from
the north slope of Sierra Guadalupe (=11 mi. S, 7 mi. W General Cepeda),
7800 ft., April 20, 1953, weight, 10 gms., testes not enlarged.

The Black and White Warbler is an uncommon visitant or migrant in
Coahuila. Miller (1955a: 172) remarked that Marsh took a fall migrant
on September 1 in Chuperosa Canyon in the Sierra del Carmen. Burleigh
and Lowery (1942:202) secured a female Black and White Warbler "in an
orchard on the outskirts of Saltillo" on April 20 and a male "near the top of
Diamante Pass on April 23."

Vermivora celata celata (Say). — Specimens examined: total 2: sex ? 31091
from the Rio Grande (=17 mi. S Dryden, Terrell Co., Texas, in Coahuila),
600 ft., March 19, 1952, measvirements: wing, 57 mm.; tail, 47 mm.; weight,
7.7 gms.; and $ 31092 from 4 mi. W Hacienda La Mariposa, 2300 ft., March
25, 1952, measurements: wing, 62 mm.; tail, 48 mm.; weight, 9.2 gms.

This subspecies of the Orange-crowned Warbler is an uncommon migrant

494 Unr-ersity of Kansas Pxjbls., Mus. Nat. Hist.

in Coahuila. In Brewster County, Texas, Van Tyne and Sutton (1937:83)
found V. c. celata "not common as a spring transient." Miller, Friedmann,
Griscom, and Moore (1957:239) recorded V. c. cehta from Coahuila. The
quality of the pale yellow color and the sizes of Nos. 31091-31092 suggest
that they are representatives of V. c. celata.

Vermivora celata orestera Oberholser. — Burleigh and Lowery (1942:202)
found V. c. orestera "only on infrequent occasions ... in the open
woods surrounding the summit of Diamante Pass" where they obtained one
specimen.

Vermivora virginiae (Baird). — Miller (1955a:172) took a male Virginia's
Warbler in Boquillas Canyon in the Sierra del Carmen "in scattered scnibby
oak growth with grass and cactus beneath." This species in the Sierra del
Carmen is considered "casual" by Miller, Friedmann, Griscom, and Moore
(1957:241).

** Vermivora crissalis (Salvin and Godman). — Specimen examined: one,
S 31590, from 20 mi. S Ocampo, 7000 ft., April 5, 1954, weight, 10 gms.

The Colima Warbler is common locally in Coahuila. Burleigh and Lowery
(1942:203) found Uiis species fairly common on the steep, rugged slopes
above the summit of Diamante Pass and saw none below an elevation of ap-
pro.ximately 7500 feet. Bangs (1925:251) stated that Nelson and Goldman
secured a specimen of the Colima Warbler at Sierra Guadalupe on April 25.

* Vermivora superciliosa mexicana (Bonaparte). — Specimen examined: one,
S 31591, from 13 mi. E San Antonio de las Alazanas, 9350 ft., April 10, 1954.

Hartlaub's Warbler is uncommon in Coahuila and seems to occur only in
the southeastern section of the State; No. 31591 is the first record of the
species in Coahuila. The size of the testes (5x2 mm.) of No. 31591 and
the fact that the bird was singing when first seen suggest the possibility that
V. s. mexicana breeds in southeastern Coahuila. Breeding there is not un-
e.\pected because the species has been found breeding in Nuevo Leon
(Miller, Friedmann, Griscom, and Moore, 1957:242).

Parula americana (Linnaeus). — Miller (1955a: 172) obtained a migrant
Parula Warbler in an oak grove at 7000 feet on April 16 in the Sierra del
Cannen, and remarked that it was "apparently the first record of this species
in Coahuila."

*Parula pitiatjumi nigrilora Coues. — The AOU Check-list Committee (1957:
486) recorded this subspecies of the OUve-backed Warbler as a resident at
Sabinas.

* Peucedramus taeniatus arizonae Miller and Griscom. — This subspecies of
the Olive Warbler is locally common in Coahuila. Miller (1955a:172) found
P. t. arizonae common in the pine timber above 6800 feet in the Sierra del
Carmen and suggested that this warbler breeds in these mountains. Burleigh
and Lowery (1942:203) found the Ohve Warbler in a thick pine wood at
an elevation of 9500 to 10,000 feet and remarked that this species was "de-
cidly uncommon in the Diamante Pass area." They (Inc. cit.) obtained a
female (not identified to subspecies) on April 23 that had a "well developed

Birds From Coahuila, Mexico 495

brood patch and was unquestionably incubating eggs." Sutton and Burleigh
(1939a:40) took a single female at Diamante Pass on March 6 which also
was not identified to subspecies. Dickerman saw OHve Warblers in the Sierra
de la Madera on December 13, 1953, and 13 mi. E San Antonio de las
Alazanas on April 10, 1954.

Dendroica petechia morcomi Coale. — Miller, Friedmann, Griscom, and
Moore (1957:246) reported this subspecies of the Yellow Warbler as having
been recorded from Coahuila.

Dendroica auduboni auduboni (Townsend). — Specimens examined: total

2: $ 31094 (skeleton only) from Fortin (=33 mi. N, 1 mi. E San Geronimo),
3300 ft., March 29, 1952; and $ 31093 from 4 mi. W Hacienda La Mariposa,
2300 ft., March 25, 1952, weight, 12.3 gms.

Audubon's Warbler is a common winter visitant and migrant in Coahuila.
Miller (1955a: 173) recorded D. a. auduboni as a migrant from April 7 to 26
in the Sierra del Carmen; he found no suggestion of breeding by the Audubon's
Warbler in the northwestern section of the State. One individual that MiUer
( loc cit. ) obtained was extensively black and approached the characters of the
subspecies nigrifrons of Chihuahua. He (loc. cit.) suggested that the black
individual was taken from "part of a cline of blackness and size in which D. a,
auduboni of the northwest and D. a. nigrifrons of Mexico are extremes." Bm--
leigh and Lowery (1942:203) remarked that Audubon's Warbler "is doubt-
less a common winter bird in the area around Saltillo." Two specimens ob-
tained by Burleigh and Lowery {loc. cit.) "might be considered intermediate"
between auduboni and memorabilis. Sutton and Burleigh (1939a: 40) saw
Audubon's Warbler "in some numbers near San Pedro . . . [on] January
29 and 30." Dickerman saw Audubon's Warblers 13 mi. E San Antonio de
las Alazanas on April 10, 1954. Miller (1955a: 173) also obtained, in the
Sierra del Carmen, a hybrid between D. coronata and D. auduboni.

Dendroica auduboni memorabilis Oberholser. — Oberholser (1921:246) re-
corded D. a. memorabilis from Saltillo on April 17. This subspecies seems
to winter commonly in western Mexico and less commonly in the Central
Plateau and Sierra Madre Oriental (Miller, Friedmann, Griscom, and Moore,
1957:249-250).

Dendroica nigrescens (Townsend). — Specimen examined: one, $ 31095,
from Fortin (=33 mi, N, 8 mi. W San Geronimo), 3300 ft., March 28, 1952,
weight, 9.3 gms.

The Black-throated Gray Warbler is an uncommon spring and possibly fall
migrant in Coahuila. Miller (1955a: 173) found D. nigrescens uncommon in
the Sierra del Carmen. He saw and heard a spring migrant singing on April
12 at 7000 feet and obtained a male on April 16.

Dendroica townsendi (Townsend). — Townsend's Warbler is a spring and
fall migrant in Coahuila. Miller (1955a: 173) recorded D. townsendi on Sep-
tember 2 at Jardin del Sur in Chuperosa Canyon in the Sierra del Carmen.
Burleigh and Lowery (1942:203) saw two individuals of Tovrasend's Warbler
at Diamante Pass on April 14. Amadou and Phillips (1947:578) secured this
species "in mesquite about twenty miles west of Saltillo on August 28." Dick-
erman saw Townsend's Warblers in the Sierra de la Madera on December 13,

496 Uxn-ERSiTY of Kansas Publs., Mus. Nat. Hist.

1953; 20 mi. S Ocampo on April 4, 1954; and 13 mi. E San Antonio de las
Alazanas on April 10, 1954.

Dendroica virens (Gmelin). — Dickerman saw one Black-throated Green
Warbler 13 mi. E San Antonio de las Alazanas, 9350 feet, on April 10, 1954,
in a white pine-Douglas fir-aspen association. This seems to be the first record
of this species in Coahuila.

*Dendroica chrtjsoparia Sclater and Salvin. — Miller, Friedmann, Griscom,
and Moore (1957:251) listed the Golden-cheeked Warbler from Hipohto on
July 3.

Dendroica occidentalis (Townsend). — Specimen examined: one, S 31592,
from 13 mi. E San Antonio de las Alazanas, April 10, 1954.

The Hermit Warbler seems to be an uncommon spring and probably fall
migrant in Coahuila. Miller, Friedmann, Griscom, and Moore (1957:252)
reported D. occidentalis from the State. No. 31592, whose testes were not
enlarged, was obtained in a pine-siiruce-aspen association.

Geothlypis trichas brachidactylus (Swainson). — Miller, Friedmann, Gris-
com, and Moore (1957:261) listed this subspecies of the Yellowthroat as
recorded from Coahuila.

Geothlypis trichas occidentalis Brewster. — Burleigh and Lowery (1942:204)
found the Yellowthroat "only on the outskirts of Saltillo, where . . . [this
species] was not uncommon in the thickets, especially around a small stand of
marsh grass bordering a pond." Three males that Burleigh and Lowery (loc.
cit.) obtained were tentatively identified as G. t. occidentalis. Dickerman saw
one male Yellowthroat at San Marcos (=20 mi. S Cuatro Cienegas) on May
4, 1954.

Geothlypis nelsoni nelsoni Richmond. — Burleigh and Lowery (1942:204)
noted the Hooded Yellowthroat "only on the open slopes above tlie summit
of Diamante Pass at an elevation of about 8,000 feet" where they obtained
an adult male.

*Icteria virens virens (Linnaeus). — Specimens examined: total 3: $ $
32105-32106 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 19, 1952; and
9 32107 from 2 mi. W Jimenez, 850 ft., June 20, 1952.

From the paucity of records in the literature, I judge that the Yellow-
breasted Chat is uncommon in Coahuila. Miller, Friedmann, Griscom, and
Moore (1957:265) listed I. v. virens from Coahuila. Findley saw a Yellow-
breasted Chat 2 mi. S and 3 mi. E San Juan de Sabinas on June 22, 1952.
The sizes of the testes (7.x4 mm.; 12 mm.) of Nos. 32105-32106, the size of
the largest ovum (2 mm.) of No. 32107, and the dates (June 19, 20) on
which these specimens were obtained indicate breeding by 7. v. virens in
northeastern Coahuila, an area southwest of the previously documented breed-
ing range.

**Icteria virens auricollis (Dcppe). — Miller, Friedmann, Griscom, and
Moore (1957:264) listed this subspecies of the YcUow-breastcd Chat as re-
corded from Coahuila.

Birds From Coahuila, Mexico 497

Wilsonia pusilla pileolata (Pallas). — Specimens examined: total 3: $
31501 and 9 31500 from Sierra del Pino (=5 mi. S, 3 mi. W Acebuches),
May 13 and 14, 1954, measurements: wing, 59, 55 mm.; tail, 50, 49 mm.;
culmen, 8, 8.5 mm.; tarsus, 16, 16 mm.; weight: 6, 7 gms.; and $ 31663 from
the north foot of Sierra Guadalupe (=10 mi. S, 5 mi. W General Cepeda),
6500 ft., April 21, 1953, measurements: wing, 58 mm.; tail, 46 mm.; culmen,
8.5 mm.; tarsus, 16 mm.; weight, 7 gms.

Wilson's Warbler is a common spring and probably fall migrant in Coa-
huila. Miller (1955a: 173) took spring migrants of W. p. pileolata from April
9 to April 27; he found W. p. pileolata at 4800 feet and at 7000 feet. Amadon
and Philhps (1947:579) saw a Wilson's Warbler at Las Delicias on August
17. Burleigh and Lowery (1942:204) found Wilson's Warbler to be the
most abundant of the warblers that they recorded in southeastern Coahuila.
They saw W. p. pileolata on the top of the high ridges and in the arid desert
country in the southeastern section of the State. Several specimens were col-
lected by Burleigh and Lowery (loc. cit.) including an immature male from
the Chorro del Agua on April 19. Dickerman saw Wilson's Warblers 16 mi.
E and 18 mi. N Ocampo on May 7, 1954, and at San Marcos (=20 mi. S
Cuatro Cienegas) on May 4, 1954. The sizes of our specimens as well as their
color (bright olive-green above, bright yellow below) are typical for the sub-
species pileolata.

Setophaga ruticilla ruticilla (Linnaeus). — The American Redstart seems
to be uncommon in Coahuila. Miller, Friedmann, Griscom, and Moore ( 1957:
268) listed one specimen of S. r. ruticilla from the State.

* Setophaga picta picta Swainson. — Specimens examined: total 2: sex ?
31096 from 26 mi. W Santa Teresa, 7050 ft., April 5, 1952; and $ 31671
from Caflon d. Meco in Sierra Guadalupe (=10 mi. S General Cepeda), April
23, 1953, weight, 11 gms.

The Painted Redstart seems to be locally common in Coahuila. Miller
(1955a: 173) found these warblers breeding in the Sierra del Carmen and
said that they were "common from 6000 to 7500 feet in canyon bottom growth,
in oaks, and in mixed pines and oaks;" however, he did not see any of these
warblers above 7500 feet where conifers tended to dominate the vegetation.
Marsh and Stevenson (1938:287) obtained a male Painted Redstart in annual
molt on August 11 in oak and juniper forest at Vivoras Spring in the Sierra
del Carmen and reported seeing two other Painted Redstarts at 9000 feet (see
also Miller, 1955a: 173). Dickerman also saw seven Painted Redstarts 20
mi. S Ocampo on April 4, 1954. The size of the testes (7x4 mm.) of No.
31671 suggests breeding by S. p. picta in the Sierra Guadalupe.

** Passer domesticus domesticus (Linnaeus). — Burleigh and Lowery (1942:
204) remarked that the House Sparrow was not "a common bird around Sal-
tillo" although they noticed P. d. domesticus regularly. They reported House
Sparrows also from the Chorro del Agua and in the high mountain valley
south of Diamante Pass. Miller, Friedmann, Griscom, and Moore (1957:275)
recorded P. d. domesticus from Sabinas. Baker captured House Sparrows in
a bat net 12 mi. E San Antonio de las Alazanas, 9950 feet, on July 5, 1955.
Although there are no other records, the House Sparrow is probably fairly
common in the villages and towns of the State.

498 Uniyersity of Kansas Publs., Mus. Nat. Hist.

[**SturneUa magna hoopesi Stone. — The Eastern Meadowlark is uncommon
in Coahuila. Tlie AOU Check-list Committee (1957:523) listed this sub-
species of the Eastern Meadowlark from northern Coahuila.]

** Sturnella neglecta neglecta Audubon. — Specimen examined: one, 9
31098, from the Rio Grande (=17 mi. S Dryden, Terrell Co., Texas, in Coa-
huila), 600 ft., March 18, 1952, weight, 71.8 gms.

This subspecies of the \^^estern Meadowlark seems to be locally common
in the open country of Coahuila. Burleigh and Lowery (1942:205) found
this meadowlark common "about Saltillo" where a "small series" of S. neglecta
was obtained. Miller, Friedmann, Griscom, and Moore (1957:295) recorded
the Western Meadowlark from El Diamante on July 7. To my knowledge,
no specific breeding records of this meadowlark from Coahuila exist.

**Xanthocephalus xanthocephalus (Bonaparte). — Specimen examined: one,

$ 32494 (skeleton only) from Las Margaritas, August 4, 1955.

The Yellow-headed Blackbird occurs in Coahuila in migration. Miller
(1955a: 173) found this blackbird at Noria "in the flats immediately east of
the Sierra del Carmen" on April 28, and reported also that Marsh took a
male in worn breeding plumage on July 24 at Tanque de los Melones on La
Bavia Ranch east of Fresno Mesa. Amadon and PhiUips (1947:579) took
two adult males at Las Delicias on August 11 and 15. Dickerman also saw
a female 8 mi. E and 2 mi. S Americanos on May 18, 1954. Van Hoose
saw a male at Torreon on July 2, 1955.

* Agelaius phoeniceus megapotamus Oberholser. — Specimens examined:
total 5: S 32124, 9 32126, and 9 32128 from 12 mi. N, 12 mi. W Jimenez,
850 ft., June 19, 1952; $ 32125 from 2 mi. W Jimenez, 850 ft., June 20,
1952; and 9 32127 from 9 mi. S, 11 mi. E Sabinas, June 14, 1952.

This subspecies of the Redwinged Blackbird is common in eastern Coahuila.
There are no records of the species from western Coahuila. Burleigh and
Lowery (1942:205) obtained a male at "the edge of Saltillo" on April 24.
Oberholser (1919a: 23) recorded A. p. megapotamus from Porfirio Diaz on
June 2, 5, and 6. The presence of juveniles (32126, 32128) from 12 mi. N
and 12 mi. W Jimenez and (32125) from 2 mi. W Jimenez, respectively, and
the dates (June 14, 19, 20) on which the University of Kansas specimens
were obtained are evidence of breeding by A. p. megapotamus in northeastern
Coahuila.

* Icterus spurius (Linnaeus). — Specimens examined: total 8: $ $ 31536-
31537 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 19, 1952; $ 31538
from 2 11. i. S, 3 mi. E San Juan dc Sabinas, 1160 ft., June 23, 1952; S S
32605-32607 from Parras, July 4, 1955; $ 32604 and 9 32603 from Hacienda
San Lorenzo, July 3, 1955, weights, 19.4, 18.5 gms.

The Orchard Oriole seems to occur fairly commonly in eastern and south-
ern Coahuila and breeds in the State. Amadon and Phillips (1947:579)
reported that Orchard Orioles were common in the desert "about Las Delicias"
in August and September and probably were migrants. Dickerman collected
Nos. 32605-32606 along an irrigated field-edge that consisted of cottnnwood
and oak; he obtained Nos. 32603-32604 in an irrigated pecan orchard. The
sizes of the testes (10x5 mm.; 10x5 mm.; 8x7 mm.; 8.\7 mm.; 10x8 mm.) of

Birds From Coahuila, Mexico 499

Nos. 31536, 31537, 32605, and 32604, respectively, and the size of the largest
ovum (2 mm.) of No. 32603 as well as the dates (June 19, 23; July 3, 4)
on which all these specimens were collected indicate breeding by this species
in the State.

* Icterus cucullatus cucuUatus Swainson. — Specimens examined: total 2:
$ 32123 from 2 mi. W Jimenez, 850 ft., June 21, 1952; and $ 32122 from
2 mi. S, 3 mi. E San Juan de Sabinas, 1160 ft., June 23, 1952.

The Hooded Oriole apparently is uncommon in Coahuila. Miller, Fried-
mann, Griscom, and Moore (1957:290) listed I. c. cucullatus from Sabinas.
The size of the testes (11x6 mm.) of No. 32122 and the dates (June 21,
23) on which Nos. 32123 and 32122 were collected as well as the juvenile
male (tail, 80.5 mm.) from 2 mi. W Jimenez suggest breeding by this sub-
species in Coahuila.

* Icterus parisorum Bonaparte. — Specimen examined: one, 5 32121, from
7 mi. S, 4 mi. E Bella Union, 7200 ft., June 25, 1952.

Scott's Oriole is common in Coahuila. Miller (1955a: 173) found this
oriole breeding in the canyons at the base of the Sierra del Carmen. Burleigh
and Lowery (1942:205) found this oriole limited to the higher altitudes above
7000 feet, and took specimens at Diamante Pass and at the Chorro del Agua
on April 19. Amadon and Philhps (1947:579) found Scott's Oriole "not
uncommon in the arroyos near Las Dehcias" and reported a juvenile "barely
out of the nest and able to fly only a few feet . . ." on August 15. No.
32121 had an egg in its oviduct. Dickerman saw Scott's Orioles in the Sierra
del Pino on May 12, 1954, and 16 mi. E and 18 mi. N Ocampo on May 7,
1954.

** Icterus wagleri wagleri Sclater. — Ridgway (1902:268) recorded Icterus
wagleri from Saltillo. Hellmayr (1937:122-123) referred this record of
Wagler's Oriole to I. w. wagleri.

*lcterus bullockii bullockii (Swainson). — This subspecies of Bullock's Oriole
was listed as breeding by Miller, Friedmann, Griscom, and Moore (1957:282)
at Monclova on May 12-19.

Euphagus cyanocephalus (Wagler). — Brewer's Blackbird is a common mi-
grant in Coahuila. Miller (1955a: 174) found a few as migrants in the
Sierra del Carmen on April 27. Burleigh and Lowery (1942:205) remarked
that "this blackbird was characteristically a bird of the towms and villages,
the scattered flocks being invariably seen feeding in the streets and near the
houses." They ( loc. cit. ) obtained three specimens at Diamante Valley on
April 23 and remarked that tlie departure of these birds in spring was "ex-
tremely late."

*Cassidix mexicanus prosopidicola Lowery. — Specimens examined: total 3:

9 32893 from Parras, July 4, 1955, weight, 98.8 gms.; and $ $ 35418-35419
from Torreon, January 8, 1954.

This subspecies of Boat-tailed Grackle has been recorded from several
localities in Coahuila. In southeastern Coahuila, Burleigh and Lowery ( 1942:
205-206) noted this grackle "somewhat local in its distribution; it was found
in cultivated fields about the towns and villages." These authors noted it

500 University of Kansas Publs., Mus. Nat. Hist.

at Saltillo, the Chorro del Agua, and "occasionally in the open valley south
of Diamante Pass" and obtained specimens from "near Saltillo" and Diamante
Valley. On August 18 Amadon and Phillips (1947:579) found C. m.
prosopidicola at Las Delicias where "a fledged young was noticed begging
for food. . . ." Lowery (1938:4) recorded one specimen of C. m.
prosopidicola from Monclova. Findley saw Boat-tailed Crackles 2 mi. W
Jimenez on June 19, 1952, and 2 mi. S and 11 mi. E Nava on June 15, 1952.

The distribution and intergradation of Boat-tailed Crackles in Coahuila is
presently poorly understood. C. m. prosopidicola from southeastern Coahuila
may approach C. m. mcxicanus, and there is probable intergradation of
prosopidicola with rnonsoni in northwestern Coahuila (Phillips, 1950:78).

Molothrus ater ater (Boddaert). — Specimen examined: one, $ 31513, from
21 mi. S, 11 mi. E AustraUa, 4400 ft, May 3, 1954, measurements: wing,
102 mm.; tail, 64 mm.; tarsus, 26 mm.; culmen, 17 mm.

This subspecies of the Brown-headed Cowbird is uncommon in Coahuila.
Amadon and Phillips (1947:579) took an adult male and a juvenal female
M. a. ater at Las Delicias on August 15, both of which were considered early
migrants. Dickerman obtained No. 31513 from a flock of eight cowbirds.
Although the measurements of this specimen agree equally well with those of
M. a. ater and M. a. artemisiae (Crinnell, 1909:275-281), the shape of the
bill most closely resembles that of ater. Crinnell ( 1909:278) said that "ater has
a tumid bill, broad and high at [the] base with [a] conspicuously arched
culmen" whereas "artemisiae has a longer and relatively much slenderer bill,
vertically shallow at [the] base and laterally compressed, with the culmen
in its greater portion straight or even slightly depressed." The size of the
ovary (8x4 mm.) of No. 31513 and the date (May 3) on which it was
obtained suggest that this individual was a late migrant, still south of the
breeding range of M. a. ater.

Molothrus ater artemisiae Crinnell. — This subspecies of the Brown-headed
Cowbird is an uncommon migrant in Coahuila. Amadon and PhiUips ( 1947:
579) obtained, at Las Delicias, a juvenal male on August 15 and an adult
male on August 17.

*Molothrus ater obscurus (Cmelin). — Specimens examined: total 18: $ $
32112-32115 from 12 mi. N, 12 mi. W Jimenez, 850 ft, June 18 and 19, 1952;
$ $ 32108-32111, $ 32116, and 9 9 32117-32120 from 2 mi. W Jimenez,
850 ft., June 20, 1952; 9 32491 from Las Margaritas, August 4, 1955; $ 31511
and 9 31510 from 16 mi. E, 18 mi. N Ocampo, Mav 8 and 7, 1952; and S S
35409-35410 (skeletons only) from 4 mi. N San Isidro, May 11, 1954.

This subspecies of the Brown-headed Cowbird is common in Coahuila and
breeds there. Amadon and Phillips (1947:579) suggested that M. a. obscurus
breeds at Las Delicias. Burleigh and Lowery (1942:206) saw the Brown-
headed Cowbird in "small numbers on the outskirts of Saltillo. . , ."

Measurements of the adult males that I have examined are: wing, 101.1
mm. (97-106); tail, 66.5 mm. (62-69); tarsus, 25.6 mm. (24.5-28); culmen,
17.3 mm. (16-18.5). Measurements of the adult females that I have examined
are: wing, 92.3 mm. (90-97); tail, 60.1 mm. (56.5-62.5); tarsus, 23.5 mm.
(22.5-24); culmen, 14.4 mm. (14-15). The sizes of the testes of three of the
males (6-7 mm. long) and of the largest ova of four of the females (6-9

Birds From Coahuila, Mexico 501

mm. in diameter) indicate breeding by this subspecies in Coahuila, as does
the small size of one of the juvenal males (tail, 33.5 mm. long).

Piranga ludoviciana (Wilson). — In Coahuila the Western Tanager occurs
fairly commonly as a migrant. There are no records of it breeding in the
State. Miller (1955a: 174) remarked that Marsh took a migrant Western
Tanager at Jardin del Sur in the Sierra del Cannen on September 7. Amadon
and Phillips (1947:579) took an adult male Western Tanager at Las Delicias
on August 12. Miller, Friedmann, Griscom, and Moore (1957:305) also re-
corded this tanager from 12 mi. W Saltillo. Dickerman saw Western Tanagers
in the Sierra del Pino on May 12, 1954, and 16 mi. E and 18 mi. N Ocampo
on May 7, 1954.

*Piranga flava dextra Bangs. — Specimen examined: one, $ 31526, from
Sierra del Pino (=5 mi. S, 3 mi. W Acebuches), May 12, 1954, weight, 41 gms.

Miller (1955a: 174) found this subspecies of the Hepatic Tanager "in the
pine-oak belt at 7000 feet on April 12 [in the Sierra del Carmen], when a
male was seen and a female taken." No. 31526 was with a female when taken;
this male was not fat and its testes were not enlarged. The size of the wing
(105.5 mm.) of No. 31526 represents the extreme maximum in this sub-
species.

Piranga flava hepatica (Swainson). — Miller, Friedmann, Griscom, and
Moore (1957:303) stated that P. /. hepatica is found in northwestern and
central Arizona and southwestern New Mexico south into the highlands of
Mexico, west of the Sierra Madre Oriental, to Oaxaca and in winter and
migration extends eastward and south to Chiapas. These authors remarked
also that P. /. dextra occurs in the mountains east of the continental divide in
New Mexico and western Texas south through eastern Mexico to Chiapas.
Specimens of the Hepatic Tanager from Coahuila in winter might well be
either P. /. hepatic or P. /. dextra. Miller, Friedmann, Griscom, and Moore
( loc. cit. ) recorded migrants of P. /. hepatica from the Sierra de Guadalupe
on April 24-27.

*Piranga rubra rubra (Linnaeus). — Specimens examined: total 6: S 32129,
S 32132, and 9 32133 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 18
and 19, 1952; $ 32130 from 2 mi. W Jimenez, 850 ft, June 20, 1952; and
$ 32131 and 9 32134 from 2 mi. S, 11 mi. E Nava, June 15, 1952.

This Summer Tanager occurs in northeastern Coahuila. The specimens
from 12 mi. N and 12 mi. W Jimenez, 2 mi. W Jimenez, and 2 mi. S and 11
mi. E Nava are typical representatives of P. r. rubra. The large testes ( 12 mm. )
of No. 32129 and the well-developed brood patch of No. 32134 are evidence
of breeding by this subspecies in the State. Heretofore this subspecies has
not been recorded from Coahuila.

* Piranga rubra cooperi Ridgway. — Specimens examined: total 4: $ $
32828-32829, $ 32831, and $ 32830 from Parras, July 4, 1955.

This subspecies of Summer Tanager seems to occur throughout Coahuila
except in the northeastern section of the State. Miller, Friedmann, Griscom,
and Moore (1957:302) hsted P. r. cooperi from Sabinas and Sierra de Guada-
lupe. Miller (1955a: 174) saw a Summer Tanager in a canyon in the Ser-
ranias de Burros, about 40 miles east of the Sierra del Cannen on April 28.

502 University of Kansas Publs., Mus. Nat. Hist.

Although there are no other records of P. r. cooperi from northwestern Coahuila,
Van Tyne and Sutton (1937:96) recorded this tanager as a common nesting
species in Brewster County, Texas, in cottonwood, mesquite, or willow trees.
I suspect that P. r. cooperi is a common nesting bird in northwestern Coahuila
as well.

Nos. 32828-32831 approach P. r. rubra. The measurements of No. 32829
are: wing, 98 mm.; tail, 80 mm.; the measurements of No. 32831 are: wing,
98 mm.; tail, 79 mm. The specimens of P. r. cooperi from Parras are some-
what small and seemingly approach P. r. rubra. The sizes of the testes (8x5
mm.; 9x5 mm.) of Nos. 32829 and 32831, respectively, and the size of the
largest ovum (4x4 mm.) of No. 32830 indicate breeding by this subspecies in
southern Coahuila, as does the presence of No. 32828, a juvenal male.

* Richmondena cardinalis canicaudus ( Chapman ) . — Specimens examined:
total 3: $ 31099 from 1 mi. N Boquillas, 700 ft., March 8, 1952, weight, 45.3
gms.; $ 32135 from 2 mi. W Jimenez, 850 ft., June 20, 1952; and $ 32136
from 2 mi. S, 3 mi. E San Juan de Sabinas, June 22, 1952.

In Coahuila the Cardinal is common. Miller (1955a: 174) found it singing
in the Boquillas drainage of the Sierra del Carmen at 4800 feet, and gave evi-
dence that the Cardinal breeds there. Sutton and Burleigh (1939a:43) found
the species to be common in the low country "east of Saltillo." Hellmayr
(1938:69) recorded R. c. canicaudus from Sabinas. The sizes of the testes
(9 mm.; 7x4 mm.) of Nos. 32135-32136 indicate breeding by this subspecies
in northeastern Coahuila.

*Pyrrhuloxia sinuata sinuata (Bonaparte). — Specimens examined: total 4:
S 31100 from 10 mi. S, 5 mi. E Boquillas, 1500 ft., March 4, 1952, weight,
37.3 gms.; $ 32137 from 5 mi. N, 19 mi. W Cuatro Cienegas, 3250 ft, July
5, 1952; $ 35403 (skeleton only) from San Marcos, May 5, 1954; and S 30234
from 3 mi. SE Torre6n, 3800 ft., January 12, 1951.

In Coahuila, the Pyrrhuloxia is common. Hellmayr (1938:76) Hsted it
from Sabinas. Ridgway (1901:628) recorded P. s. texana (=sinuata) from
La Ventura. Burleigh and Lowery (1942:206) found the species "only in the
open desert country west of Saltillo where, on April 22, several pairs were
seen in a small arroyo." Amadon and Phillips (1947:579) took an im-
mature P. s. sinuata at Las DeUcias; Sutton and Burleigh ( 1939a:43-44)
found this subspecies fairly common in the San Pedro district on January 29
and 30. Miller, Friedmann, Griscom, and Moore (1957:330) recorded
breeding by P. s. sinuata at Hipolito on July 2. The size of the testes ( 8 mm. )
of No. 32137 indicates breeding in central Coahuila.

* Pheucticus melanocephalus melanocephalus ( Swainson ) . — Specimen ex-
amined: one, $ 31664, from Sierra Guadalupe (=10 mi. S, 5 mi. W General
Ccpeda), 7500 ft., April 30, 1953, measurements: wing, 105.5 mm.; tail, 78
mm.; culmen, 19 mm.; weight, 48 gms.

Miller (1955a: 174) reported that the Black-headed Grosbeak first ap-
peared in the Sierra del Carmen on April 13 and was soon seen patrolling
territories. He remarked that these specimens from the Sierra del Carmen
conformed adequately with the rather poorly differentiated race P. m. melano-
cephalus and stated that Marsh took an immature male at Jardin del Sur on
September 7. Oberholser (1919b:416) listed Ilcdymclcs melanocephalus
papago i=P. m. melanocephalus) from Sierra Guadalupe on April 27.

Birds From Coahuila, Mexico 503

Diclcerman saw Black-headed Grosbeaks in the Sierra del Pino on May 12,
1954, and 16 mi. E and 18 mi. N Ocampo on May 7, 1954. The size of No.
31664 represents the characters of P. m. melanocephalus as presented by
Oberholser (1919b:413). No specimen of P. m. melanocephalus from Coa-
huila, to my knowledge, approaches P. m. maculatus. The size of the testes
(7x5 mm.) of No. 31664 suggests breeding by this subspecies in the Sierra
Guadalupe.

*Guiraca caerulea interfusa Dwight and Griscom. — Specimens examined:
total 2: $ 32138 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 18, 1952;
and 5 32139 from 2 mi. W Jimenez, 850 ft., June 21, 1952.

Burleigh and Lowery (1942:206) recorded this Blue Grosbeak from
"about twenty miles west of Saltillo" on April 22. Miller (1955a: 174) stated
that Marsh obtained a male G. c. interfusa at Vivoras Spring on August 3.
Miller, Friedmann, Griscom, and Moore (1957:334) remarked that the Blue
Grosbeak breeds in the northern and eastern sections and reported G. c.
interfusa from Hipohto on July 2. Amadon and Phillips (1947:580) re-
ported examining material from Sabinas referable to G. c. interfusa. Dicker-
man saw Blue Grosbeaks 4 mi. N San Isidro on May 10, 1954. Findley saw
Blue Grosbeaks 4 mi. W Jimenez on June 19, 1952, and 2 mi. S and 3 mi. E
San Juan de Sabinas on June 22, 1952. Nos. 32138-32139 are typical rep-
resentatives of G. c. interfusa. The size of the testes ( 12 mm. ) of No. 32138,
the size of the largest ovum (2 mm.) of No. 32139, and the dates (June 18, 21)
on which they were collected are evidence of breeding by this subspecies.

**Guiraca caerulea eurhyncha Coues. — Amadon and Phillips (1947:580)
obtained an adult male of this Blue Grosbeak from Las Delicias on August 12.
This subspecies, according to Miller, Friedmann, Griscom, and Moore (1957:
334), is resident at middle and lower elevations through most of central and
southern Mexico. Except for the occurrence of intergrades of G. c. interfusa,
caerulea, and eurhyncha in southern Nuevo Leon and Tamauhpas (Miller,
Friedmann, Griscom, and Moore, 1957:335), the record from Las Delicias
represents the northern limit of the range of the subspecies eurhyncha.

* Passerina cyanea (Linnaeus). — Specimen examined: one, $ 31544, from
San Marcos, May 5, 1954.

The Indigo Bunting is rare in Coahuila. Van Hoose (1955:303) reported
that No. 31544 seems to provide the first record of the species in the State.
The Indigo Bunting is a summer resident in southwestern Oklahoma and
southeastern Texas (Miller, Friedmann, Griscom, and Moore, 1957:336).
No. 31544 seemingly represents a large extension in the summer range of
the Indigo Bunting. Van Hoose {loc. cit.) stated that No. 31544 was ob-
served chasing another male, and the pursuer in turn was followed by a
female; he thought that the territorial behavior of this bird suggested
breeding.

* Passer ina versicolor versicolor (Bonaparte). — Specimens examined: total
2: $ 35414 from 6 mi. N, 2 mi. W Castillon, 3750 ft., June 30, 1953, weight,
8 gms.; and $ 35415 from 5 mi. S Castillon, 4150 ft., June 28, 1953.

Although the Varied Bunting has been recorded only from northwestern
Coahuila, I suspect that this bird is locally common throughout most of the

504 Unrtrsity of Kansas Publs., Mus. Nat. Hist.

State. Miller (1955a: 174) stated that the habitat of this species consisted of
catclaw-covered bottom lands at the base of the Sierra del Carmen at 4700
feet. Miller's records indicate incipient breeding by P. v. versicolor in the
Sierra del Carmen on April 26. The size of the testes (11 mm.) of No. 35415
and the dates (June 28, 30) on which Nos. 35414-35415 were collected are
strong evidence of breeding by the Varied Bunting 6 mi. N and 2 mi. W
CastiUon and 5 mi. S Castillon,

* Passerina ciris pallidior Meams. — Specimens examined: total 6: $ 32141
and 9 32142 from 2 mi. S, 11 mi. E Nava, 810 ft., June 15 and 16, 1952;
$ 35416 from 6 mi. N, 2 mi. W Castillon, 3750 ft., June 29, 1953, weight,
15 gms.; $ 32140 from 2 mi. S, 3 mi. E San Juan de Sabinas, June 22, 1952;
<5 31547 from 16 mi. N, 14 mi. E Ocampo, May 10, 1954, weight, 16 gms.;
and $ 31546 from San Marcos, May 5, 1954.

The Painted Bunting is a common summer resident in Coahuila. Marsh
and Stevenson (1938:287) wrote that Painted Buntings were common in sum-
mer in the foothills of the Sierra del Carmen, and they took a male at Piedra
Blanca on July 25 and a female at Jardin del Sur on September 1. Amadon
and Phillips (1947:580) remarked that two immature Painted Buntings, "ap-
parently migrants, were taken at Las Delicias on August 10 and 11." Miller,
Friedmann, Griscom, and Moore (1957:339) recorded breeding by P. c. palli-
dior at Hipolito, 4200 feet, on July 1. Findley saw Painted Buntings 2 mi.
S and 11 mi. E Nava on June 15, 1952. Dickerman saw Painted Buntings 4
mi. N San Isidro on May 10, 1954. Hardy saw Painted Buntings at Parras
on July 4, 1955.

All the University of Kansas specimens are typical of the larger subspecies
pallidior. The dates (May 5, 10; June 15, 16, 22, 29) on which these speci-
mens were collected, the sizes of the testes (9x6 mm.; 6x5 mm.; 7x3 mm.) of
Nos. 32140, 31547, and 31546, respectively, and the size of the largest ovum
(2.5 mm.) of No. 32142 indicate breeding by P. c. pallidior in Coahuila.

Carpodacus cassinii Baird. — Cassin's Finch is an uncommon winter migrant
in Coahuila. Miller, Friedmann, Griscom, and Moore (1957:314) listed C.
cassinii from Sierra Guadalupe.

* Carpodacus mexicanus potosinus Griscom. — Specimens examined: total 8:
$ 35417 from 13 mi. S, 5 mi. W Castillon, 4000 ft., June 28, 1953, testes, 6
mm., weight, 19 gms.; ? 35411 (skeleton only) from Sierra del Pino (=5 mi.
W, 3 mi. S Accbuches), 6200 ft.. May 14, 1954; $ 32145 from 5 mi. N, 19
mi. W Cuatro Cionegas, 3250 ft., July 5, 1952; $ 32144 and 9 32147 from 3
mi. S, 3 mi. E Bella Union, 6750 ft., June 27, 1952, $ gonads, 7 mm.; $ 32143
and 9 32146 from 7 mi. S, 4 mi. E Bella Union, 7200 ft., Jime 24, 1952; and
$ 31675 from the west foot of Pico de Jimulco, 5000 ft., April 4, 1953, weight,
20 gms.

The House Finch is common in Coahuila. Miller (1955a: 174) reported
a sparse population of this species in the foothills adjoining Boquillas Canyon
at 4800 feet of the Sierra del Carmen and observed young just out of the
nest on April 25. He remarked also that specimens of the House Finch from
the Sierra del Carmen seem to show no intergradation toward frontalis. Bur-
leigh and Lowcry (1942:206) noted C. m. potosinus at "Saltillo, in the desert
country west of there, at the Chorro del Agua, and in the open valley south
of Diamante Pass." Miller, Friedmann, Griscom, and Moore (1957:316)
listed breeding by C. m. potosinus at El Diamante on July 6. Dickerman

Birds From Coahuila, Mexico 505

also saw the House Finch at San Marcos on May 4, 1954, and Hardy saw it
at Parras on July 4, 1955. No. 32147 had a distinct brood patch; the largest
ovum of No. 32146 was 7 mm. in diameter. No. 32145 was a juvenal male.
The University of Kansas specimens agree well with descriptions of C. m.
potosimis as given by Moore (1939:195). No approach toward C. m. frontalis,
centralis, or nigrescens is exhibited by any of these specimens. No. 31675,
from Pico de Jimulco in southwestern Coahuila, is paler above and below than
any other specimens of C. m. potosinus. Also the crown of No. 31675 is suffused
with more red than in typical representatives of C. m. potosinus.

** Spinus pinus pinus (Wilson). — Specimen examined: one, sex ? 33219
(skeleton only) from 13 mi. E San Antonio de las Alazanas, 9950 ft.,
July 6, 1955.

On April 5, 7, and 21, Miller (1955a: 175) recorded winter visitant or
vagrant flocks of Pine Siskins in the Sierra del Carmen. Miller, Friedmann,
Griscom, and Moore (1957:318) hsted S. p. pinus south to Sierra Guadalupe.
Dickerman saw Pine Siskins 13 mi. E San Antonio de las Alazanas on April 10,
1954.

Spinus pinus macropterus (Bonaparte). — Miller, Friedmann, Griscom, and
Moore (1957:319) recorded a vagrant S. p. macropterus from 50 mi. S
Monclova, 2850 feet, on November 9.

Spinus tristis pallidus Mearns. — Specimen examined: one, 9 31101, from
Fortin (=33 mi. N, 8 mi. W San Geronimo), 3300 ft., March 28, 1952,
weight, 11.5 gms.

The American Goldfinch is an uncommon migrant or winter visitant in
Coahuila. Hellmayr (1938:296) recorded S. t. pallidus from Sabinas. Fortin
and Sabinas are the only places in Coahuila where S. t. pallidus has been col-
lected. No. 31101 is a typical representative of S. t. pallidus.

** Spinus psaltria psaltria (Say). — Specimens examined: total 7: $$
32148-32149 and $ 32151 from 12 mi. N, 12 mi. W Jimenez, 850 ft., June 19,
1952; $ 32150 from 2 mi. W Jimenez, June 20, 1952; $ 33220 (skeleton
only) from Parras, July 4, 1955; and S 32939 and $ 32940 from Mesa de las
Tablas, July 7, 1955, weights, 9.5, 11 gms.

In Coahuila, the Lesser Goldfinch seems to be common. Although Miller
(1955a: 175) did not find it in the Sierra del Carmen, he reported that
Marsh took a specimen on August 22 in Chuperosa Canyon that was "pre-
sumed to" be S. p. psaltria. Burleigh and Lowery (1942:206) observed the
Lesser Goldfinch "on the outskirts of Saltillo in an orchard on April 20."

Nos. 32148-32151 and 32940 were typical for the subspecies psaltria.
A partial albino (32939), which was obtained from a pine-oak -wheat field
edge, has upper parts that lack the black coloring of typical representatives of
S. p. psaltria. Instead the crown and back of No. 32939 is yellow, re-
sembling the color of its underparts, the wing coverts are white, and its
primaries are black with white edgings.

* Atlapetes pileatus dilutus Ridgway. — Specimens examined: total 2: $
32942 and ? 32943 from 13 mi. E San Antonio de las Alazanas, 9950 ft.,
July 6, 1955.

The Rufous-capped Atlapetes occurs uncommonly in southeastern Coahuila.
The male and female A. p. dilutus from 13 mi. E San Antonio de las Alazanas

506 Unr'ersity of Kansas Publs., Mus. Nat. Hist.

represent the first records of occurrence of this subspecies in the State. The
smaller size, grayer upper parts, and pale yellow color of the underparts of
Nos. 32942-32943 characterize the subspecies dilutus. The size of the testes
(7x6 mm.) of No. 32942 and the date (July 6) on which both specimens were
collected indicate breeding by this species in Coahuila.

*Arremonops rufivirgata rufivirgata (Lawrence). — Specimens examined:
total 2: <5 32152 and 2 32153 from 2 mi. S, 3 mi. E San Juan de Sabinas,
June 22 and 23, 1952.

The Olive Sparrow is uncommon in Coahuila. Miller, Friedmann, Griscom,
and Moore (1957:349) listed A. r. rufivirgata from Sabinas on February 10.
Sabinas and soutlieast of San Juan de Sabinas seem to be the only localities in
Coahuila where the Olive Sparrow has been collected and also are at the
westernmost extremity of range of this species. The size of the testes (9x7
mm.) of No. 32152 and the dates of collection indicate breeding by the Ohve
Sparrow in Coahuila.

Chlorura chlorura (Audubon). — Specimen eocamined: one, $ 30238, from
1 mi. N San Lorenzo, 4200 ft., February 5, 1951.

The Green-tailed Towhee is a common migrant and winter visitant in
Coahuila; the species has been found at several localities. Miller (1955a: 175)
noted several migrants "each day in the last week of April ... at the
mouth of Boquillas Canyon" of the Sierra del Carmen. Burleigh and
Lowery (1942:207) noted the species "in an arroyo in the open desert
country about twenty miles west of Saltillo on April 22." Miller, Friedmann,
Griscom, and Moore (1957:351) hsted the Green-tailed Towhee from 12 mi.
W Saltillo on September 28 and from Sabinas.

* Piplio erythrophthalmus gaigei Van Tyne and Sutton. — Specimens ex-
amined: total 3: $ 31102 from Fortin (=33 mi. N, 8 mi. W San Geronimo),
3300 ft., March 28, 1952, weight, 38.8 gms.; $ 35412 (skeleton only) from
Sierra del Pino (=5 mi. W, 3 mi. S Acebuches), 6200 ft.. May 15, 1954,
testes enlarged, weight, 34 gms.; and $ 31593 from 17 mi. S Ocampo, 5300
ft., April 7, 1954, weight, 38 gms.

The Rufous-sided Towhee is locally common in Coahuila; P. e. gaigei is
present in northern Coahuila. Miller (1955a: 175) remarked that "between
6800 and 7500 feet these towhees were sparsely distributed in areas of
scattered low ceanothus and hawthorne, chiefly in canyon bottoms, but also
on slopes where ceanothus was intermingled with downed timber and young
pines." He {loc. cit.) indicated that the adult obtained by Marsh from
Vivoras Canyon on August 25 was P. e. gaigei rather than P. maculatus mon-
tanus (see also Sibley, 1950:127). Dickerman saw Rufous-sided Towhees
in the Sierra de la Madera on December 13, 1953. I have referred Nos. 31 102
and 31593 to gaigei although both are close to orientalis. The size of the
testes (14x8 mm.) of No. 31593 suggests breeding.

* Piplio erythrophthalmus orientalis Sibley. — Specimens examined: total 3:

S 32154 from 7 mi. S, 4 mi. E Bella Uni6n, 7200 ft., June 25, 1952; S 33223
(skeleton only) from 13 mi. E San Antonio de las Alazanas, 9950 ft., July 6,
1955; and $ 31630 from Mesa de Tablas, 8600 ft., January 15, 1954, weight,
46 gms.

Birds From Coahuila, Mexico 507

This subspecies of the Rufous-sided Towhee occurs in southeastern Coahuila.
Burleigh and Lowery (1942:207) stated that it "was Hmited in its distribu-
tion to the mountain sides . . . [and was] noted in the arroyos at the
base of the mountains and from there up to about 8,000 feet." These authors
identified their specimens from Diamante Pass and from Saltillo as Pipilo
maculatus gaigei. Sibley (1950:129) reidentified them, as well as a series
from Sierra de Guadalupe, as P. e. orientalis. The size of the testes ( 12x7 mm. )
of No. 32154 as well as the date (June 25) on which it was obtained suggests
breeding by the Rufous-sided Towhee in southeastern Coahuila.

* Pipilo fuscus potosinus Ridgway. — Specimens examined: total 3: $ 32155
from 7 mi. S, 4 mi. E Bella Union, 7200 ft., June 25, 1952; S 31676 from the
west foot of Pico de Jimulco, 5000 ft., April 5, 1953, weight, 45 gms.; and sex ?
29560 from 7 mi. S, 1 mi. E Gomez Farias, 6500 ft., November 19, 1949,
weight, 46.5 gms.

The subspecies potosinus has been recorded from several localities in Coa-
huila. Davis (1951:70) Hsted the following localities in the State from which
P. f. potosinus has been collected: Muralla, San Lazaro Mountains, 50 mi. S
Monclova, 2850 ft., Saltillo; Saltillo (Chorro de Agua); 19 mi. W Saltillo;
Cresta Blanca, 12 mi. W Saltillo, 5500 ft.; Diamante Pass, 11 mi. S Saltillo,
6000-8000 ft.; and Cameros. Miller, Friedmann, Griscom, and Moore (1957:
358) recorded a "small juvenile" from El Diamante on July 5. Sutton and
Burleigh ( 1939a :45) recorded P. f. texanus from Diamante Pass on March 6.
I suspect that Davis (op. cit.) reidentified the specimen concerned from Dia-
mante Pass as P. /. potosinus. Burleigh and Lowery (1942:207) indicated
that P. f. potosinus "was fairly common and of general distribution in the area,
occurring both on the arid plateau about Saltillo and on the mountain sides
up to an elevation of about 8,000 feet." Nos. 32155, 31676, and 29560 are
typical for P. /. potosinus. The size of the testes ( 14x7 mm. ) of No. 32155
suggests breeding by the Brown Towhee in southeastern Coahuila.

* Pipilo fuscus texanus van Rossem. — Specimen examined: one, 9 31103,
from 10 mi. S, 5 mi. E Boquillas, 1500 ft., March 5, 1952, weight, 46.9 gms.

This subspecies of the Brown Towhee occurs in northwestern Coahuila
south through the Sierra del Carmen. Miller (1955a: 176) reported that his
series of Brown Towhees from the Sierra del Carmen agreed satisfactorily
with texanus although revealing some sign of intergradation with potosinus.
Davis (1951:70) thought that P. /. potosinus is present in northern as well as
southern Coahuila. Miller (1955a: 176), however, remarked that P. /. texanus
is more characteristic of the population of Brown Towhees of northwestern
Coahuila. He indicated that a single juvenile taken by Marsh on August 28
from Jardin del Sur and allocated to P. f. potosinus by Davis probably is P. /.
texanus.

Calamospiza melanocorys Stejneger. — Specimen examined: one, $ 30239,
from 10 mi. E Torreon, 3700 ft., January 9, 1951.

The Lark Bunting is an uncommon winter visitant in Coahuila. Other
than the present specimen, the only record of the Lark Bunting in Coahuila
is that of Burleigh and Lowery (1942:207), who reported a small flock of
this species from which several specimens were collected "on April 20 in a field
on the edge of Saltillo."

5—7905

508 Uni\'ersity of Kansas Publs., Mus. Nat. Hist.

Passerculus sandwichensis oblitus Peters and Griscom. — Miller, Friedmann,
Griscom, and Moore (1957:364) recorded P. s. oblitus from Sabinas on Feb-
ruary 25 and March 18 and 24. These records represent sparse winter visi-
tants to Coahuila.

Passerculus sandwichensis brooksi Bishop. — Burleigh and Lowery (1942:
208) recorded P. s. brooksi from Diamante Pass in April; Miller, Friedmann,
Griscom, and Moore (1957:363) remarked that this specimen is perhaps best
regarded as a variant of one of the races normally wintering in this area.

Passerculus sandwichensis anthinus Bonaparte. — Specimens examined: total
3: S 31104 from 10 mi. E Hacienda La Mariposa, 2000 ft., March 30, 1952,
weight 15 gms.; 9 31105 from 8 mi. N, 4 mi. W Muzquiz, 1800 ft, April 1,
1952, weight, 19.0 gms.; and ? 31594 from 17 mi. S Ocampo, 5300 ft., April
7, 1954, weight, 16 gms.

Burleigh and Lowery (1942:208) recorded P. s. anthinus from Diamante
Pass. This subspecies is not uncommon in Coahuila. The University of
Kansas specimens showed no indication of breeding.

Passerculus sandwichensis nevadensis Grinnell. — This subspecies of the
Savannah Sparrow is uncommon in Coahuila; Hellmayr (1938:490) listed
one specimen from Sabinas.

Passerculus sandwichensis brunnescens (Butler). — Burleigh and Lowery
(1942:208) recorded this subspecies of the Savannah Sparrow from Diamante
Pass in April,

Ammodramus savannarum perpallidus (Coues). — Specimen examined: one,
9 31562, from 3 mi. N, 4 mi. E San Francisco (=25 mi. N Ocampo), 4850
ft., May 16, 1954, weight, 15 gms.

The Grasshopper Sparrow is an uncommon spring and possibly fall mi-
grant in Coahuila. Miller, Friedmann, Griscom, and Moore (1957:367) re-
corded A. s. perpallidus from Sabinas on March 12. No. 31562, which was
obtained in a yucca and acacia association, had little fat.

Ammodramus bairdii (Audubon). — Miller, Friedmann, Griscom, and
Moore (1957:368) remarked that Baird's Sparrow is a rare winter visitant to
the northern states of Mexico and recorded A. bairdii from Saltillo on May 8.

Pooecetes gramineus confinis Baird. — The Vesper Sparrow seems to be an
uncommon winter visitant in Coahuila. Miller (1955a: 176) found P. g.
confinis "on two occasions in the grass of the dry cienega at tlie head of Corte
Madera Canyon at 7500 feet" on April 9 and 14 in the Sierra del Carmen.
In April, Burleigh and Lowery (1942:208) found P. g. confinis only in Dia-
mante Valley where this sparrow "appeared to be quite uncommon." Sutton
and Burleigh (1939a:45) took a male P. g. confinis at San Pedro on Jan-
uary 29.

* Chondestes grammacus strigatus Swainson. — Specimen examined: one, $
32156, from 8 mi. N, 2 mi. W Piedras Negras, June 18, 1952.

The Lark Sparrow is not uncommon in Coahuila. Miller (1955a:176)
saw one male, "apparently on a breeding territory, on April 27 in an open,
rather barren desert flat adjoining the lower part of Boquillas wash at 4600
feet." He reported that Marsh took a young of the year, still largely in

Birds From Coahuila, Mexico 509

Juvenal plumage, on September 6 in the Sierra del Carmen. Amadon and
Phillips (1947:580) remarked that Lark Sparrows were common "about Las
DeHcias" after August 18. Findley saw Lark Sparrows 2 mi. W Jimenez on
June 19, 1952, and 2 mi. S and 11 mi. E Nava on June 15, 1952. Dicker-
man saw Lark Sparrows at San Marcos on May 4, 1954. The pale and nar-
rowly streaked upperparts of No. 32156 are typical for C. g. strigatus. The
size of the testes (9x4 mm.) of No. 32156 and the date (June 18) on which
it was obtained suggest breeding by the Lark Sparrow in northeastern Coa-
huila.

* Aimophila ruficeps tenuirostris Burleigh and Lowery. — This subspecies of
the Rufous-crowned Sparrow is resident in the northern part of Coahuila.
Miller (1955a: 176) remarked that the species "ranged up to 7000 feet on
open south-facing slopes within the oak belt" of the Sierra del Carmen. Speci-
mens collected by him showed no approach to boucardi of southern Mexico
and seem to be closest to tenuirostris. Miller referred the specimen that
Marsh and Stevenson (1938:287) took on August 22 in Chuperosa Canyon
to tenuirostris rather than boucardi. Miller, Friedmann, Griscom, and Moore
(1957:376) listed A. r. tenuirostris from 50 mi. S Monclova on November
8 and 10.

* Aimophila ruficeps boucardi (Sclater). — This subspecies of the Rufous-
crowned Sparrow is common in southern Coahuila. Burleigh and Lowery
(1942:208) indicated that A. r. boucardi was common throughout the moun-
tainous areas and to some extent in the arroyos of the open desert country of
southeastern Coahuila. Sutton and Burleigh (1939a:46) took a female A. r.
boucardi "near Diamante Pass." Ridgway (1901:252) listed A. r. boucardi
from Carneros. A. r. tenuirostris and boucardi seem to intergrade in central
and even southern Coahuila. Miller, Friedmann, Griscom, and Moore (1957:
376) recorded intennediate populations of the Rufous-crowned Sparrow from
12 mi. W Saltillo.

* Aimophila cassinii (Woodhouse). — Specimens examined: total 2: $
32157 from 9 mi. S, 11 mi. E Sabinas, June 14, 1952; and $ 32158 from
18 mi. S, 14 mi. E Tanque Alvarez, 4000 ft., July 6, 1952.

Cassin's Sparrow seems to be common in Coahuila. The AOU Check-list
Committee (1957:603) listed A. cassinii from 10 mi. E Saltillo. Miller, Fried-
mann, Griscom, and Moore (1957:378-379) recorded Cassin's Sparrow from
Sabinas, on April 25; from 25 mi. SW Monclova, on November 20; from 12
mi. W Saltillo, on September 28; and from 10 mi. NE Saltillo, on July 3.
These authors stated that Cassin's Sparrow was breeding 10 mi. NE Saltillo.
The sizes of the testes (5x3 mm.; 7x5 mm.) of Nos. 32157-32158, respectively,
and the dates (June 14, July 6) on which they were obtained are additional
evidence of breeding by Cassin's Sparrow in Coahuila.

* Amphispiza bilineata bilineata (Cassin). — Specimens examined: total 5:
$ 32159 and 9 32163 from 2 mi. W Jimenez, 850 ft., June 20, 1952; $ 32160
from 8 mi. N, 2 mi. W Piedras Negras, June 18, 1952; S 32161 and sex ?
32162 from 5 mi. N, 19 mi. W Cuatro Cienegas, 3250 ft., July 5, 1952.

The Black-throated Sparrow is common in Coahuila. Typical representa-
tives of A. b. bilineata occur in eastern Coahuila. The center of the State is
occupied by intergrades between A. b. bilineata, opuntia, and grisea. Miller,

510 University of Kansas Publs., Mus. Nat. Hist.

Friedmann, Griscom, and Moore (1957:381) reported A. b. bilineata from the
"Saltillo area." Hellmayr (1938:539) recorded A. b. bilineata from Sabinas.
The sizes ( 13, 14 mm. ) of the white spot on the lateral tail feathers of Nos.
32163 and 32160, respectively, suggest A. b. bilineata. The short wing
(61-64.5 mm.) and the lighter and browner color of the backs of Nos. 32163,
32160, and 32161 are suggestive of A. b. bilineata. Nevertheless, the size
(8 mm.) of the white spot on the lateral tail feather of No. 32161 indicates
intergradation with A. b. opuntia.

The sizes of the testes (6x4 mm.; 8 mm.) of Nos. 32160 and 32161, the
size of the largest ovum (2 mm. in diameter) of No. 32163, and the presence
of the juveniles from 2 mi. W Jimenez and 5 mi. N and 19 mi. W Cuatro
Cienegas indicate breeding by A. b. bilineata in Coahuila.

* Amphispiza bilineata opuntia Burleigh and Lowery. — Specimens examined:
total 2: 9 31106 from 10 mi. S, 5 mi. E Boquillas, 1500 ft., March 3, 1952;
weight, 12.3 gms.; and $ 31108 from 35 mi. S, 14 mi. E Boquillas, 2350 ft.,
March 12, 1952.

This subspecies of the Black-throated Sparrow occurs in northwestern Coa-
huila. Miller (1955a: 176) stated that the Black-tliroated Sparrow was mod-
erately common in the open desert scrub at the base of the Sierra del Carmen
below 4800 feet. He said that the series of Black-throated Sparrows from
the Sierra del Carmen "resembles most the race opuntia of western Texas
. but shows some intergradation toward grisea of southern Coahuila and
toward A. b. bilineata of eastern Coahuila." He remarked also that the speci-
men from Jardin del Sur, which Marsh and Stevenson (1938:287) reported
as A. b. grisea, was in extremely worn, dirty summer plumage and contributed
nothing reliable to racial determination.

*Amphispiza bilineata grisea Nelson. — Specimen examined: one, S 31665,
from the north foot of Sierra Guadalupe ( = 10 mi. S, 5 mi. W General Ccpeda),
6500 ft., April 25, 1953.

A. b. grisea is the subspecies of Black-throated Sparrow in southern
Coahuila. Burleigh and Lowery (1942:208) saw this sparrow "frequently
on the arid plateau around Saltillo" and obtained specimens there that were
identified as A. b. grisea. Amadon and Phillips (1947:581) saw individuals
on August 8 and 28 that were feeding "fledged young near Saltillo." The
size of its wing ( 68.5 mm. ) , the slaty color of its back, and the size ( 8 mm. )
of the white spot on its lateral tail feather suggest that No. 31665 is
characteristic of A. b. grisea. The size of the testes (4.5x3 mm.) of No.
31665 indicates that A. b. grisea may breed in southern Coahuila.

*Junco phaeonotus palliatus Ridgway. — Specimens examined: total 3: $
35402 (skeleton only) from 13 mi. E San Antonio de las Alazanas, 9345 ft.,
April 10, 1954; $ 33226 (skeleton only) from 13 mi. E San Antonio de las
Alazanas, 9950 ft., July 6, 1955; and S 31633 from Mesa de Tablas, 8600 ft.,
January 16, 1954, weight, 22 gms.

In Coahuila the Mexican Junco seems to be common. Miller (1955a: 177)
found it in the conifers of the upper Corte Madera drainage at 7500 feet
and up to 8800 feet on Loomis Peak in the Sierra del Carmen. Marsh and
Stevenson (1938:287) took an adult in Vivoras Canyon on August 14 in the
Sierra del Carmen. Sutton and Burleigh (1939a:46) found small flocks at
Diamante Pass in March. Burleigh and Lowery (1942:208-209) noted /. p.

BiBDS From Coahuila, Mexico 511

palliatus as a common bird of the mountain slopes above an elevation of
about 7000 feet in southeastern Coahuila, and obtained specimens at Diamante
Pass. Ridgway (1901:300) recorded /. p. phaeonotus from Sierra Encar-
nacion. His record should be of /. p. palliatus as indicated by Miller,
Friedmann, Griscom, and Moore (1957:386). Dickerman saw Mexican Juncos
in the Sierra de la Madera on December 13, 1953.

No. 31633 shows no sign of intergrading with /. p. phaeonotus to the
south. The outermost rectrix of No. 31633 is wholly white; the second
rectrix is nearly as white. No. 31633 is paler than representatives of /. p.
phaeonotus from the southern part of the Central Plateau of Mexico.

Spizella passerina arizonae Coues. — Specimens examined: total 4: S 31110
from the Rio Grande (=17 mi. S Dryden, Terrell Co., Texas, in Coahuila),
600 ft., March 18, 1952, weight, 10.7 gms.; S 31111 and $ 31112 from 4 mi.
W Hacienda La Mariposa, 2300 ft., March 24, 1952, weights, 11.0, 11.7 gms.;
and $ 31666 from the north foot of Sierra Guadalupe (=10 mi. S, 5 mi. W
General Cepeda), 6400 ft., April 19, 1953, weight, 14.0 gms.

The Chipping Sparrow is a common spring and possibly fall migrant in
Coahuila. Miller (1955a: 177) noted small flocks from April 21 to 27 in the
Sierra del Carmen where specimens were taken. Burleigh and Lowery
(1942:209) indicated that S. p. arizonae was "quite common and of general
distribution" at Saltillo and Diamante Pass and remarked that specimens were
taken at these localities in April. Amadon and Phillips (1947:581) took two
Chipping Sparrows "near Las Delicias on August 17." Dickerman saw in-
dividuals in the Sierra del Pino on May 12, 1954, and at San Marcos on
May 4, 1954. Our specimens, which are typical representatives of S. p.
arizonae, are pale; the ground color of their backs is grayish buff.

Spizella pallida (Swainson). — The Clay-colored Sparrow is a migrant or
winter visitant in Coahuila. Burleigh and Lowery (1942:209) saw "large
flocks of sparrows, mostly of this species, ... on frequent occasions in
the cultivated fields and orchards on the outskirts of Saltillo." Four specimens
were taken by Burleigh and Lowery {loc. cit.) on April 19 and 20 at Saltillo.

Spizella breweri breweri Cassin. — Specimens examined: total 2: $ 31115
from the Rio Grande (=17 mi. S Dryden, Terrell Co., Texas, in Coahuila),
600 ft., March 18, 1952, weight, 8.2 gms.; and $ 31114 from 28 mi. S, 11 mi.
E Boquillas, 2000 ft., March 12, 1952, weight, 9.7 gms.

Brewer's Sparrow is probably a winter resident in much of Coahuila.
Miller, Friedmann, Griscom, and Moore (1957:389) recorded S. b. breweri
from 25 mi. NW Monclova on November 20 and from 8 mi. S Cuatro
Cienegas on November 15. The only definite records obtained by Burleigh
and Lowery (1942:209) of S. b. breweri are those of a female and a male
taken "near Saltillo" on April 16 and 18. The size of the dorsal area of
sandy buff with narrow streakings of Nos. 31114-31115 suggests S. b. breweri.

Spizella pusilla arenacea Chadboume. — Specimen examined: one, $ 31116,
from 4 mi. W Hacienda La Mariposa, 2300 ft., March 24, 1952, weight,
13.1 gms.

The Field Sparrow is an uncommon spring and probably fall migrant in
Coahuila. Other than No. 31116, S. p. arenacea has only been recorded from
Sabinas in March, when three specimens were obtained (Miller, Friedmann,

512 University of Kansas Publs., Mus. Nat, Hist.

Griscom, and Moore, 1957:390). The coloration of No. 31116 is much grayer
and tlie black streaks on its back are much narrower than on typical repre-
sentatives of S. p. pusilla.

** Spizella wortheni wortheni Ridgway. — The single specimen of Worthen's
Sparrow obtained by Burleigh and Lowery (1942:209) "just outside the
limits of Saltillo on April 16" represents the only record of occurrence of this
species in Coahuila.

** Spizella atrogularis atrogularis (Cabanis). — Miller, Friedmann, Griscom,
and Moore (1957:391) remarked that the Black-chinned Sparrow is a com-
mon resident of the Central Plateau from Durango and southern Coahuila
southward. Burleigh and Lowery (1942:212) noted the species only at the
foot of the mountains of southeastern Coahuila where, at 6000 feet, scattered
pairs were found. Miller (1955a: 177) observed a male on April 23 on a
slope at the mouth of Boquillas Canyon of the Sierra del Carmen; he pre-
siuned it to be a transient.

Zonotrichia leucophrys leucophrys (Forster). — Specimen examined: one,
S 30243, from 1 mi. SW San Pedro de las Colonias, 3700 ft., February 8, 1951.

The White-crowned Sparrow is a fairly common migrant or winter visitant
in Coahuila. Miller, Friedman, Griscom, and Moore (1957:393) recorded
Z. I. leucophrys from Sabinas on March 16 and February 23. Burleigh and
Lowery (1942:212) remarked that "small flocks of White-crowned Sparrows
were seen at infrequent intervals in thickets and stretches of underbrush on
the outskirts of Saltillo on April 18 and again on April 24 . . . [and] near
a small town some twenty miles west of Saltillo on April 22." Specimens
that Burleigh and Lowery (loc. cit.) collected "near Saltillo" were identified
as Z. I. leucophrys.

Zonotrichia leucophrys gamhelii (Nuttall). — Specimen examined: one, $
31117, from Sierra de la Encantada (=38 mi. S, 23 mi. E Boquillas), 4400
ft., March 15, 1952.

Miller (1955a: 177) noted Z. I. gambelii on April 27 at 4600 feet in Bo-
quillas wash in the Sierra del Carmen. Hellmayr (1938:568) hsted Z. /.
gamhelii from Sabinas.

Zontrichia leucophrys oriantha Oberholser. — This subspecies has been re-
corded from Sabinas on April 25 and Hip6hto on November 5 (Miller, Fried-
mann, Griscom, and Moore, 1957:393).

Melospiza lincolnii lincolnii (Audubon). — Specimen examined: one, $
31595, from 20 mi. S Ocampo, 7000 ft., April 5, 1954, measurements: wing,
58 mm.; tail, 50 mm.; weight, 12 gms.

Lincoln's Sparrow seems to be a fairly common migrant or winter visitant
in Coahuila; M. I. lincolnii is the common subspecies. Burleigh and Lowery
(1942:212) found this sparrow only in a grain field situated between a
small pond and a narrow stream on the outskirts of Saltillo; the four specimens
collected were identified as M. I. lincolnii. No. 31595 was obtained in a pine-
oak association.

Melospiza lincolnii alticola (Miller and McCabe). — Miller, Friedmann,
Griscom, and Moore (1957:398) listed M. I. alticola, which seems to be un-
common in Coahuila, from Sabinas on March 14.

Birds From Coahuila, Mexico 513

Melospiza lincolnii gracilis (Kittlitz). — Miller (1955a: 177) took an un-
sexed representative of M. I. gracilis on April 7 in Carboneras Canyon of the
Sierra del Carmen at 6700 feet. This occurrence is at the extreme eastern
range of this subspecies.

Melospiza georgiana ericrypta Oberholser. — In Coahuila this subspecies of
the Swamp Sparrow has been recorded as a migrant or winter visitant. Miller,
Friedmann, Griscom, and Moore ( 1957:399) recorded it from Sabinas on Febru-
ary 22 to March 8 and from 8 mi. S Cuatro Cienegas on November 4.

LITERATURE CITED

Aldrich, J. W., and Duvall, A. J.

1955. Distribution of American galHnaceous game birds. U. S. Dept.
Int., Fish and Wildlife Ser., Circular 34:ii+30 pp.

Amadon, D., and Phillips, A. R.

1947. Notes on Mexican birds. Auk, 64:576-581, October.

American Ornithologists' Union Check-list Committee

1957. Check-Ust of North American birds. 5th ed. xiv4-691 pp.

Baird, S. F.

1858. Reports of e\-plorations and surveys for a railroad route from the

Mississippi River to the Pacific Ocean. 9(pt. II — Birds):lvi-J-
1005 pp.

Baker, R. H.

1956. Mammals of Coahuila, Mexico. Univ. Kansas Publ., Mus. Nat. Hist.,
9:125-335, June 15.

Bangs, O.

1898. Some new races of birds from eastern North America. Auk,

15:173-183, April.
1925. The history and characters of Vermivora crissalis (Salvin and

Godman). Auk, 42:251-253, April.

Blake, E. R.

1953. Birds of Mexico. Univ. Chicago Press, Chicago, xxx-f 644 pp.

Brodkorb, p.

1935. A new flycatcher from Texas. Occas. Papers Mus. Zool, Univ,
Michigan, 306:1-3, January 30.

Burleigh, T. D., and Low^ry, G. H., Jr.

1942. Notes on the birds of southeastern Coahuila. Occas. Papers Mus.
Zool., Louisiana State Univ., 12:185-212, March 4.

Cory, C. B.

1919. Catalogue of birds of the Americas. Field Mus. Nat. Hist. (Publ.
203), Zool. Ser., 13 (Pt. II, no. 2):317-607, December 31.

Cory, C. B., and Hellmayr, C. E.

1927. Catalogue of birds of the Americas and the adjacent islands. Field
Mus. Nat. Hist. (Publ. 242), Zool. Ser., 13(Pt. V):vi-|-517 pp.,
April 11.

Davis, J.

1951. Distribution and variation of the Brown Towhees. Univ. Cali-
fornia Publ. Zool., 52:1-120, October 30.

514 University of Kansas Publs., Mus. Nat. Hist.

Dexon, K. L.

1955. An ecological analysis of the interbreeding of Crested Titmice in
Texas. Univ. California Publ. Zool., 54:125-206, December 8.

EvENDEN, F. G., Jr.

1952. Notes on Mexican bird distribution. Wilson Bull, 64:112-113,
June.

Friedmann, H.

1950. The birds of North and Middle America. U. S. Nat. Mus., Bull.
50 (Pt. XI):xiv-f793 pp.

Friedmann, H., Griscom, L., and Moore, R. T.

1950. Distributional check-list of the birds of Mexico, Pt. 1. Pacific
Coast Avifauna, 29:1-202, June 30.

Goldman, E. A.

1951. Biological investigations in Mexico. Smithsonian Miscel. Coll., 115:
xiv-f476 pp., July 31.

Goldman, E. A., and Moore, R. T.

1946. The biotic provinces of Mexico. Jour. Mamm., 26:347-360,
February.

Grinnell, J.

1909. A new cowbird of the genus Molothrus with a note on the prob-
able genetic relationships of the North American fonns. Univ.
Cahfornia, Publ. Zool., 5:275-281, December 31.

Griscom, L.

1950. Distribution and origin of the birds of Mexico. Bull. Mus. Comp.
Zool., 103:341-382.

Hardy, J. W., and Dickerman, R. W.

1955. The taxonomic status of the Maroon-fronted Parrot. Condor, 57:
305-306, September-October.

Hellmayr, C. E.

1929. Catalogue of birds of the Americas and the adjacent islands. Field
Mus. Nat. Hist. (Publ. 266), Zool. Sen, 13(Pt. VI):vi+258 pp.,
November 14.

1934. Catalogue of birds of the Americas. Ibid. (Publ. 330), Zool. Ser.,
13(Pt. VII):vi-f531 pp., November 15.

1935. Ibid. (Publ. 347), Zool. Ser., 13(Pt. VII):vi+541 pp., Septem-
ber 16.

1937. Catalogue of birds of the Americas and the adjacent islands.
Ibid. (Publ. 381), Zool. Ser., 13(Pt. X):vi+228 pp., April 12.

1938. Ibid. (Publ. 430), Zool. Ser., 13 (Pt. XI):vi-f662 pp., December 31.

Hellmayr, C. E., and Conover, B.

1942. Catalogue of birds of the Americas. Field Mus. Nat. Hist. (Publ.
514), Zool. Ser., 13 (Pt. I, no. l):vi-l-635 pp., April 30.

LOWERY, G. H.

1938. A new grackle of the Cassidix mcxicanus group. Occas. Papers
Mus. Zool., Louisiana State Univ., 1:1-11, May 4.

Marsh, E. G., Jr., and Stevenson, J. O.

1938. Bird records from northern Coahuila. Auk, 55:286-287, April.

Birds From Coahuila, Mexico 515

Mayb, E.

1946. History of the North American bird fauna. Wilson Bull., 58:3-41,
March.

Merriam, C. H.

1898. Life zones and crop zones of the United States. U. S. Dept.
Agric, Div. Biol. Surv., 10:1-79.

Miller, A. H.

1931. Systematic revision and natural history of the American shrikes
(Lanius). Univ. California Publ. Zool., 38:11-242, October 24.

1955a. The avifauna of the Sierra del Carmen of Coahuila, Mexico. Con-
dor, 57:154-178, May-June.

1955b. A hybrid woodpecker and its significance in speciation in the
genus Dendrocopos. Evolution, 9:317-321, September.
Miller, A. H., Friedmann, H., Griscom, L., and Moore, R. T.

1957. Distributional check-list of the birds of Mexico, Pt. 2. Pacific
Coast Avifauna, 33:1-436, December 20.
Moore, R. T.

1939. A review of the House Finches of the subgenus Burrica. Condor,
41:177-205, September-October.

1941. Three new races in the genus Otus from central Mexico. Proc.
Biol. Soc. Washington, 54:151-159, November 17.

1947. New species of parrot and race of quail from Mexico. Proc. Biol.
Soc. Washington, 60:27-28, April 3.

MULLER, C. H.

1947. Vegetation and climate of Coahuila, Mexico. Madrono, 9:33-57,
April.

Oberholser, H. C.

1902. A review of the larks of the genus Otocoris. Proc. U. S. Nat. Mus.,

24 (1271): 801-883, June 9.
1912. A revision of the forms of the Ladder-backed Woodpecker (Dryo-

bates scalaris [Wagler]). Proc. U. S. Nat. Mus., 41 (1847): 139-

159, May 20.
1919a. Distribution of a new Red-winged Blackbird from Texas. Wilson

Bull. 31:20-23, March.
1919b. The geographic races of Hedymeles melanocephalus Swainson.

Auk, 36:408-416, July.
1921. A revision of the races of Dendroica auduboni. Ohio Jour. Sci.,

21:240-248, May.

Packard, R. L.

1957. Broad-winged Hawk in Coahuila. Wilson Bull., 69:370-371,
December.

Phillips, A. R.

1950. The Great-tailed Crackles of the southwest. Condor, 52:78-81,
March.

PiTELKA, F. A.

1948. Notes on the distribution and taxonomy of Mexican game birds.
Condor, 50:113-123, May.

516 University of Kansas Publs., Mus. Nat. Hist,

RiDGWAY, R.

1901. The birds of North and Middle America. U. S. Nat. Mus., Bull.
50(Pt. I):.xxxii + 715 pp., October 24.

1902. Ibid. ( Pt. II ) :xx + 834 pp., October 16.
1904. Ibid. ( Pt. Ill ) :xx + 801 pp., December 31.
1907. Ibid. ( Pt. IV ) :.xxii + 973 pp., July 1.
1914. Ibid. ( Pt. VI ) :xx + 882 pp., AprU 8.
1916. Ibid. ( Pt. VII ) :xiv + 543 pp.. May 5.

RiDGWAY, R., and Friedmann, H.

1946. The birds of North and Middle America. U. S. Nat. Mus., Bull.
50 (Pt. X) :xii + 484 pp., December 18.

Sclater, P. L.

1857. Review of the species of the South American subfamily Tityrinae.
Proc. Zool. Soc. London, Pt. XXV, pp. 67-80.
Selander, R. K., and Baker, J. K.

1957. The Cave Swallow in Texas. Condor, 59:345-363, November-
December.

Sibley, C. G.

1950. Species formation in the Red-eyed Towhees of Mexico. Univ.
California Publ. Zool., 50:109-194, November 24.

Sutton, G. M., and Burleigh, T. D.

1939a. A list of birds observed on the 1938 Semple Expedition to north-
eastern Mexico. Occas. Papers Mus. Zool., Louisiana State Univ.,
3:15-46, April 5.
1939b. A new Screech Owl from Nuevo Leon. Auk, 56:174-175, April.

Van Hoose, S. G.

1955. Distributional and breeding records of some birds from Coahuila.
Wilson Bull, 67:302-303, December.

Van Rossem, A. J.

1934. Notes on some types of North American birds. Trans. San Diego
Soc. Nat. Hist, 7:347-362, May 31.

Van Tyne, J., and Sutton, G. M.

1937. The birds of Brewster County, Texas. Univ. Michigan Mus. Zool.,
Miscel. Publ., 37:1-115, August 24.

Webster, J. D., and Orr, R. T.

1958. Variation in the Great Homed Owls of Middle America. Auk,
75:134-142, April.

Wetmore, a.

1948. The Golden-fronted Woodpeckers of Texas and northern Mexico.
Wilson Bull., 60:185-186, September.

Transmitted February 27, 1959.

n

27-7905

(Continued from inside of front cover)

Vol. 8. 1. Life history and ecology of the five-lined skink, Eumeces fasciatus. By Henry
S. Fitch. Pp. 1-156, 26 figs, in text. September 1, 1954.

2. Myology and serology of the Avian Family Fringillidae, a taxonomic study.
By William B. Stallcup. Pp. 157-211, 23 figures in text, 4 tables. November
15, 1954.

3. An ecological study of the collared lizard (Crotaphytus coUaris). By Henry
S. Fitch. Pp. 213-274, 10 figures in text. February 10, 1956.

4. A field study of the Kansas ant-eating frog, Gastrophryne olivacea. By Henry
S. Fitch. Pp. 275-306, 9 figures in text. February 10, 1956.

5. Check-list of the birds of Kansas. By Harrison B. Tordoff. Pp. 307-359, 1
figure in text. March 10, 1956.

6. A population study of the prairie vole (Microtus ochrogaster) in northeastern
Kansas. By Edwin P. Martin. Pp. 361-416, 19 figures in text. April 2,
1956.

7. Temperatiu-e responses in free-living amphibians and reptiles of northeastern
Kansas. By Henry S. Fitch. Pp. 417-476, 10 figures in text, 6 tables. June
1, 1956.

8. Food of the crow, Corvus brachyrhynchos Brehm, in south-central Kansas. By
Dwight Piatt. Pp. 477-498, 4 tables. June 8, 1956.

9. Ecological observations on the woodrat, Neotoma floridana. By Henry S.
Fitch and Dennis G. Rainey. Pp. 499-533, 3 figures in text. June 12, 1956.

10, Eastern woodrat, Neotoma floridana: Life history and ecology. By Dermis
G. Rainey. Pp. 535-646, 12 plates, 13 figures in text. August 15, 1956.
Index. Pp. 647-675.

Vol. 9. 1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 fig-
tires in text. December 10, 1955.

2. Additional records and extensions of ranges of mammals from Utah. By
Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80.
December 10, 1955. i

3. A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rol-
lin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.

4. Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming.
By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956.

5. The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6
figures in text. May 19, 1956.

6. Additional remains of the multituberculate genus Eucosmodon. By Robert
W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.

7. Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures
in text. June 15, 1956.

8. Comments on the taxonomic status of Apodemus peninsulae, with description
of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 337-346,
1 figure in text, 1 table. August 15, 1956.

9. Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp.
347-351. August 15, 1956.

10. A new bat (Genus Leptonycteris) from Coahuila. By Howard J. Stains.
Pp. 353-356. January 21, 1957.

11. A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexica
By Robert J. Russell. Pp. 357-361. January 21, 1957.

12. Geographic variation in the pocket gopher, Thomomys bottae, in Colorado.
By PhiUip M. Youngman, Pp. 363-387, 7 figures in text. February 21,
1958.

13. New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones,
Jr. Pp. 385-388. May 12, 1958.

14. Pleistocene bats from San Josecito Cave, Nuevo Leon, Mexico. By J. Knox
Jones, Jr. Pp. 389-396. December 19, 1958.

15. New Subspecies of the rodent Baiomys from Central America. By Robert L.
Packard. Pp. 397-404. December 19, 1958.

16. Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405-
414, 1 figure in text. May 20, 1959.

17. Distribution, variation, and relationships of the montane vole, Microtus mon-
tanus. By Emil K. Urban. Pp. 415-511. 12 figs, in text, 2 tables. August
1, 1959.

More numbers will appear in volume 9.

(Continued on outside of back cover)

(Continued from inside of back cover)

Vol. 10. 1. Studies of birds lolled in nocturnal migration. By Harrison B. Tordoff and
Robert M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 1956.

2. Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 Gguie. December 20, 1956.

3. The forest habitat of the University of Kansas Natural History Reservation.
By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figmres
in text, 4 tables. December 31, 1956.

4. Aspects of reproduction and development in the prairie vole (Microtus ochro-
gaster). By Henry S. Fitch. Pp. 129-161, 8 figmes in text, 4 tables. Decem-
ber 19, 1957.

5. Birds found on the Arctic slope of northern Alaska. By James W. Bee.
Pp. 163-211, pis. 9-10, 1 figure in text. March 12, 1958.

6. The wood rats of Colorado: distribution and ecology. By Robert B. Finley,
Jr. Pp. 213-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958.
More numbers will appear in volume 10.

Vol. 11. 1. The systematic status of the colubrid snake, Leptodeira discolor Gilnther.
By William E. Duellman. Pp. 1-9, 4 figs. July 14, 1958.

2. Natural history of the six-lined racerunner, Cnemidophonis sexlineatus. By
Henry S. Fitch. Pp. 11-62, 9 figs., 9 tables. September 19, 1958.

3. Home ranges, territories, and seasonal movements of vertebrates of the
Natm-al History Reservation. By Henry S. Fitch. Pp. 63-326, 6 plates, 24
figures in text, 3 tables. December 12, 1958.

4. A new snake of the genus Geophis from Chihuahua, Mexico. By John M.
Legler. Pp. 327-334, 2 figures in text. January 28, 1959.

5. A new tortoise, genus Gopherus, from north-central Mexico. By John M.
Legler. Pp. 335-343, 2 plates. AprU 24, 1959.

6. Fishes of Chautauqua, Cowley and Elk counties, Kansas. By Artie L.
Metcalf. Pp. 345-400, 2 plates, 2 figures in text, 10 tables. May 6, 1959.

7. Fishes of the Big Blue River Basin, Kansas. By W. L. Minckley. Pp. 401-442,
2 plates, 4 figures in text, 5 tables. May 8, 1959.

8. Birds from Coahuila, Mexico. By Emil K. Urban. Pp. 443-516. August 1,
1959.

More numbers will appear in Volume 11.
Vol. 12. 1. Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry
A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text, July 8, 1959.
2. The ancestry of modern Amphibia: a review of the evidence. By Theodore
H. Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, 1959.
More nmnbers will appear in volume 12.

-^I\ - L

University of Kansas Publication; i
Museum of Natural History

MUS. COMF. 7001
LiSIUJiy

=Sffi=3 19591

HARVARD
l/NfVERSITy

Volume 11, No. 9, pp. 517-525, 2 pis., 1 fig.
August 14, 1959

Description of a New Softshell Turtle
From the Southeastern United States

BY

ROBERT G. WEBB

University of Kansas
Law^rence
^ 1959

University of Kansas Publications, Museum of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 11, No. 9, pp. 517-525, 2 pis., 1 fig.
Published August 14, 1959

University of Kansas
Lawrence, Kansas

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA. KANSAS

I 9S»

27-8360

Description of a New Softshell Turtle
From the Southeastern United States

m. tm, zooi

SEP -3 1959

UNIVERSITY

BY

ROBERT G. WEBB

Examination of softshell turtles allied to Trionyx muticus from
the southeastern United States discloses the presence of an un-
described subspecies inhabiting river systems of the Gulf Coast.

The author is indebted to Mr, Roger Conant for constructive criti-
cism of the manuscript. I am grateful also to many fellow students
for assistance in field work or for other courtesies, especially Wil-
liam E. Erode, Franklin Sogandares-Bernal, Ernest A. Liner,
Donald W, Tinkle, Paul K. Anderson, and John K. Greer. The
photographs were provided through the cooperation of Roger and
Isabelle Hunt Conant and John M. Legler.

Collections from which specimens were obtained are as follows:
TU (Tulane University), USNM (United States National Mu-
seum), MCZ (Museum of Comparative Zoology, Harvard College),
CNHM (Chicago Natural History Museum), KU (Museum of
Natural History, University of Kansas), UI (Museum of Natural
History, University of Illinois ) .

Measurements ( in millimeters ) were made with a Vernier caliper
and a metal tape; those of the holotype were made to the nearest
one-tenth millimeter. Plastral length was measured from the
posterior edge of the plastron to the anteriormost edge of the
ventral surface; other measurements were maximal. Depth of shell
was taken only on hatchlings and an immature female. Hatchlings
were arbitrarily designated as specimens having plastrons shorter
than 44 mm; sex of all specimens except adult males was determined
by dissection unless otherwise noted.

Trionyx muticus calvatus new subspecies
Gulf Coast Smooth Softshell

Amyda mutica (in part), Stejneger, Bull. Mus. Comp. Zool., 94(l):23-24,

1944.
Amyda muticus (in part), Cook, Jour. Mississippi Acad. Sci., 1941-1947,

p. 185, 1946.
Trionyx muticus Anderson, Copeia, 3:211, August 28, 1958.

Holotype. — UI 31071, hatchling, sex undetermined, from the Pearl River,
Roses Bluff, 14 miles northeast Jackson, Rankin County, Mississippi; obtained
by William F. Childers on August 25, 1952 (Plate 1).

Paratypes. — A total of 20 alcoholic specimens: TU 17301, hatchhng male
(Plate 2). TU 17302-.1, 16682, three hatchling females, and TU 13473,

(519)

520 University of Kansas Publs., Mus. Nat. Hist.

adult female, from the Escambia River, 2 miles east and 1 mile north of
Century, Escambia County, Florida; TU 17306, adult female, from the Pearl
River, 9 miles south of Monticello, Lawrence County, Mississippi; USNM
7655, hatchhng, sex undetermined, and KU 47117-19, three adult males, from
the Pearl River, 1 mile south to 4 miles north of Monticello, Lawrence County,
Mississippi; TU 17303-.4, 17304-.3, five hatchling males and four hatchling
females, from the Pearl River, Vamado, Washington Parish, Louisiana; TU
17305, immature female, no data.

Diagnosis and definition. — A subspecies of softshell turtle most closely allied
to Trionyx muticus muticus but differing from that subspecies in having:
( 1 ) a Juvenal pattern of large, circular spots, ( 2 ) no stripes on dorsal surface
of snout, and (3) postocular stripe with thick, black borders immediately be-
hind eye in adult males. T. m. calvatus resembles T. m. muticus, and differs
from the several subspecies of Trionyx spinifer in having: (1) no enlarged
tubercles on anterior edge of carapace, (2) no ridge projecting from nasal
septum, and (3) a smooth dorsal surface on carapace in adult males.
T. m. calvatus and T. m. muticus resemble T. ferox in having a smooth dorsal
surface on carapace in adult males, but differ from T. ferox in having: (1) no
tubercles along anterior edge of carapace, and (2) no ridge projecting from
nasal septum.

Description of Iwlotype. — Carapace circular, widest at region of bridge;
margin entire; dorsal surface smooth; anterior margin of carapace lacking
tubercles; blunt vertebral ridge evident anteriorly; maximum length, 53.1 mm;
greatest width, 46.3 mm; greatest depth, 11.5 mm.

Plastron small, extending slightly farther forward than carapace; anterior
lobe truncate with slight midventral indentation; posterior lobe rounded,
sides forming acute angle; certain features of bony elements of plastron visible
through overlying skin; width of bony bridge, 4.5 mm; maximum length of
plastron, 37.5 mm.

Head extended to level of eyes; head terminating in long, rounded, flexible
snout; nostrils rounded with no ridges projecting from nasal septum; jaws
closed, each covered by fleshy lips except anteriorly where horny portions of
jaws are exposed; iris with dark stripe through pupil.

Forefeet and hind feet well- webbed and with five digits each; each limb
with nails on first three digits; dorsal surface of each forelimb with four
cornified areas, three of which have a free edge; each hind limb with two
comified areas, one smooth on posterodorsal surface and other with free edge
on posteroventral surface.

Tail terminating in flexible point and not extending beyond posterior edge
of carapace; anus to tip of tail, 2.6 mm; anus to posterior edge of carapace,
8.1 mm.

In preservative: Ground color of carapace dark tan having pattern of 49
brownish spots; 47 spots circular; two spots noticeably elongate, one rep-
resenting fusion of two circular spots; 17 spots on carapace not exceeding
2.0 mm in diameter, whereas 32 spots range from 2.5 to 4.0 mm in diameter;
periphery of carapace pale except anteriorly; maximum width of pale margin
(posteriorly), 3.3 mm; junction of pale margin and dorsal ground color formed
by rough-edged line composed of small, closely-set dots; pattern of fine
punctations and other marks on dorsal surface of forelimbs and hind limbs.

Description of a New Softshell Turtle 521

Ground color of underparts whitish, lacking markings; top of head and snout
gray, lacking markings; lower eyelids with small dark dots.

Description of paratypes. — Adult females (2 specimens). No striping on
dorsal surface of snout; pale postocular stripe not distinct, dark borders
obscure (head not extended in TU 13473); carapace circular, pale brown with
mottled pattern; carapace lacking pattern of large spots; dark marks present
in pale margin of carapace; dorsal surface of soft parts of body finely stippled,
larger marks on hind Hmbs and on anterior surface of forelimbs near their in-
sertions; plastron and ventral surface of soft parts of body without markings.
Maximal measurements, respectively, are: length of plastron, 172 and 180 mm;
length of carapace, 238 and 263 mm; width of carapace, 203 and 218 mm;
width of head, 28 and ? mm.

Immature female (1 specimen). Carapace circular having juvenal pattern
of large spots, some of which have borders darker than their centers and are
best described as ocelli; junction of pale margin and ground color of carapace
formed by ill-defined, ragged dark border; dorsal surface of forelimbs and
hind limbs finely streaked and dotted, larger marks occurring toward inser-
tions of forelimbs; lower border of pale postocular stripe in contact with
upper margin of postlabial pale stripe; no stripes on dorsal surface of snout;
fine markings on dorsal surface of neck. Maximal measurements are: length of
plastron, 56 mm; length of carapace, 82 mm; width of carapace, 77 mm;
depth of shell, 13 mm; width of head, 12 mm.

Adult males (3 specimens). No striping on dorsal surface of snout; pale
postocular stripe with thick, black borders immediately behind eye; width of
black borders equal to approximately one-half width of pale postocular stripe;
dorsal surface of soft parts of body with indistinct markings that are larger
on hind limbs; plastron and ventral surface of soft parts of body without
markings; small dark spots posteriorly along ventral edge of carapace; pale
margin of carapace lacking markings or having few small black spots; carapace
circular with or without pattern of large spots. Maximal measurements of
smallest and largest specimens, respectively, are: length of plastron, 108 and
118 mm; lengtli of carapace, 160 and 177 mm; width of carapace, 142 and
152 mm; width of head, 21 mm.

Hatchhngs ( 14 specimens ) . These paratypes resemble the holotype in
all features mentioned; markings on neck tend to form longitudinal streaks in
TU 17303 and 17304. There are no secondary sexual differences in hatchling
turtles.

There is some variation in hatchling turtles. Four from the Escambia River
have dorsal spots 3 mm or larger in greatest diameter and on three specimens
the dorsal spots number 27, 37 and 37 (total number not discernable in
TU 16682); none of the dorsal spots is ocellate. Maximal measurements of
these three hatchlings, respectively, are: length of plastron, 35, 36 and 37 mm;
length of carapace, 50, 50 and 52 mm; width of carapace, 44, 45 and 47 mm;
depth of shell, 11 mm; width of head, 9 mm. Nine hatchlings from the Pearl
River at Varnado have more (all small) dorsal spots, which may be ocellate.
The dorsal spots and ocelli do not exceed 2 mm in their greatest diameter
except that some of those of TU 17304 are 3 mm; the spots range in number
from 38 (TU 17303) to 63 (TU 17304). Maximal measurements of the
smallest and largest specimens, holotype excepted, are: length of plastron, 30

522

University of Kansas Publs., Mus. Nat. Hist.

and 33 mm; length of carapace, 42 and 46 mm; width of carapace, 37 and 43
mm; depth of shell, 9 and 10 mm; width of head, 9 and 10 mm. The holo-
type resembles hatchlings from the Escambia River in having large, non-ocel-
late dorsal spots 3 mm in greatest diameter, and larger measurements.

One other specimen (not designated as a paratype), consisting of a head
with a few attached cervical vertebrae, was obtained on a sand bank of the
Escambia River, Florida. The postocular stripe, bright yellow with black
borders, was especially vivid in this adult male (KU 47116).

Range. — Trionyx m. calvatus is knowTi from the Pearl, Pascagoula and
Escambia river drainages and is to be expected in the Tombigbee-Alabama
river drainage (Fig. 1). Tinkle (1958:41, fig. 53, stippled) has indicated
the probable range of calvatus. This subspecies is unknown from the Missis-
sippi and Tennessee river drainages, which are inhabited by T. m. muticus. The

Fig. 1. Map of southeastern United States showing record stations of Trionyx
muticus calvatus (solid symbols) and Trionyx m. muticus (open symbols).
Circles indicate specimens examined; triangles indicate records in the literature.
The question mark refers to a specimen oearing catalogue number 17236 in
the collection of Tulane University (see comments on page 524 concerning

No. 17236 from the Amite River).

Description of a New Softshell Turtle 523

western limit of distribution is the Pearl River drainage and probably those
streams of the Florida Parishes of Louisiana that drain into Lake Ponchartrain.
The most easterly record of occurrence for T. m. calvatus is in the Escambia
River drainage; the eastern extent of geographic range is not known.

I have seen three preserved young turtles having the characteristic spotted
pattern from the Pascagoula drainage in eastern Mississippi. These specimens
are uncatalogued and in the collections at Mississippi Southern College,
Hattiesburg, Mississippi.

There is a specimen of T. m. muticus labeled as from Mobile, Alabama
(MCZ 1596), for which I believe the locahty datum is incorrect. It is a
young turtle having a well-defined pattern on the carapace and is without
doubt a representative of T. m. muticus. Mobile is in the large drainage basin,
of the Tombigbee, Black Warrior, Coosa and Alabama rivers, which is be-
tween the Escambia and Pearl rivers.

Yarrow (1882:28) reported a specimen of Amyda mutica, USNM 11630,
from Gainesville, Florida. This record was questioned by Cahn (1937:179),
and has been disregarded by subsequent authors. Stejneger (1944:23) lists
this specimen number with uncertainty from Mt. Carmel, Ilhnois. The exact
geographic provenance of this specimen is seemingly unknown.

Habitat. — I have collected eggs of T. m. calvatus on sand banks of the
Escambia River, Florida. The Escambia River has a sand-gravel bottom,
extensive sandy banks, a moderately-rapid current, and is flanked by a thick
riparian forest. It is approximately 80 feet wide with fallen trees and brush
intermittently emergent along the shoreline. The sand bar-habitat along the
Pearl River has been mentioned by Anderson (1958:212). All records thus
far are from lotic habitats.

Comparisons. — Trionyx m. calvatus is most closely related to Trionyx m.
muticus. Both subspecies have the following characteristics: (1) no en-
larged tubercles on the anterior edge of the carapace, (2) no ridge projecting
from the nasal septum, and (3) a smooth carapace in adult males. These
characters distinguish these two subspecies from the several subspecies of
T. spinifer, and, except for the smooth carapace in adult males, from T. ferox.
Another feature of T. m. calvatus and T. m. muticus, not known to be definitive
or diagnostic but noticed on occasion, is the pale orange cast, in hfe, of the
dorsal surface of the carapace and soft parts of the body in young of these
turtles.

The spotted pattern of juveniles of calvatus is easily distinguished from
the pattern of muticus (small dots, streaks and dashes) figured by Agassiz
(1857, vol. 2, pt. 3, pi. 6, fig. 6), Smith (1950:154, fig. 104), Conant (1938:
192, pi. 21, fig. 1; 1958, pi. 11, opposite p. 94), and Cahn (1937:177, pi. 24C),

Unfortunately, the distinctive dorsal spotting in young calvatus becomes
obscure or absent in some adults of both sexes. Spotting in large males is
not so well-defined as in juveniles; it may be absent (TU 17306.3), or in-
dicated by two obscure spots (KU 17117), but is usually evident, at least
posteriorly. The spotted pattern is absent in large females, which have a pale,
mottled and blotched pattern of Uchen-like figures; dorsal spots are obscure
in TU 17305 (length of plastron, 56 mm).

Two additional features are, so far as known, universal in calvatus; these
are: (1) the absence of striping on the dorsal surface of the snout, and (2)

524 University of Kansas Publs., Mus. Nat. Hist.

the presence of thick, black borders of the postocular stripe in adult males.
These features have also been observed in some specimens of muticus; their
presence in muticus cannot be properly evaluated at this time, and is seem-
ingly not due to individual variation. These two characters, however, coupled
with the distinctive juvenile pattern of spots, serve, in combination, to dis-
tinginsh calvatus from muticus.

Discussion. — The two populations are recognized as subspecies
because: (1) there is close resemblance, (2) the diagnostic char-
acters pertaining to pattern are few and superficial, and (3) the
geographic ranges are allopatric, but juxtaposed. It is probable
that muticus and calvatus would be capable of interbreeding if they
were not spatially isolated. It should be pointed out, however,
that there is no evidence of intergradation between muticus and
calvatus in the lower Mississippi Valley as has been reported for
the subspecies of T. spinifer ( Conant and Goin, 1948 ) , and that the
degree of difference between calvatus and muticus is greater than
that between some subspecies of T. spinifer.

Specimens examined. — All the localities listed below are plotted on tlie
distribution map (Fig. 1). Only those specimens of T. muticus muticus are
listed that serve to delimit the range of T. m. calvatus. Fortunately, the
identification of the specimens of muticus is certain as all show the character-
istic juvenile pattern, except the large female, TU 7543, from southeastern
Louisiana. USNM 95133-34 (carapaces and plastrons only) and TU 17236
are females, which lack the diagnostic spotted pattern of calvatus; the former
are referred to this subspecies on geographic grounds ( Pearl River at Columbia,
Mississippi). TU 17236, from the Amite River, is dubiously relegated to
calvatus on the supposition that this river and others in the Lake Ponchartrain
drainage will yield the characteristic juveniles.

Trionyx m. calvatus (33 specimens): TU 13473, 16682, 17301, 17302-.1,
KU 47116 (skull only), Escambia River, 2 miles east, 1 mile north Century,
Escambia Co., Florida; TU 17303-.4, 17304-.3, Pearl River, Vamado, Wash-
ington Par., Louisiana; TU 17306-.3, Pearl River, 9 miles south Monticello,
Lawrence Co., Mississippi; TU 16956, KU 47117-19, USNM 7655, Pearl
River, vicinity of Monticello, Lawrence Co., Mississippi; TU 17236?, Amite
River, near Baton Rouge, Louisiana; TU 13795, Bogue Chitto River, Enon,
Washington Par., Louisiana; TU 17305, no data, Louisiana; USNM 95133-34,
Pearl River, Columbia, Marion Co., Mississippi; UI 31071 Pearl River, 14
miles northeast Jackson, Rankin Co., Mississippi; Uncatalogued, see page
523, Leaf River, 3 miles southeast New Augusta, Perry Co., Mississippi.

Trionyx m. muticus ( 6 specimens ) : TU 5989, Ouachita River, Monroe,
Ouachita Par., Louisiana; TU 7543, Vacherie, St. James Par., Louisiana;
CNHM 7845, Gayles, Caddo Par., Louisiana; USNM 92605, Greenville, Wash-
ington Co., Mississippi; USNM 113228, Jonesville, Catahoula Par., Louisiana;
USNM 118167, Wheeler Reser\'oir, Tennessee River, Alabama.

Records in the Literature. — USNM 113228, referred to above as Trionyx m.
muticus is listed by Stejneger (1944:56) as Amyda s. spinifera; four of the
specimens listed above (USNM 7655, 92605, 95133-34) are recorded by
Stejneger (op. ci*.: 23-34) as Amyda mutica. Cook (1946:185) records seven
specimens of the muticus group from Mississippi as follows: 1, no data
1, Vicksburg, Warren Co.; 3, Forrest Co.; 1, Crawford Bridge, Jones Co.
1, Lake Park, Columbus, Lowndes Co. I have not seen these specimens

PLATE 13

Trionyx muticus calvatus new subspecies, hatchling, UI 31071,

holotype (X 1.3). Top, dorsal view. Bottom, ventral view.

Photographs by John M. Legler.

PLATE 14

Trionyx inttticus calvatus new subspecies, liatcliling male, TV 17301, para-
typc (X 1.3). Top, dorsal view. Bottom, lateral view of left side. Photo-
graphs by Isabellc Hunt Conant.

Description of a New Softshell Turtle 525

they are plotted on the distribution map — the one from Vicksburg as muticus
and the others as calvatus on geographic grounds. The hatchUngs of
Trionyx muticus referred to by Anderson {loc. cit.) include the nine para-
types from Varnado, Louisiana.

LITERATURE CITED
Anderson, P. K.

1958. The photic responses and water-approach behavior of hatchling
turtles. Copeia, 1958, 3:211-215, 5 figs., August 28.
Agasse, L.

1857. Contributions to the natural history of the United States. Vol. II,
Part III. Embryology of the turtle. Little, Brown and Co., Bos-
ton, pp. 451-643, 27 pis.
Cahn, a.

1937. The turtles of Illinois. Illinois Biol. ?vIonogr., 16(1-2): 1-218, 31
pis., 15 figs., 20 maps, August 31.

CONANT, R.

1938. The reptiles of Ohio. Amer. Midi. Nat., 20(l):l-200, 26 pis., 38
maps, July.

1958. A field guide to reptiles and amphibians of eastern North America.
Houghton Mifflin Co., Boston, pp. vii-|-366, 40 pis., 62 figs.,
248 maps.
CoNANT, R., and C. J. Coin.

1948. A new subspecies of soft-shelled turtle from the central United
States, with comments on the application of the name Amyda.
Occas. Pap. Mus. Zool, Univ. Michigan, 510:1-19, 2 pis., 1 map,
June 15.
Cook, F. A.

1946. Distribution of species of Amyda in Mississippi. Joum. Mis-
sissippi Acad. Sci., 1941-1947:185-190.
Smith, H. M.

1950. Handbook of amphibians and reptiles of Kansas. Univ. Kansas
Mus. Nat. Hist., Misc. Publ., 2:1-336, 233 figs., September 12.
Stejneger, L.

1944. Notes on the American soft-shell turtles with special reference to
Amyda Agassizii. Bull. Mus. Comp. Zool., 94(l):l-75, 30 pis.
Tinkle, D. W.

1958. The systematics and ecology of the Sternothaerus carinatus com-
plex (Testudinata, Chelydridae). Tulane Stud. Zool., 6(l):l-56,
57 figs.
Yarrow, H. C.

1882. Check list of North American Reptilia and Batrachia, with cata-
logue of specimens in the U. S. National Museum. Bull. U. S.
Nat. Mus., 24:1-249.

Transmitted April 30, 1959.

n

27-8360

y/f

-Lf^WR:

University of Kansas Publications
Museum of Natural History

Volume 11, No. 10, pp. 527-669, 16 pis., 29 figs.
March?, 1960

'A "1 O fi/'Wi^ ,

I f^

Natural History of the Ornate Box Turtle,
Terrapene omata ornata Agassiz

BY
JOHN M. LEGLER

University of Kansas

Lawrence

1960

UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY

Institutional libraries interested in publications exchange may obtain this
series by addressing the Exchange Librarian, University of Kansas Library,
Lawrence, Kansas. Conies for individuals, persons vi'orking in a particular
field of study, may be ootained by addressing instead the Museum of Natural
History, University of Kansas, Lawrence, Kansas. There is no provision for
sale of this series by the University Library, which meets institutional requests,
or by the Museum of Natiual History, which meets the requests of individuals.
When individuals request copies from the Museum, 25 cents should be in-
cluded, for each separate number that is 100 pages or more in length, for the
purpose of defraying the costs of wrapping and mailing.

* An asterisk designates those numbers of which the Museum's supply (not the Library'!
gupply) is exhausted. Numbers published to date, in this series, are as follows:

Vol. 1. Nos. 1-26 and index. Pp. 1-638. 1946-1950.
•Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140
figures in text. April 9, 1948.
Vol. 3. *1. The avifaima of Micronesia, its origin, evolution, and distribution. By Rol-
lin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951.
*2. A quantitative study of the nocturnal migration of birds. By George H.
Lowery, Jr. Pp. 361-472, 47 figures in text. June 29. 1951.

3. Phvlogeny of the wax^vings and allied birds. By M. Dale Arvey. Pp. 473-
530, 49 figures in text, 13 tables. October 10, 1951.

4. Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and
Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10,
1951.

Index. Pp. 651-681.
*Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31
figures in text. December 27, 1951.
Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953.
♦Vol. 6. (Complete) Mammals of Utah, taxonomtf and distribution. By Stephen D.
Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952.
Vol. 7. *1. Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, 73 figures in
text, 37 tables. August 25, 1952.

2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry
S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 figures in text. August
24, 1953.

3. The silky pocket mice (Perognathus flavus) of Mexico. By RoUin H. Baker.
Pp. 339-347, 1 figure in text. February 15, 1954.

4. North American jumping mice (Genus Zapus). By Philip H. Knitzsch. Pp.
349-472. 47 figures in text, 4 tables. April 21, 1954.

5. Mammals from Southeastern Alaska. By Rollin H. Baker and James S.
Findley. Pp. 473-477. April 21. 1954.

6. Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479-
487. April 21, 1954.

7. Subspeciation in the montane meadow mouse, Microbis montanus, in Wyo-
ming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text.
July 23, 1954.

8. A new subspecies of bat (Myotis vellfer) from southeastern California and
Arizona. By Terry A. Vaughan. Pp. 507-512. July 23, 1954.

9. Mammals of the San Gabriel mountains of California. By Terry A. Vaughan,
Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954.

10. A new bat (Genus Pipistrellus) from northeastern Mexico. By Rollin H.
Baker. Pp. 583-586. November 15, 1954.

11. A new subspecies of pocket mouse from Kansas. By E. Raymond Hall. Pp.
587-590. November 15, 1954.

12. Geographic variation in the pocket gopher, Cratogeomys castanops, in Coa-
huila, Mexico. By Robert J. Russell and Rollin H. Baker. Pp. 591-608.
March 15, 1955.

13. A new cottontail (Sylvilagus floridanus) from northeastern Mexico. By Rollin
H. Baker. Pp. 609-612. April 8, 1955.

14. Taxonomy and distribution of some American shrews. By James S. Findley.
Pp. 613-618. June 10, 1955.

15. The pigmy woodrat, Neotoma goldmani. Its distribution and systematic posi-
tion. By Dennis G. Rainey and Rollin H. Baker. Pp. 619-624, 2 figures in
text. June 10. 1955.

Index. Pp. 625-651.

(Continued on inside of back cover)

University of Kansas Publications
Museum of Natural History

Volume 11, No. 10, pp. 527-669, 16 pis., 29 figs.
March?, 1960

Natural History of the Ornate Box Turtle,
Terrapene ornata ornata Agassiz

BY
JOHN M. LEGLER

University of Kansas

Lawrence

1960

University of Kansas Publications, Museum of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 11, No. 10, pp. 527-669, 16 pis., 29 figs.
Published March 7, 1960

University of Kansas
Lawrence, Kansas

MUS. CQMP. ZOOL
LIBRARY

MAR 2 4 1960

PRINTED IN

THE STATE PRJNTING PLANT

TOPEKA. KANSAS

IS60

28-773

Natural History of the Ornate Box Turtle,
Terrapene ornata ornata Agassiz

BY

John M. Legler

CONTENTS

PAGE

Introduction 531

Acknowledgments 531

Systematic Relationships and Distribution 532

Fossils 534

Economic Importance 534

Study Areas 535

Materials and Methods 537

Terminology 539

Habitat and Limiting Factors 539

Habitat in Kansas 542

Reproduction 543

Mating 543

Insemination 545

Sexual Cycle of Males 545

Sexual Cycle of Females 549

Nesting 554

Eggs 558

Embryonic Development 560

Fertility and Prenatal Mortahty 564

Reproductive Potential 565

Number of Reproductive Years 565

Growth and Development 565

Initiation of Growth 565

Size and Appearance at Hatching 566

Growtli of Epidermal Laminae 568

Growth of Juveniles 575

Growth in Later Life 578

Annual Period of Growth 580

Environmental Factors Influencing Growth 580

Number of Growing Years 584

Longevity 585

Weight 586

(529)

530 University of Kansas Publs., Mus. Nat. Hist.

PACE

Bony Shell 586

Color and Markings 593

Wear 595

Sexual Dimorphism 596

Temperature Relationships 598

Optimum Temperature 599

Basking 600

Toleration of Thermal Maxima and Minima 601

Hibernation 611

Diet 617

Populations 623

Movements 626

Locomotion 627

Daily Cycle of Activity 629

Seasonal Cycle of Activity 630

Home Range 632

Homing Behavior 636

Social Relationships 637

Injuries 638

Repair of Injuries to the Shell 641

Ectoparasites 643

Predators 646

Defence 648

Discussion of Adaptations 650

Summary 656

Literature cited 663

Natural History of the Ornate Box Turtle 531.

INTRODUCTION

The ornate box turtle, Terrapene o. ornata Agassiz, was studied
more or less continuously from September, 1953, until July, 1957.
Intensive field studies were made of free-living, marked populations
in two small areas of Douglas County, Kansas, in the period 1954 to
1956. Laboratory studies were made, whenever possible, of phe-
nomena diflBcult to observe in the field, or to clarify or substantiate
field observations. Certain phases of the work ( for example, studies
of populations and movements ) were based almost entirely on field
observation whereas other phases (for example, growth and game-
togenic cycles) were carried out almost entirely within the labora-
tory on specimens obtained from eastern Kansas and other localities.

A taxonomic revision of the genus Terrapene was begun in 1956
as an outgrowth of the present study. The systematic status of
T. ornata and other species is here discussed only briefly.

Objectives of the study here reported on were: 1) to learn as
much as possible concerning the habits, adaptations, and life history
of T. o. ornata; 2) to compare the information tlius acquired with
corresponding information on other emyid and testudinid chelo-
nians, and especially with that on other species and subspecies of
Terrapene; 3) to determine what factors hmit the geographic
distribution of ornate box turtles; and, 4) to determine the role of
ornate box turtles in an ecological community.

Acknowledgments

The aid given by a number of persons has contributed substantially to the
present study. I am grateful to my wife. Avis J. Legler, who, more than any
single person, has unselfishly contributed her time to this project; in addition to
making all the liistological preparations and typing the entire manuscript, she
has assisted and encouraged me in every phase of the study. Dr. Henry S.
Fitch has been most helpful in offering counsel and encouragement. Thanks
are due Professor E. Raymond Hall for critically reading the manuscript.

Special thanks are due also to the following persons: Professor A. B. Leonard
for helpful suggestions dealing with photography and for advice on several
parts of the manuscript; Professor William C. Young for the use of facihties
at the Endocrine Laboratory, University of Kansas; Professor Edward H. Taylor
for permission to study specimens in his care; Dr. Richard B. Loomis for
identifying chigger mites and offering helpful suggestions on tlie discussion of
ectoparasites; Mr. Irwin Ungar for identification of plants; and, Mr. Wilham
R. Brecheisen for allowing me to examine his field notes and for assistance with
field work. Identifications of animal remains in stomachs were made by Pro-
fessor A. B. Leonard (moUusks, crustaceans), Dr. George W. Byers (arthro-
pods), and Dr. Sydney Anderson (mammals).

Miss Sophia Damm generously permitted the use of her property as a study
area and Mr. Walter W. Wulfkuhle made available two saddle horses that

532 University of Kansas Publs., Mus. Nat. Hist.

greatly facilitated field work. The drawings (with the exception of Fig. 21)
are by Miss Lucy Jean Remple. All photographs are by the author.

I am grateftJ also to the Kansas Academy of Science for three research
grants (totaUng $175.00) that supported part of the work. The brief dis-
cussion of taxonomic relationships and distribution results partly from studies
made by means of two research grants (totaling $150.00), from the Graduate
School, University of Kansas, for which I thank Dean John H. Nelson.

Systematic Relationships and Distribution

Turtles of the genus Terrapene belong to the Emyidae, a family com-
prising chiefly aquatic and semiaquatic species. Terrapene, nevertheless, is
adapted for terrestrial existence and di£Fers from all other North American
emyids in having a hinged and movable plastron and a down-turned (although
often notched) maxillary beak. Emydoidea blandingi, the only other North
American emyid with a hinged plastron, lacks a down-turned beak. The
adaptations of box turtles to terrestrial existence (reduction of webbing be-
tween toes, reduction in number of phalanges, reduction of zygomatic arch,
and heightening of shell) occur in far greater degree in true land tortoises
of the family Testudinidae. Four genera of emyid turtles in the eastern
hemisphere (Cuora, Cyclemys, Emtjs, and Notochelys) possess terrestrial adap-
tations paralleling those of Terrapene but (with the possible exception of
Cuora) the adaptations are less pronounced than in Terrapene. A movable
plastron has occurred independently in two groups of emyids in the New
World and in at least three groups in the Old World.

The genus Terrapene, in my view, contains seven species, comprising 11
named kinds. Of these species, five are poorly known and occur only in
Mexico. Terrapene mexicana (northeastern Mexico) and T. yucatana (Yu-
catan peninsula) although closely related, diflFer from each other in a number
of characters. Similarly, Terrapene klauberi (southern Sonora) and T. nehoni
(Tepic, Nayarit — known from a single adult male) are closely related but are
considered distinct because of their morphological differences and widely
separated known ranges. Terrapene coahuila, so far found only in the basin of
Cuatro Cienegas in central Coahuila, is the most primitive Terrapene known;
it differs from other box turtles in a number of morphological characters and
is the only member of the genus that is chiefly aquatic.

Two species of Terrapene occur in the United States. Terrapene Carolina,
having four recognized subspecies, has a nearly continuous distribution from
iouthom Maine, southern Michigan, and southern Wisconsin, southward to
Florida and the Gulf coast and westward to southeastern Kansas, eastern
Oklahoma and eastern Texas, and characteristically inhabits wooded areas.

Terrapene omata is a characteristic inhabitant of the western prairies of
the United States, and ranges from western and southern Illinois, Missouri,
Oklahoma, and all but the extreme eastern part of Texas, westward to south-
eastern Wyoming, eastern Colorado, eastern and southern New Mexico, and
southern Arizona, and, from southern South Dakota and southern Wisconsin,
southward to northern Mexico (Fig. 1). It is the only species of tlie genus
that occurs in both Mexico and the United States. The northeastemmost
populations of T. omata, occurring in small areas of prairie in Indiana and
Illinois, seem to be isolated from the main range of the species. The ranges
of T. omata and T. Carolina overlap in the broad belt of prairie-forest ecotone
in the central United States. Interspecific matings under laboratory conditions

Natural History of the Ornate Box Turtle

533

are not uncommon and several verbal reports of such matings under natural
conditions have reached me. Nevertheless, after examining many specimens
of both species and all alleged "hybrids" recorded in the literature, I find no
convincing evidence that hybridization occurs under natural conditions.

Terrapene ornata differs from T. Carolina in having a low, flattened carapace
lacking a middorsal keel (carapace highly arched and distinctly keeled in
Carolina), and in having four claw^s on the hind foot (three or four in Carolina),
the claw of the first toe of males being widened, thickened, and turned in
(first toe not thus modified in Carolina). Terrapene ornata is here considered
to be the most specialized member of the genus by virtue of its reduced pha-
langeal formula, hghtened, relatively loosely articulated shell, reduced plastron,
and hghtly built skull, which completely lacks quadratojugal bones (Fig. 2);
most of these specializations seem to be associated with adaptation for terrestrial
existence in open habitats.

scale of miles

Fig. 1. Geographic distribution of Terrapene ornata. Sohd symbols indicate
the known range of T. o. ornata and hollow symbols the known range of
T. 0. luteola. Half-circles show the approximate range of intergradation be-
tween the two subspecies. Triangles indicate locahties recorded in Uterature;
specimens were examined from all other localities shown. Only peripheral

locahties are shown on the map.

Two subspecies of T. ornata are recogni/ed. Terrapene o. luteola, Smith
and Ramsey (1952), ranges from northern Sonora (Guaymas) and southern
Arizona (southern Pima County) eastward to southeastern New Mexico and
Trans-Pecos, Texas, where it intergrades with T. a. ornata; the latter subspecies
is not yet known from Mexico but almost surely occurs in the northeastern part

534

University of Kansas Publs., Mus. Nat. Hist.

of that country. The subspecies luteola differs from omata in being slightly
larger and in having more pale radiations on the shell (11 to 14 radiations on
the second lateral lamina in luteola, five to eight in omata). In individuals of
luteola the markings of the shell become less distinct with advancing age and
eventually are lost; shells of most old individuals are uniform straw color or
pale greenish-brown; this change in coloration does not occur in T. o. ornata.

Fig. 2. Dorsal and lateral views of skull of T. a. omata
(a and b) (KU 1172, male, from 6 mi. S. Gamett, Ander-
son Co., Kansas) and of T. Carolina (c and d) (KU 39742,
from northern Florida). Note the relatively higher brain-
case and the incomplete zygomatic arch in T. o. omata.
All figures natural size.

Fossils

Of the several species of fossil Terrapene described (Hay, 1908b: 359-367,
Auffenberg, 1958), most are clearly allied to Recent T. Carolina. One species,
Terrapene longinsulae Hay, ( 1908a: 166-168, PI. 26) from ". . . the Upper
Miocene or Lower Pliocene . . ." of Phillips County, Kansas, however, is
closely related to T. omata (if not identical). I have examined the type speci-
men of T. longinsulae. Stock and Bode (1936:234, Pi. 8) reported T. ornata
from sub-Recent deposits near Clovis, Curry County, New Mexico.

Economic Importance

Ornate box turtles, referred to as "land terrapins" or "land tortoises" over
most of the range of the species, are regarded by most persons whom I have
queried as innocuous. These turtles occasionally damage garden crops and

Natural History of the Ornate Box Turtle 535

have been known to eat the eggs of upland game birds. Terrapene ornata is
seldom used for food. A. B. Leonard told me the species was eaten occasionally
by Arapaho Indians in Dewey County, Oklahoma. Several specimens in the
University of Kansas Archeological Collections were found in Indian middens
in Rice County, Kansas, from a culture dated approximately 1500 to 1600 a. d.
The flesh of T. ornata occasionally may be toxic if the turtle has eaten toxic
fimgi as has been recorded for T. Carolina (Carr, 1952:147).

Study Areas

Preliminary studies and collections of specimens were made at a number of
locahties in northeastern Kansas in 1953 and 1954. Two small areas were
finally selected for more intensive study. One of these areas, the University
of Kansas Natural History Reservation, five and one-half miles north-northeast
of Lawrence in the northeastemmost section of Douglas County, Kansas, is
a tract of 590 acres maintained as a natural area for biological investigations.
Shghtly less than two thirds (338 acres) of the Reservation is wooded; the
remainder consists of open areas having vegetation ranging from undisturbed
prairie grassland to weedy, partly brushy fields (Fitch, 1952). Although
ornate box turtles were not numerous at the Reservation, the area was selected
for study because: 1) there was a minimum of interference there from man
and none from domestic animals; 2) the vegetation of the Reservation is
typical of areas where T. ornata and T. Carolina occur sympatrically (actually
only one specimen of T. Carolina has been seen at the Reservation); and, 3)
availability of biological and chmatological data there greatly facilitated the
present study. Actual field work at the Reservation consisted of studies of
hibernation and long-term observations on movements of a few box turtles.

A much larger number of individuals was intensively studied on a tract of
land, owned by Sophia Damm, situated 12 miles west and one and one-half
miles north of Lawrence in the northwestern quarter of Douglas County,
Kansas. The Damm Farm lies on the southern slope of a prominence —
extending northwestward from Lawrence to Topeka — that separates the Kan-
sas River Valley from the watershed of the Wakarusa River to the south. The
prominence has an elevation of approximately 1100 feet and is dissected on
both sides by small valleys draining into the two larger river valleys.

The Damm Farm (see Pi. 15) has a total area of approximately 220 acres.
The crest of a hill extends diagonally from the middle of the northern edge
approximately two thirds of the distance to the southwestern comer. Another
hill is in the extreme northwestern comer of the study area.

The northeastern 22 acres were wooded and had small patches of over-
grazed pasture. Trees in the wooded area were Black Walnut (Juglans
nigra). Elms (Ulmus americana, U. rubra), Cottonwood (Populus deltoides),
and Northern Prickly Ash (Xanthoxylum americanum). The areas used as
pastiu-e had thick growths of Buckbush (Symphoricarpos orbiculatus) mixed
with short grasses {Bromus japonicus, Mtihlenbergia Schreberi, and Poa
pratensis). Farm buildings were situated in the wooded area at the end of
an entry road. The southeastern 74 acres were cultivated; com, wheat, and
milo were grown here and fallow fields had a sparse growth of weeds.

Most of the western two thirds of the study area, comprising 124 acres,
was open roUing prairie (hereafter referred to as "pasture") upon which beef-
cattle were grazed (Pi. 16, Fig. 1; Pi. 17, Fig. 1; Pi. 18, Fig. 2). Rock

536 University of Kansas Publs., Mus. Nat. Hist.

fences (PI. 17, Fig. 2) two to four feet high bordered the northern edge,
southern edge, and one half of western edge of the pasture. A wagon track
lead from a gate on the entry road, along the crest of the hill, to a gate in the
southern fence. Except for the latter gate and for ocassional under-cut places
in low areas, there were no openings in the rock fences through which box
turtles could pass. A few trees — American Elm, Hackberry {Celtis occiden-
talis), Red Mulberry (Morus rubra), Osage Orange (Madura pomifera). Black
Cherry (Prunus serotina), Box-Elder (Acer Negundo), and Dogwood (Comus
Drummondi) — were scattered along fences at the borders of the pasture and
in ravines. Larger trees in a small wooded creek-bed at the southwestern
edge of the pasture were chiefly Cottonwood, American Elm, Red Mulberry,
and Black Willow ( Salix nigra ) . The only trees growing on the pasture itself
were a few small Osage Orange, none of which bore fruit.

Paths were worn along fences by cattle and in several places near the
fence, usually beneath shade trees, there were large bare places where cattle
congregated. Vegetation near paths and bare places was weedy and in some
places there were tall stands of Smooth Sumac (Rhus glabra).

Rich stands of prairie grasses occurred along the top of the hill in the
pasture; bluestems (Andropogon gerardi, A. scoparius) were the dominant
species and Switchgrass (Panicum virgatum) and Indian grass (Sorghastrum
nutans) were scattered throughout. A number of small areas on top of the
hill were moderately overgrazed, as indicated by mixture of native grasses
with an association of shorter plants consisting chiefly of Ragweed (Ambrosia
artemisiifolia var. elatior), Mugwort (Artemisia ludoviciana) , Japanese Chess
(Bromus japonicus), and Asters (Aster sp.).

The upper parts of the hillsides were overgrazed moderately to heavily.
Limestone rocks of various sizes were partly embedded in soil or lay loose
at the surface. Depressions beneath rocks provided shelter for box turtles
as well as for other small vertebrates. Native grasses were sparse in this
area and gave way to Sideoats Grama (Bouteloua curtipendula) , extensive
patches of Smooth Sumac, and scattered colonies of Buckbrush.

Tall grasses were dominant on the lower hillsides and small patches of
Slough grass (Spartina pectinata) grew in moist areas. Ravines originated
at small intermittent springs on the sides of the hill. The banks of ravines
were high and steep and more or less bare of vegetation. High, dense stands
of Slough grass grew at intermittent springs and along the courses of ravines;
sedges (Carex, sp. ) grew where small pools of water formed and created
marshy conditions. Prairie grasses along the tops of ravine embankments
formed a narrow overhanging canopy of vegetation that was accentuated
in many places where the sod was undercut by erosion or by the activities
of burrowing animals (PI. 18, Fig. 1). Box turtles frequently sought shelter
beneath this vegetational canopy or burrowed beneath the sod.

On the highest part of the pasture near the entry road several small areas
were nearly bare, presumably because of heavy overgrazing; grasses (except
for scattered clumps of Bouteloua curtipendula and Setaria lutescens) were
absent and dominant vegetation consisted of Bufi^alo-bur (Solanum rostratum).
Blue Vervain (Verbena hastata). Mullein (Verbascum Thapsus), Ragweed,
Asters, and a few Prickly Pear (Opuntia humifusa). Two small areas on the
pasture completely lacked vegetation; these may have been wallows or the
sites of old salt-licks.

Natural History of the Ornate Box Txjrtle 537

Three shallow stock ponds, behind earthen dikes in ravines, were present
on the pasture. The pond near the farm buildings ("House Pond") and
that in the southwestern part of the pasture ("Far Pond") were present when
studies of box turtles were begun. The largest pond, in a deep ravine in
the northern part of the pasture, was constructed in June, 1956, and became
filled in approximately one month (Pis. 16 and 18). Pond embankments
were chiefly bare of vegetation because of trampling by cattle; in a few places
at the edge of the water, or in places too steep for cattle to walk, there were
small patches of weeds, sedges, and Slough Grass. The ponds contained some
water at all times of the year. The only vertebrates permanently inhabiting
the ponds in the course of my studies were Bullfrogs {Rana catesbeiana) and
Leopard frogs {Rana pipiens).

The three parts of the pasture in which studies were concentrated were
designated as separate subdivisions. The northwest corner area (28 acres)
was triangular and bounded on two sides by rock fences and on its third side
by a deep ravine. The southern ravine area ( 17 acres ) constituted the part of
the lower southern hillside drained by a series of ravines. The house pond
area (seven acres) surrounded "House Pond." Habitat in these three sub-
divisions of the pasture was especially favorable for box turtles.

Materials and Methods

Observations were made at the Damm Farm on 102 days in the two-year
period beginning in Autumn, 1954; observations were concentrated in the
period from May to October although some observations were made in every
month, January and February excepted. Field work was done chiefly in day-
light hours but a few trips were made to the study area at night.

Routine handling of each turtle captured at the Damm Farm consisted of:
marking, weighing and measuring turtle; recording the exact place of capture,
body temperature and environmental temperature; and, recording miscellaneous
items such as the presence of ectoparasites, injuries, distinctive markings, and
in some instances, the approximate age of the turtle.

Excursions on the Damm Farm were made on foot in 1954 and 1955, and,
in 1956, on horseback. By using a horse, more ground could be covered per
unit of time, a better view could be obtained of immediate surroundings, and,
cattle on the area, being accustomed to horses, did not become agitated as
they would when unmounted persons were nearby.

The entire study area could not be inspected thoroughly in a single day.
It was usually more profitable to find and mark turtles along fences, in ravines,
or in other open areas, and subsequently to follow their movements away from
these areas by means of trailing threads. Turtles could be observed from a
distance through binoculars. Cultivated areas were regularly scanned with
binoculars but turtles were seldom seen there. Behavior was observed by
sitting motionless on rock fences or in a blind on top of a stepladder.

No box tiutles were removed from the study area. Specimens obtained in
other areas were used for studies of growth, reproduction, and food habits.
Measurements, weights, and data concerning temperature and ectoparasites
were obtained from specimens collected elsewhere as well as from individuals
on study areas.

Turtles were obtained by hand-collecting and in unbaited traps; the number
captured in a single day ranged from 12 to none. Traps, like those used by

538 Unatrsity of Kansas Publs., Mus. Nat. Hist.

Packard (1956:9) for tree squirrels, were set in the mouths of burrows and
dens, or — with leads to channel animals into the trap — along ravines and rock
fences. Traps set in the open were covered to prevent death of turtles from
overheating in direct sunbght. Live-trapping provided much valuable data,
although quail, rabbits, opossums, and box turtles were caught with about
equal frequency in the traps.

Turtles were marked by notching the marginal scutes of the carapace by
means of a hacksaw blade, following the code system described by Cagle
(1939). Notches, one eighth to one quarter of an inch deep and wide could
be cut more quickly than filed and were more evident than drilled holes which
often became plugged with soil and obscured. Hatchlings and juveniles were
notched with a sharp knife.

Movements of individual turtles were studied by means of a turtle-traihng
device — similar to the kind first described by Breder ( 1927 ) and later modified
by Stickel (1950:355-356) — a tin can, cut to fit the shell of a turtle, with an
axle that bore a spool of thread ( PI. 27, Fig. 1 ) . The device was taped to the
turtle; the free end of the thread was tied to a stationary object. Thread payed
out from the spool through a guide-loop and marked the course of the turtle as
it moved away from the starting point. Because of its great strength and
elasticity (as compared to cotton), nylon sewing thread was used in trailers.
Ordinarily, turtles were unable to break the thread if it became snarled or was
expended. Cattle frequently tangled the thread and displaced it but did not
often break it. Ordinary spools were cut down on a lathe so they would hold
600 to 800 yards of thread. Turtle-trailing provided an accurate record of
where and how far a turtle had traveled, and to a lesser extent, the sort of
activity in which the turtle had been engaged (evidence of feeding, forms, or
trial nest holes). Trailers seemed not to alter the normal activity of turties.

Prominent landmarks were rare or wanting in most places on the pasture.
Locations of captures (or reference points in the movements of trailer-turtles)
were determined by triangulation with a Brunton compass, using trees along
fences as known points of reference. Rough maps were made in the field and
used later, along with compass readings and measurements, to make a more
precise record of movements and captures on a large map (scale, 100 feet to
one inch) of the study area. Mapped points of capture in grassy areas were
accurate within ten to twenty feet; points of capture in areas where landmarks
were nearby were nearly exact. Areas were measured with a planimeter;
distances traveled by individuals were measured with a cartomcter.

Turtles were measured in the field to the nearest milUmcter with large
wooden calipers (of the type used by shoe salesmen) and a clear plastic ruler.
Measurements in the laboratory, especially in studies of growth, were made,
to the nearest tenth of a millimeter with dial calipers. Measurements made
on each specimen examined in the field were: length of carapace, width of
carapace, length of plastron (sum of lengths of forelobe and hind lobe), width
of plastron (at hinge), and height. All measurements were made in a straight
line. A spring scale of 500 gram capacity, used in the field, gave weights
accurately within three grams. A triple-beam balance was used in the lab-
oratory. Unless otherwise noted, measurements are expressed in miUimetcrs
and weights are expressed in grams.

Body temperatures were taken by means of a quick-reading Schultheis
thermometer inserted into the distal portion of the large intestine with the

Natural History of the Ornate Box Turtle 539

bulb directed ventrally to avoid puncturing the bladder. Body temperature of
turtles were altered little or not at all in the few seconds the turtles were held
and no attempt was made (except for small juveniles) to insulate them from
the warmth of my hands. Data recorded with body temperature were: air
temperature (in shade, approximately one inch from turtle); ground tempera-
ture (or water temperature); behavior of turtle; weather conditions; natvure of
vegetation or other cover; and, time of day. Unless otherwise noted, tempera-
tures are ex-pressed in degrees Centigrade.

A maximum-minimum thermometer was installed near the buildings at the
Damm Farm. Notes on general weather conditions were made on each visit
to the study area. Additional climatological data were obtained from the
U. S. Weather Stations in Topeka and Lawrence, from records at the Reserva-
tion, and from oflBcial bulletins of the U. S. Weather Bureau.

Stomachs and gonads were removed and preserved by standard techniques
soon after specimens were killed. The dates given to gonads were, in all in-
stances, the dates when the specimens were killed. Eggs were prepared for
incubation in the manner described by Legler (1956). Females laying or con-
taining eggs used in studies of incubation were preserved for fiu±her studies
and comparison with young hatched from the eggs. Histological preparations
were fixed in ten per cent formahn or Bouin's fluid, embedded in paraffin, and
stained with hematoxahn and eosin.

Terminology

Names used for the epidermal and bony parts of the shell follow the classi-
fication proposed by Carr (1952:35-39). The terms "scute," 'lamina," and
"scale" are used here more or less interchangeably for the epidermal parts as are
the terms "plate," "bone," and "element" for the bony parts of the shell.

The term "form" is used here in the same sense that Stick el (1950:358)
used it in her study of T. Carolina — to indicate a depression or cavity made by
a turtle in vegetation or soil. Forms correspond closely in shape and size to
shape and size of the turtle. Forms of T. ornata differ from those of T. Carolina
chiefly in being made most often in soil, over which there is a minimum of
vegetational cover. The term "den" refers to natural cavities (or cavities of
unknown origin) beneath rocks, in rock fences, or in cut banks. The term
"burrow," unless otherwise noted, refers to burrows made by animals other
than box turtles.

HABITAT AND LIMITING FACTORS

The known range of T. ornata includes the southern half of the
Grassland Biome, part of the Desert Biome, and that part of the
Temperate Deciduous Forest Biome known as the Prairie-Forest
Ecotone. The species is found in microhabitats that differ widely in
food supply, temperature, moisture, and kind of soil. In spite of its
relatively high degree of morphological specialization, T. ornata is
remarkably versatile in regard to habitat requirements.

Ornate box turtles are relatively inconspicuous in natural sur-
roundings and collectors seldom seek out and obtain specimens
under completely natural conditions as may be done with certain

540 University of Kansas Publs., Mus. Nat. Hist.

other reptiles and amphibians by turning rocks, tearing apart logs,
or setting traps. Most series of specimens are obtained by hunting
after rains on roads or other natural breaks in vegetational cover.
Detailed information on habitat preferences is lacking.

Low temperature seems to be an important factor limiting the dis-
tribution of T. ornata in the northern part of its range. Box turtles,
hke nearly all other reptiles occurring at these latitudes, spend the
winter in underground hibemacula. The depth to which the ground
freezes in the coldest part of the winter is therefore a critical factor.
The ground freezes to an average depth of 30 inches or less over
most of the range of the species; only in the extreme northern part
of the range (southern South Dakota, southeastern Wyoming) does
the ground freeze to an average depth of as much as 35 inches.
Average depth of freezing is, in fact, less than 15 inches over more
than one half the range of the species. The average number of
frost-free days per year ranges from 130 to 140 days in the northern
part of the range to more than 250 days in the southwestern part of
the range.

Terrapene ornata occurs from near sea level to elevations of more
than 5000 feet. Both subspecies are found at both high and low
elevations but luteola is more consistently taken at high elevations
than ornata. The latter subspecies commonly occurs at elevations
above 4000 feet on the high plains in extreme western Kansas and
eastern Colorado; the highest elevation from which I have examined
specimens of T. o. ornata is between 4600 and 4700 feet near Akron,
Washington County, Colorado. The greater part of the known
range of T. o. luteola lies above 3000 feet.

Norris and Zweifcl (1950:1) observed T. o. luteola on the Jor-
nada del Muerto, an elongate plain approximately 4500 feet high,
in southeastern Socorro County, New Mexico; box tm-tles were
abundant on the level part of the plain and on the bordering foot-
hills but not at higher elevations where the substratum was rocky.
The authors otherwise noted no preference for any kind of soil.
The principal elements of the plant associations in which the turtles
were found were creosote bush, yucca, mesquite, juniper, tarbush,
and grasses. Lewis (1950:3) reported that T. ornata luteola in-
habited the yucca-grassland zone in Dona Ana County, New Mex-
ico; he stated {op. cit.: 10) that individuals were commonly found
on roads after rains and in cloudy weather. No specimens were
taken at altitudes higher than 4300 feet.

I have examined specimens of luteola from elevations of approxi-
mately 5500 feet in Cochise County, Arizona, and Lincoln County,

Natural History of the Ornate Box Turtle 541

New Mexico. These localities are probably at or near the maximum
elevation at which the species occurs. The texture of the substrate
is the most important factor limiting vertical distribution. Ornate
box turtles, like nearly all other turtles, excavate nests; T. ornata is
a burrower, at least for purposes of hibernation. Populations of the
species, therefore, could not survive in areas of hard unyielding
substrata. Such substrata seem to be the most important factor
limiting altitudinal distribution.

Most of the area in which T. ornata occurs is semiarid or arid.
Average precipitation in the warm season (April through Septem-
ber) varies from approximately 25 inches in the northeast to less
than ten inches in the southwest. In drier parts of the range,
precipitation is unevenly distributed over the warm season. Long,
hot, dry periods are unfavorable for reptihan activity. T. ornata,
like many other reptiles inhabiting dry regions, survives long pe-
riods without water by seeking shelter (usually underground) and
remaining quiescent. Populations of the subspecies luteola live
under far more rigorous conditions in this respect than do the
more northern populations. Specimens of luteola from Arizona
that were kept for several years in the laboratory under dry con-
ditions and fed adequately, but at infrequent intervals, were able
to remain healthy and even to grow whereas examples of ornata
kept under the same conditions soon languished and died; luteola
seems to be physiologically adapted for existence under arid con-
ditions, where normal activity is sometimes possible for only a
few weeks in the year.

The prairies of Nebraska, Kansas, Oklahoma, and northern Texas
seem to provide the most nearly optimum habitat for the species;
in these regions box turtles are active on a large majority of the
days from April to October in years having average or better than
average precipitation and population density seems to be greater
than in tlie more arid parts of the range.

Activities of man have probably affected the density of popula-
tions of the ornate box turtle in many parts of its range but appear
not to have acted as limiting factors except in certain areas along
the northern edge of the range (Blanchard, 1923:19-20, 24) where
disruption of grassland through intensive cultivation probably has
excluded the species. Unlike certain other reptiles of the Great
Plains (Fitch, 1955:64), T. ornata seems not to have been affected
— either by direct decimation of populations or by disruption of
habitat — by intensive zoological collecting in restricted areas. En-
vironmental changes such as those resulting from overgrazing and

542 University of Kansas Publs., Mus. Nat. Hist.

erosion, or from protection of tlie habitat from grazing could be
expected to cause long-term changes in populations of ornate box
turtles.

Terrapene o. omata is an omnivorous, opportunistic feeder, pri-
marily insectivorous but able to subsist on nearly any sort of ani-
mal or vegetable food. The general food habits of luteola are
poorly known but probably resemble those of omata. Although
kind of food available probably does not limit the distribution of
T. omata there are indications that it influences population density.
In Kansas, for example, dung insects are an important staple in the
diet and box turtles were found always to be more numerous in
areas where domestic cattle provided an abundant supply of dung
than elsewhere. A similar relationship probably existed in former
times between box turtles and native ungulates. Near extinction
of buffalo in the Great Plains possibly caused a decrease in popula-
tions of box turtles, Henry S. Fitch told me that the number of
T. omata at the Reservation gradually declined after cattle were
removed from the area in 1948.

In summary, the distribution of T. omata seems to be limited
by: 1) Presence of a substrate too hard to permit digging of nests
and forms (southwestern and western edges of range); 2) tem-
peratures causing the ground to freeze deep enough (approxi-
mately 30 inches) to kill turtles in hibernacula (northern edge of
range); and, 3) the lack of one or more relatively wet periods in
the course of the warm season, preventing at least temporary emer-
gence from quiescence (southwestern edge of range).

HABITAT IN KANSAS

Clarke (1958:40-45) reported T. o. omata in all terrestrial com-
munities studied in Osage County; he considered the subspecies
to be characteristic of the ". . . cultivated-field community
. . ." and to be of frequent occurrence in (but not characteristic
of) the ". . . Oak- Walnut Hillside Forest . . ., Buckbrush-
Sumac . . ., and Prairie communities . . .". Brennan
(1937:345) found T. o. omata to be equally abundant in mixed
prairie and prairie-streamside habitats in Ellis County; the sub-
species was much rarer on rocky hillsides and in the habitat sur-
rounding prairie ponds. Carpenter (1940:641) listed T. o. omata
as an inhabitant of ". . . tall and mixed-grass prairies . . ."
(also in Oklahoma and Nebraska). Fitch (1958:99) found the
order of preference for habitats at the Natural History Reservation
to be grazed pasture land, woodland, open fields with undisturbed

Natural History of the Ornate Box Turtle 543

prairie vegetation, and fallow fields with a rank growth of weeds.

At the Damm Farm the greatest number of box turtles was
collected on the pasture, especially in three areas designated in
Plate 1 as the "northwest corner," "southern ravine," and "house
pond" areas. These three areas had several features in common.
All contained ravines and rocky slopes that provided many places
of concealment (dens, burrows of larger animals, and suitable sub-
strate for the excavation of earthen forms). All contained water
(in ponds and intermittent streams) for most of the year; and, all
were frequented daily by cattle that left an abundant supply of
dung in which box turtles foraged. In addition, each of the three
areas contained at least one mulberry tree, under which fruit was
abundant in the months of June and July.

The relative numbers of box turtles found in difiFerent areas on
the Damm Farm were, of course, governed to some extent by my
activity in these areas and by the relative ease with which box
turtles were seen in different types of vegetational cover. Turtles
were more easily seen in the pasture (especially in sparsely
vegetated or denuded areas) where much of my field work was
done on horseback, than in the wooded areas, where excursions
were usually made on foot. It was evident, however, after mapping
known ranges and studying patterns of movement in marked turtles,
that concentrations in the three above-mentioned areas of pasture
were an indication of actual preference by turtles for the more
favorable habitat in these areas rather than the result of incomplete
sampling.

REPRODUCTION

Mating

Mating takes place throughout the season of activity but is most
common in spring — soon after emergence from hibernation — and
in autumn. Turtles frequently copulated in the laboratory in spring
and autumn. Copulation was observed under natural conditions
on several occasions but only once at the Damm Farm.

Norris and Zwiefel (1950:4) saw two captive individuals of T. o.
luteola copulating on 12 August; copulation lasted two hours.
Brumwell (1940:391-2) gave the following description of mating in
T. o. ornata. A male pursued a female for nearly half an hour, first
nudging the margins of her shell and later approaching her rapidly
from the rear and hurHng himself on her back in an attempt to
mount, at the same time emitting a stream of liquid from each
nostril. The liquid was presumably water; both sexes had imbibed

2—773

544 University of Kansas Publs., Mus. Nat. Hist.

water in a pond just before courtship began. Brumwell suggested
that pressure on the plastron of the male had forced the water out
his nostrils. The pair remained in the coital position for 30 minutes
after the male had achieved intromission. In anotlier instance,
Brumwell {loc. cit.) saw four males pursuing a single female, the
males exhibiting the same behavior ( nudging and lunging ) outlined
above. Males that attempted to mount other males were repelled
by defensive snapping of the approached male. The female also
snapped at some of the males that tried to mount her. One male
was finally successful in mounting and was henceforth unmolested
by the other males. Brumwell suggested that shell biting and
tapping may be methods of sex-recognition.

In the several instances of mating that I observed, the male,
after mounting the shell of the female (PI. 28), gripped her, with
the first claws of his hind feet, just beneath her legs or on the skin
of the gluteal region and, with the remaining three claws, gripped
the posterior edges of her plastron. In most instances the female
secured the male's legs by hooking her own legs around them.
The coital position of T. ornata seems to differ from that of T.
Carolina, at least in regard to the position of the male's legs. The
coital positions of T. Carolina illustrated by Cahn (1937:94, Fig.
13) are physically impossible for T. ornata.

In T. ornata the pressure exerted on the male's legs by the female
probably impairs circulation and probably is painful to the male,
especially after coitus, when the male falls backward but is still
held by the female. The heavily developed musculature of the
legs of males may be an adaptation to strengthen the legs for this
temporary period of stress. Evans (1953:191) and Cahn and
Conder (1932:87-88) observed the hind legs of males of T. Caro-
lina to be noticeably weakened after copulation, causing the males
to remain inactive for several hours.

Evans ( op. cit. ) observed 72 matings of T. Carolina and divided
the process into three phases as follows: 1) circling, pushing and
biting by the male; 2) mounting (female with shell closed); and,
3) coition (female with shell open). Penn and Pottharst (1940:26)
reported that captive T. Carolina in New Orleans mated chiefly
under conditions of optimum temperature (21 to 27° C. ) and high
humidity; some matings took place in a pool of water. Males pushed
females about after mating, often rolling them over several times.

Because ornate box turtles observed by me were able easily to
right themselves from an inverted position on substrata of all

Natueal History of the Ornate Box Turtle 545

kinds, males left lying on their backs after copulation are probably
in no danger of perishing in this position, as was suggested by
Allard (1939) for T. Carolina.

Insemination

Oviducts of several females w^ere flushed by means of a pipette
to determine w^hether they contained sperm. Approximately half
of die females captured in May, 1956, had sperm in their oviducts,
but females captured in June and July did not. Sperm flushed from
the oviducts were in clumps of several hundred and showed no
sign of motility a few minutes after the female was anesthetized
with chloroform. No sperm were found in the oviducts of imma-
ture females but one female of nearly adult size was observed in
copulation with a mature male.

Thorough examination of microscopic sections of oviduct (taken
at various times in the season of activity) usually revealed a few
sperm lodged in the folds (PI. 19, Fig. 8) of the cephalic as well
as the caudal portion of the tube, but no specialized seminal re-
ceptacles such as occur in snakes (Fox, 1956) were present. Fer-
tilization without reinsemination probably occurs in T. ornata.
Evidng (1943) and Finneran (1948:126) reported that females of
T. Carolina produced fertile eggs for periods of four and two years,
respectively, after being removed from all contact with males.

Sexual Cycle of Males

Testes were preserved in each month from April to October. The
following description of spermatogenesis is based chiefly on material
collected in 1955, although testes were preserved also in 1954.
Comparison of material obtained in 1954 and 1955 revealed that
spermatogenesis began earlier and was more advanced on any
given date in 1955 than in 1954.

Testes of mature individuals are pale yellow and slightly oblong.
The epididymis is ordinarily dark brown or black and contrasts
sharply with the color of the testes. Size of testes was expressed as
the average lengtii (greatest diameter) of both testes. Testes are
smallest in April, immediately after emergence from hibernation,
and largest in early September (PI. 20, Figs. 3-4). They are nearly
spherical when of maximum size; increase in bulk, therefore, is
relatively greater than the increase in size shown in Figure 3. They
increase in size from April until early June, recede during most of
June, and again increase in size in July and August. They remain

546

University of Kansas Publs., Mus. Nat. Hist.

large from early September until hibernation is begun, becoming
only slightly smaller in late September and October,

Increase in size following emergence from hibernation may be
due in part to proliferation of the sustentacular cytoplasm. De-
crease in size in early June is correlated with the end of the period
of most active mating; maximal size is coincident with the peak of
the spermatogenic cycle in early September.

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April

May

June July

Aug.

Sept. Oct.

Fig. 3. Seasonal fluctuations in size (average greatest diameter) of testes
in T. o. ornata as determined by examination ot 40 specimens from eastern

Kansas.

Spermatogenesis {refer to PI. 19, Figs. 1-5) begins in early May
when a few spermatogonia appear in the seminiferous tubules. The
histological appearance of testes preserved in April and May is much
the same. Nuclei of Sertoli cells, which outnumber the sperma-
togonia, are evident at the periphery of the tubules and the clear
cytoplasm of the cells extends into and nearly fills the lumina. The
few darkly stained spermatids that are present in April are cells
that probably were produced in the previous summer. Sperm are
present in small groups within the sustentacular cytoplasm, but
ordinarily are absent in the lumina.

Natural History of the Ornate Box Turtle 547

Primary spermatocytes appear in the tubules from mid-May to
early June. By mid-May there are practically no sperm at any
place in the tubules. The sustentacular cytoplasm has a less com-
pact arrangement in late May than in April.

Spermatogenesis is well under way by mid-June; at this time,
two or tliree distinct layers of primary and secondary spermatocytes
are present and these cells outnumber the Sertoli cells. The lumina
are filled with cellular detritus and are no longer bordered by a
clear ring of sustentacular cytoplasm. No sperm are present.

Spermatids appear in late June and a few of them undergo meta-
morphosis in early July; by mid-July, spermatids and secondary
spermatocytes are the dominant cells in the seminiferous tubules,
although spermatogonia are still active.

By late August, clusters of sperm and metamorphosing spermatids
surround the Sertoli cells; large numbers of sperm as well as
sloughed cells representing various spermatogenic stages are present
in the lumina. Secondary spermatocytes are still evident near the
periphery of the tubules but they are much less numerous than
spermatids. The germinal epithelium is still semiactive and small
groups of primary spermatocytes are present in nearly all of the
tubules.

The spermatogenic cycle is completed in the latter half of October
when most of the spermatozoa pass into the epididymides. A few
spermatozoa and spermatids remain in the seminiferous tubules
during hibernation. Although no testicular material was obtained
from hibernating turtles, comparisons of sections made in October
and April show that the germinal epithelium remains inactive from
autumn until spring. Possibly some spermiogenesis takes place in
the early phases of hibernation or in the period in late autumn
when turtles are intermittently active. It is uncertain whether
the reorganization of the sustentacular cytoplasm occurs in autumn,
in spring, or in the course of hibernation.

The seminiferous tubules of immature males are small, lack
lumina, and contain a few large but inactive spermatogonia ( PI. 19,
Fig. 6 ) . The testes of specimens that were nearly mature contained
primary and secondary spermatocytes but lacked lumina; it was
thought that such individuals would have matured in the following
summer and bred in the following autumn.

Mature sperm were found in epididymides at all times of the year
but were most numerous in spring and autumn, the period between

548 University of Kansas Publs., Mus. Nat. Hist.

spermatogenic cycles (PI. 19, Fig. 7). Sperm expelled from the
epididymides in autumn matings are seemingly replaced by others
from the seminiferous tubules; the epididymides become much
smaller when their supply of sperm is nearly exhausted after spring
mating.

Risley (1938:304) found the testes of the common musk turtle,
Sternotherus odoratus, to be largest in August and smallest in early
May. Recession of testes in spring was coincident with the period
of active breeding; increase in size, later in the season, corresponded
to increasing spermatogenic activity and enlargement of seminifer-
ous tubles. Altland (1951:600-603) found the spermatogenic cycle
of Terrapene coroUna to be nearly like that of Sternotherus odoratus.
Fox (1952) found that testes of garter snakes (Thamnophis sirtalis
and T. elegans) in CaUfornia reached a peak of spermatogenic
activity in midsummer, regressed in the latter half of the summer,
and were inactive in winter.

The spermatogenic cycle of T. ornata as here reported, differs
in no important respect from those of Thamnophis, Sternotherus
odoratus, or Terrapene Carolina, except that in T. ornata the cycle
begins and ends somewhat later in the season of activity. In most
of the lizards that have been studied (Fox, 1952:492-3), spermato-
genesis reaches a peak in spring (more or less coincident with the
mating period and with ovulation) and the germinal epithelium re-
mains active in winter. Sternotherus, Terrapene, and Thamnophis
are alike in completing spermatogenesis late in the season and stor-
ing spermatozoa, in the seminiferous tubules or in the epididymides,
during hibernation.

It is noteworthy that, in the turtles and snakes mentioned above,
sperm produced in autumn are used to fertilize eggs laid in the
following year, and mating [with the exception of Thamnophis
elegans, (Fox, 1956)] occurs in both spring and autumn. It is not
definitely known in any of these instances, whether sperm resulting
from autumn or spring inseminations (or both) fertilize the eggs.
Risley (1933:693) found motile sperm in the oviducts of female
Sternotherus odoratus that had recently emerged from hibernation;
he believed that spring mating, although it commonly occurred,
was not necessary to fertilize eggs. Disadvantages, if any, of com-
pleting spermatogenesis well in advance of ovulation seem to be
at least partly counteracted by two annual mating periods or by
mating throughout the season of activity.

Natural History of the Ornate Box Turtle

549

Sexual Cycle of Females

The following account of oogenesis is based on examination of
preserved ovaries from 68 mature specimens. The ages of most
specimens were known, inasmuch as the specimens were used in
studies of growth as well as gametogenesis. Other data were ob-
tained from adult females that were dissected but not preserved,
and from immature females.

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MARCH

APR.

MAY

JUNE JULY AUG. SEPT. OCT.

Fig. 4. Seasonal fluctuations in ovarian weight in T. o. ornata, as determined
by examination of 60 specimens from eastern Kansas.

Size of ovarian follicles was determined by means of a clear
plastic gauge containing notches 5, 10, 15, 20, and 25 millimeters
wide. The number of foUicles within a given size range could
be quickly determined by finding the smallest notch into which the
follicles fit. It was necessary to weigh all ovaries after preserva-
tion since some of them had not been weighed when fresh. Since
all ovarian samples were preserved in the same manner, weights

550

Unr'ersity of Kansas Publs., Mus. Nat. Hist.

remained relatively the same. Preserved material was lighter than
fresh by an average of 13 per cent. Follicles less than one milli-
meter in diameter were not counted. Corpora lutca and corpora
albicantia were studied under a binocular dissecting microscope.
No histological studies were made of the female reproductive sys-
tem.

Ovarian follicles and oviducal eggs were recorded separately for
the right and left sides. Each ovary was always kept associated
with the oviduct of the same side, but in some instances it was
not recorded whether the organs were left or right.

Ovaries ordinarily weighed most in October, March, and April,
when most females contained enlarged follicles, and least in August
and September when the supply of enlarged follicles was usually
exhausted (Figs, 4 and 5),

percentoge of females with
two or more large follicles

UI o en

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(4)

HTJO)

(17)
H(t4)

(5)
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March
April

May

June

July
Aug.

Sept.

Oct.

Fig. 5. The seasonal occurrence of enlarged ovarian
follicles in females of 7'. o. ornata, expressed, for each
month, as the percentage of total females that con-
tained two or more follicles having diameters gro;>.ter
than 15 mm. Total numher of females in each of the
samples is shown in parentheses at the top of each bar.

The ovarian cycle begins in July or August, after ovulation has
occurred. At that time many minute follicles form on the germinal
ridges of the ovaries. On the basis of the material that I examined,
it seems that ovarian follicles either grow to nearly mature size
in the season preceding ovulation and remain quiescent over win-
ter or grow rapidly in the period of approximately six weeks be-
tween spring emergence and ovulation, Altland (1951:603-5) re-

Natural History of the Ornate Box Turtle 551

ported that the former condition was the usual one in T. Carolina;
he suggested that possibly some of the enlarged follicles were
absorbed during hibernation.

Examination of yolks of oviducal eggs revealed that follicles
mature when they reach a diameter of 16 to 20 millimeters and a
weight of two to two and one-half grams ( Pi. 20, Fig. 1 ) .

The enlarged follicles remaining on the ovaries after ovulation
(excluding those smaller than six mm.) can be grouped according
to diameter as: large (greater than 15 mm.), medium (11 to 15
mm.), and small (six to 10 mm.). Ten females collected in the
period from June 2 to 8, after they had ovulated, all had foUicles
falling in at least one of these size groups, and eight had follicles
falling in two or more of the groups. In females having enlarged
follicles of more than one of the size groups, there were several
follicles in each of two groups and no follicles, or only one folHcle,
in the remaining group. Enlarged follicles represent future clutches
but whether the enlarged follicles will be ovulated in the same
season or in a later season is questionable.

Evidence found in the present study suggested diat at least a
few females lay more than one clutch of eggs per year. Among 34
specimens obtained in June and July, eight (24 per cent) had
corpora lutea (or easily discernible corpora albicantia) and at least
two folhcles more than 15 miUimeters in diameter; in three speci-
mens ( 9 per cent ) the ovaries bore fresh corpora lutea ( representing
recent ovulations) and a set of older corpora lutea (representing
ovulations that had occurred several weeks previously). It was
thought that each of these eleven females (33 per cent of sample)
had produced or would have produced two clutches of eggs in the
season of its capture. The number of large folhcles present after
the first set of ovulations (mean, 3.5) was fewer in most instances
than the average clutch-size (see below), indicating that second
clutches are smaller than first clutches. Smaller second clutches
were found also in T. Carolina (Legler, 1958).

Further evidence for multiple clutches was the absence of en-
larged ovarian follicles in some females obtained in September.
Atretic follicles, ordinarily orange, brown, or purplish, were ob-
served on the ovaries of many of the females examined; in most
instances, not more than two follicles of the small or medium size
groups were atretic. Atresia was in no instance great enough to
account for the complete loss of enlarged follicles.

Further study probably will show that many of the females laying

552 UNivERSiTi' OF Kansas Publs., Mus. Nat. Hist.

in May and early June lay again before the end of July, and that
eggs in the oviducts of females captured in the latter month fre-
quently represent second clutches. Under favorable conditions,
eggs laid by the end of July would have a good chance of hatching
before the advent of cold weather in autumn; turtles hatching too
late to escape from the nest could burrow into its sides and
probably escape freezing temperatures.

Cagle's findings concerning Pseudemijs scripta (1950:38) and
Chnjscnujs picfa (1954:228-9) suggest that these species lay more
than one clutch per season, at least in tlie southern parts of their
ranges. Carr (1952) indicated that multiple layings were known
in most species of marine turtle.j (families Dermochelydae and
Chelonidae) and strongly suspected in other species. Other turtles
recorded to have produced multiple clutches in a single season
(based chiefly on captive specimens or cultured populations) in-
clude: the starred tortoise, Geochelone elegans ( Deraniyagala,
1939:287); the Asiatic trionychid, Lissemys punctata {op. cii.: 304);
the diamond-backed terrapin, Malaclemys terrapin (Hildebrand
and Prytherch, 1947:2); and the Japanese soft-shelled turtle, Trionyx
japonicus (Mitsukuri, 1895, cited by Cagle, 1950:38).

There is a marked alternation of ovarian activity in T. ornata, one
ovary being more active than its partner in a given season. The less
active ovary is more active than its partner in the following season.
For example, a specimen killed in July had four corpora lutea on
the right ovary and two on the left and there were five enlarged
follicles (of the medium size group), representing the next set of
eggs to be ovulated, four on the left ovary and one on the right.
Similar alternation of ovarian activity was observed, to a greater or
lesser extent, in nearly all of the females examined. Many subadult
females that were approaching their first breeding season (as
evidenced by the presence of large ovarian follicles but no indica-
tion of fonner ovulation) had but one active ovary. This may ac-
count in part for the tendency of small, young females to lay
clutches smaller than average. One ovary may become senile in
old females before its partner does; this may explain the occasional
absence or atrophy of one ovary in large females that I have
examined.

In all the specimens examined, it was evident that ovulation had
occurred or would occur in two successive seasons. Senile or young
females might, however, be expected to skip a laying season if only
one ovary was functioning.

After ovulation, the collapsed follicle assumes a cuplike shape

Natural History of the Ornate Box Turtle 553

and becomes a glandular corpus luteum (PI. 20, Fig. 2). Corpora
lutea are approximately eight millimeters in diameter and are easily
discernible at least until the eggs are laid; they are somewhat less
distinct after preservation. Corpora lutea undergo rapid involution
following oviposition and, after two to tliree weeks, are little more
than small puckerings on the ovarian epithelium. At this stage
they are properly referred to as corpora albicantia and are dis-
cernible only after careful examination of the ovary under low
magnification. Corpora albicantia remain on the ovary until April
of the year following ovulation but disappear in May and are never
present after the new set of eggs is ovulated. Ovaries of some sub-
adults (that would have laid first in the season following capture)
contained enlarged follicles and, but for their lack of corpora lutea
and corpora albicantia, were indistinguishable from those of older,
fully mature females.

Altland (1951:605-610) gave a histological description of the
corpus luteum of Terrapene Carolina. Corpora lutea were glandular
and filled with lipoidal material until the eggs were laid. Atresia
of corpora lutea began when eggs were laid, was completed by mid-
August, and was coincident with atresia of large follicles that did
not undergo ovulation. Altland did not describe the gross external
appearance of the corpus albicans.

The corpus luteum of oviparous reptiles seems to be closely as-
sociated with the intrauterine life of the eggs and, in viviparous
reptiles, it may be an important factor in maintaining optimum
gestational environment; however, its functions in all reptiles are
poorly understood (Miller, 1948:200-201).

Information gleaned from records of gravid females and known
dates of nesting suggests that eggs are retained in the oviducts two
to three weeks before laying. Once they are ovulated, the eggs
ai'c exposed to but few hazards until laid; counts of corpora lutea
are an accurate indication of the number of eggs laid. In the gravid
females examined by me, number of corpora lutea on the ovaries
was equal, in all but one instance, to the number of oviducal eggs.
In the single instance in which an extra corpus luteum was found,
one egg had probably been laid before the specimen was captured.
The high incidence of correspondence between counts of corpora
lutea and counts of oviducal eggs indicates also that T. ornata de-
posits the entire complement of oviducal eggs at one time, not
singly or in smaller groups.

Extrauterine migration of ova, whereby eggs from one ovary pass
into the oviduct of the opposite side, is of common occurrence in

554 University of Kansas Publs., Mus. Nat. Hist.

T. ornata and is known to occur also in T. Carolina, Chrysemys
picta, Emydoidea blandingi, Fseudemys scripta, Cnemidophorus
sexJincatus, and in several mammals (Legler, 1958). This ovular
migration may serve to redistribute eggs to the oviducts when the
ovaries are functioning at unequal rates.

The eggs acquire shells soon after they enter the oviducts. No
shell-less eggs were found in oviducts but several specimens of
T. ornata had oviducal eggs, the thin, parchmentlike shells of which
lacked the outer calceous layer; in these specimens the corpora
lutea were fresh, probably not more than two days old. Eggs that
had remained in the oviducts longer had a calceous layer on the
outside of the shell. Eggs having incompletely developed shells
were successfully incubated in the laboratory. Cagle (1950:38)
found shelled but yolkless eggs in the oviducts of several Fseudemys
scripta but found no yolkless eggs in nests. No yolkless eggs were
found in specimens of T. ornata in the course of the present study.

The uterine portion of the oviducts becomes darkened (pale
gray to intense black) in the breeding season. Darkening of ovi-
ducts seemed to coincide with the period when eggs were in the
oviducts and it persisted for a variable length of time after the
eggs were laid. Oviducts of immature females were ordinarily pale.

Nesting

Ornate box turtles nest chiefly in June. Some females nest as
early as the first week of May or as late as mid-July but the nesting
season reaches its peak in mid-June. Eggs nearly ready to be laid
were in oviducts (determined by bimanual palpation in the field
or by dissection in the laboratory) of many females captured in
June; nearly half of the records so obtained were in the second
week of that month. Early records of shelled oviducal eggs were
April 25 (specimen from Ottawa County, Oklahoma), May 5, and
May 22. The two latest records are for females retaining oviducal
eggs on July 2 and 11. Known dates for nesting of free-living fe-
males were distributed rather evenly through the month of June.
It is worthy of note that all (four) of the nestings known to occur
in July were by captive females. Females of T. ornata, like those
of some other turtles (Cagle and Tihen, 1948; Risley, 1933:694),
seem to retain their eggs until conditions are suitable for nesting.
Most of the reports in the literature of nesting after mid-July rep-
resent records for captive females.

Natural History of the Ornate Box Turtle 555

Nests of T. o. ornata were so well-concealed that they were dif-
ficult to find even when a gravid female had been followed to the
approximate location by means of a trailing thread. Females
spend one to several days seeking a site for the nest, usually trav-
eling a circuitous route within a restricted area. Movements of
nest-seeking females were more extensive than those of males and
non-gravid females observed in the same periods.

Activities of one gravid female, typical in most respects of the
activities of several other gravid females observed (for periods
of one to 23 days) at the Damm Farm, illustrate pre-nesting be-
havior (Fig. 29). A trailer was attached to the female on the
morning of June 7. She was recovered early on the following
afternoon; her movements in the elapsed period had been restricted
to a small, deep, ravine 150 feet long and 20 to 30 feet wide. She
had traversed each edge of the ravine at least once and had
crossed it six or seven times, keeping mostly to areas on the upper
parts of south — or west — facing slopes where vegetation was sparse
or lacking. In six places she had dug into the ground, probably
to test the suitability of the soil for nesting. In three places she
dug beneath rocks that jutted out from the bank, and in two places
merely scratched away the upper crust of soil. Her most recent
attempt at digging (probably late the previous evening or in early
morning on the day of her capture) consisted of a flask-shaped
cavity that, but for the lack of eggs and a covering of earth, was like
a completed nest (PI. 21, Fig. 1). The cavity was 55 millimeters
deep, 80 millimeters wide at the bottom, and 60 millimeters wide
at the opening. For several inches about the opening the earth
was slightly damp. That piled on the rim of the opening was
of the consistency of thick mud, indicating that the female had
voided fluid first on the surface of the earth and again inside the
cavity to soften the soil. Subsequently during eight days her ac-
tivities were similar but not so extensive as on the day described
above. It was determined by daily palpation that she laid her eggs
somewhere in the general area of the ravine on June 15 but the
nest could not be found.

No completed nests containing eggs were discovered at the
Damm Farm but the locations of several robbed nests and partly
completed nests provided some information on preferred sites.
The nests found were on bare, well-drained, sloping areas and
were protected from erosion by upslope clumps of sod or rocks.

556 University of Kansas Publs., Mus. Nat. Hist.

The nest cavity illustrated in Plate 21 was at the edge of the sod-
line on the upper lip of the west-facing bank of a ravine. One
nest had been excavated in a shallow den beneath an overhanging
limestone rock. Three nests were on west- or south-facing slopes
and one was on the north-facing bank of a ravine. Box turtles
presumably select bare areas for nesting because of the greater
ease of digging. One female at the Damm Farm was thought to
have laid her eggs in a cultivated field and William R. Brecheisen
told me he discovered two nests in a wheat field being plowed in
July, 1955.

The repeated excavation of trial nest cavities presumably ex-
hausts the supply of liquid in the female's bladder. Frequent
imbibing of water is probably necessary if the search for a nesting
site is continued for more than a day or two. Standing water was
usually available in ponds, ravines, ditches, and other low areas at
the Damm Farm in June. Nesting in June, therefore, is advanta-
geous not only because of the greater length of time provided for in-
cubation and hatching but also because of the amount of water
available for drinking. Females can probably be more selective in
the choice of a nesting site if their explorations are not limited by
lack of water.

Females of T. ornata, in all instances known to me, began exca-
vation of their nests in early evening and laid their eggs after dark;
Allard (1935:328) reported the same behavior for T. Carolina.

William R. Brecheisen, on July 22, 1955, at his fann, two miles
south and one mile west of Welda, Anderson County, Kansas, ob-
served that a large female began digging a nest in an earth-filled
stock tank at 6:00 P. M. At first she moved her body about on the
surface of the earth, loosening it and pushing it aside with all four
legs, making a depression approximately two inches deep and large
enough to accommodate her body. At 7:30 P. M. she began digging
alternately with her hind feet at the bottom of the depression.
Digging continued until 10:00 P. M., at which time the nest cavity
was three inches deep, and tliree inches in diameter, with a smaller
opening at the top. Six eggs were laid in the next half-hour.
Covering of the nest probably took more than one hour but observa-
tions were terminated after the final egg was laid. By the following
morning the nest-site had been completely covered and was no
difi^erent in appearance from the rest of the earthen floor of the
tank. (Brecheisen observed more of the nesting than anyone else

Natural History of the Ornate Box Turtle 557

has recorded and I am obliged to him for permission to abstract,
as per the above paragraph, the notes that he wrote on the matter. )

A nest made by a captive female at the Reservation was of normal
proportions except for an accessory cavity that opened from the
neck of the nest, immediately below the surface of the ground.
This smaller cavity contained a single egg. This peculiar nest may
have resulted from the eJEorts of two difiFerent females since several
were kept in the same outdoor pen.

Ten adult females were kept in an outdoor cage in the summer of
1955. The cage was raised off the ground on stilts and its floor was
covered with 12 inches of black, loamy soil. A small pan of water
was always available in the cage and the turtles were fed greens,
fruit, and table scraps each evening. Nesting activity was first
noted on June 21, when one of the females was digging a hole in a
corner of the enclosure. She dug with alternate strokes of her fully-
extended hind legs in the manner described (Legler, 1954:141) for
painted turtles ( Chrysemys picta bellii ) . Nevertheless, digging was
much less eflficient than in Chrysemys, because of the narrow hind
foot of the female T. ornata; approximately half of the earth re-
moved by any one stroke rolled back into the nest or was pulled
back when she reinserted her leg. The female stopped digging
when I made sudden movements or held my hand in front of her.
Digging continued for approximately 45 minutes; then the female
moved away and burrowed elsewhere in the cage. The nest cavity
that she left was little more than a shallow depression. Three
other females were digging nests early in the evening on July 3,
5, and 8; in each of these instances the female stopped digging to
eat when food was placed in the cage and completed the nesting
process, unobserved, later in the evening. In each instance where
nest-digging by captive females was observed, the hind quarters
of the female rested in a preliminary, shallow depression, and the
anterior end of the body was tilted upward at an angle of 20 to
30 degrees. In late June and early July several eggs were found,
unburied, on the floor of the cage and in the pan of water.

The excavation of a preliminary cavity by captive females may
not represent a natural phenomenon. Allard (1935) made no
mention of it in his meticulous description of the nesting process
in T. Carolina. It is worthy of mention, however, that Booth
(1958:261) reported the digging of a preliminary cavity by a
captive individual of Gopherus agassizi.

558

Uni\'ersity of Kansas Publs., Mus. Nat. Hist.

-

•

•

-

130

•

•

-

length (mm)

fo

o

~

•

•

•

• •

• •

•

•
•

•
•

•
•

•

"

piastre 1
5

-

•
•

•
1 •

-

2

4

6

8

number

of

eggs

Eggs

The number of eggs in 23 clutches ranged from two to eight
(mean, 4.7 ± 1.37 a); clutches of four, five, and six eggs were
most common, occurring in IS (78 per cent) instances. The ten-
dency for large females to lay
more eggs than small females
( Fig. 6 ) was not so pronounced
as that reported by Cagle (1950:
38) for Fsctidemijs scripta. The
small size of T. ornata, in com-
parison with other emyid turtles,
seemingly limits the number of
eggs that can be accommodated
internally. The number of eggs
per clutch in T. Carolina [2 to
7, average 4.2, Allard (1935:
331 ) ] , is nearly the same as that

of T. ornata. P^^ g ^j^^ relation of plastral length

Shells of the eggs are trans- to number of eggs laid by 21 females of
lucent and pinkish or yellowish ^'- ^- ^''"'^''^ ^"^""^ ^^''^"^ ^^"^^s-
when the eggs are in the oviducts. After several days outside
the oviducts the shells become chalky-white and nearly opaque.
Eggs incubated in the laboratory retained the pinkish color some-
what longer than elsewhere on their under-surfaces, which were
in contact with moist cotton, but eventually even this part of the
shell became white. Infertile eggs remained translucent and
eventually became dark yellow, never becoming white; they could
be distinguished from fertile eggs on the basis of color alone.
Shells of infertile eggs became brittle and slimy after several weeks.
The outer layer of the shell of a freshly laid egg is brittle and
cracks when the egg is dented. After a few days, when the eggs
begin to expand, the shell becomes flexible and has a leathery
texture. The shell is finely granulated but appears smootli to the
unaided eye. The granulations are approximately the same as
those illustrated by Agassiz (1857: PI. 7, Fig. 18) for T. Carolina.
Eggs arc ellipsoidal. Data concerning size and weight (con-
sisting of mean, one standard deviation, and extremes, respectively)
taken from 42 eggs (representing 9 clutches) within 24 hours after
they were laid, or dissected from oviducts, are as follows: length,
36.06 It 2.77 (31.3-40.9); width, 21.72 ± 1.04 (20.0-26.3); and
weight, 10.09 ih 1.31 (8.0-14.3). There was a general tendency

NATxmAL History of the Ornate Box Turtle 559

for smaller clutches to have larger eggs; the largest and heaviest
were in the smallest clutch (two eggs) and the smallest were in
the largest clutch (eight eggs). Risley (1933:697) reported such
a correlation in Sternotherus odoratus, as did Allard (1935:331)
in T. Carolina. Measurements in the literature of the size of eggs
of T, ornata suggest a width greater than that stated above, prob-
ably because some eggs aheady had begun to expand when meas-
ured.

Eggs of T. ornata expand in the course of incubation, as do
other reptilian eggs with flexible shells, owing to absorption of
water. In the laboratory, 48 eggs increased by an average of ap-
proximately three grams in weight and three millimeters in width
over the entire period of incubation; increase in width coincided
with decrease in length. Cotton in incubation dishes was kept
moist enough so that some water could be squeezed from it. When
the cotton was constantly moist, eggs showed a fairly steady ex-
pansion from the first week of incubation until hatching. The
process could be reversed by allowing the cotton to dry. Eggs
that were allowed to dry for a day or more became grossly dented
or collapsed. Eggs at the periphery of the incubation dish were
ordinarily more seriously affected by drying than were those at
the center or in the bottom of the dish. A generous re-wetting
of desiccated eggs and cotton caused the eggs to swell to their
original proportions within 24 hours. Recessions occurred, however,
even in the clutches that received the most nearly even amount of
moisture. Increases in weight and size seemed to reach a peak
in the middle of the incubation period and again immediately be-
fore hatching. Infertile eggs expanded in the same manner as
fertile eggs in the first week or two of incubation, but thereafter
gradually regressed in bulk or failed to re-expand after temporary
periods of dryness. Fertile eggs that were in good condition had
a characteristically turgid, springy feel and could be bounced
off a hard surface.

Temporary lack of moisture usually did not kill embryos; pro-
longed dryness, combined with high temperatures, probably could
not be tolerated. Lynn and Ullrich (1950), by desiccating the
eggs of Chrysemys picta and Chelydra serpentina, produced ab-
normalities in the young ranging from shght irregularities of the
shell to eyeless monstrosities; eggs desiccated in the latter half of
incubation produced a higher percentage of abnormal young than
eggs that were desiccated earlier.

a— 773

560 University of Kansas Publs., Mus. Nat. Hist.

In 1956, three fertile eggs, from clutches that were at different
stages of incubation, were immersed in water for 48 hours. The
eggs rested on the bottom of the bowl in the same position in
which they had been placed in the incubation dishes; when turned,
they returned invariably to the original position. The embryos
in two of the eggs ( one and 27 days old at the time of immersion )
were still living ten days after the eggs were removed from the
water; the embryo in the remaining egg (21 days old at the time
of immersion) was dead. Eggs immersed in water increased in size
and weight at the same rate as eggs in incubation dishes, indi-
cating that absorption of water probably operates on a threshold
principle, the amount absorbed being no more than normal even
under wet conditions.

Natural nests usually are in well-drained areas, but water prob-
ably stands in some nests for short periods after heavy rains. Pro-
vided the nest cavity itself is not damaged, water in the nest is
probably more beneficial than harmful to the eggs; however, nests
that are inundated during floods probably have little chance of
survival.

Embryonic Development

Eggs were examined by transmitted light in the course of in-
cubation. At the time of laying (or removal from oviducts) no
embryonic structures were discernible even in eggs that had been
retained in the oviducts of captive females some weeks past tlie
normal time of laying; a colorless blastodisc could be seen if eggs
were opened. Embryonic structures first became visible at eight
to ten days of incubation; at this time vascularization of the blas-
todisc was evident and the eyes appeared as dark spots. Heart
beats were observed in most embryos by the fifteenth day but
were evident in a few as early as the tenth day. The pulse of a
fifteen-day-old embryo averaged 72 beats per minute at a tem-
perature of 30 degrees. Embryos at fifteen days, measured in a
straight line from cej)halic flexure to posteriormost portion of body,
were approximately nine to ten millimeters long and at 22 days
were 14 millimeters long. At approximately 35 days the eggs
became dark red; embryonic structures were discernible there-
after only in eggs that had embryos situated at one end, close to
the shell.

Incubation periods for 49 eggs (representing 12 clutches) kept
in the laboratory ranged from 56 to 127 days, depending on the
temperature of the air during the incubation period. In 1955, eggs

Natural History of the Ornate Box Turtle

561

were kept at my home in Lawrence where air temperatures were
uncomfortably hot in summer and fluctuations of 20 degrees
(Fahrenheit) or more in a 24-hour period were common. The fol-
lowing summer eggs were kept in my office at the Museum where
temperatures were but slightly cooler than in my home and subject
also to wide variation. In 1957 this part of the Museum was air-
conditioned and kept at approximately 75 degrees. The greater
lengths of incubation periods at lower temperatures are shown in
Table 1. Risley (1933:698) found the incubation period of Sterno-
therus odoratus to be longer at lower temperatures; corresponding
observations were made by Allard (1935:332) and Driver (1946:
173) on the eggs of Terrapenc Carolina. Cagle (1950:40) and
Cunningham (1939) found no distinct differences in length of in-
cubation period for eggs of Fseudemys scripta and Mahclemys
terrapin, respectively, at different temperatures within the range
tolerated by the eggs.

Most nests observed in the field were in open situations where
they would receive the direct rays of the sun for at least part of
the day; the shorter average incubation periods (59 and 70 days,
respectively), observed in 1955 and 1956, therefore, more nearly
reflect the time of incubation under natural conditions than does
the excessively long period (125 days at 75 degrees) observed in
1957 under cooler, more nearly even temperatures.

Sixty-five days seems to be a realistic estimate of a typical incu-

Table 1. — The Relationship of Temperature and DtrRATioN of Incubation
Period as Determined from Laboratory Studies of 49 Eggs of T. ornata.

Average
daily

Period of incubation
(Days)

Number

of
clutches

Number

of

eggs

Remarks

temperature
(Fahrenheit)

Mean

Range

91

59

56-64

6

24

Wide daily

fluctuations in

temperature

82

70

67-73

4

21

Wide daily

fluctuations in

temperature

75

125

124-127

2

4

Temperature

thermostatically

controlled

562 University of Kansas Publs., Mus. Nat. Hist.

bation period under natural conditions; eggs laid in mid-June
would hatch by mid-August. Even in years when summer tempera-
tures are much cooler than normal, eggs probably hatch by the
end of October. Hatchlings or eggs would have a poor chance
of surviWng a winter in nests on exposed cut-banks or in other
unprotected situations. Overwintering in the nest, hatchlings might
survive more often than eggs, since hatchlings could burrow into
the walls and floor of the nest cavity. Unsuitable environmental
conditions that delay the nesting season and retard the rate of
embryonic development may, in some years, be important limiting
factors on populations of ornate box turtles.

In areas where T. ornata and T. Carolina are sympatric (for ex-
ample, in Illinois, Kansas, and Missouri) the two species occupy
different habitats, ornata preferring open grassland and Carolina
wooded situations. Under natural conditions, the average incuba-
tion periods of these two species can be expected to differ, T. Caro-
lina having a somewhat longer period due to lower temperatures
in nests that are shaded. In the light of these speculations, the re-
mark of Cahn (1937:102) — that T. ornata nested later in the season
(in Illinois) and compensated for this by having a shorter incuba-
tion period — is understandable.

The range of temperatures tolerated by developing eggs probably
varies with the stage of embryonic development. When tempera-
tures in the laboratory were 102 to 107 degrees Fahrenheit for ap-
proximately eight hours, due to a defect in a thermostat, the young
in two eggs of T. ornata, that had begun to hatch on the previous
day, were killed, as were the nearly full-term embryos in a num-
ber of eggs of T. Carolina (southern Mississippi) kept in the same
container. A five-day-old hatchhng of T. ornata, kept in the same
container, survived the high temperatures with no apparent ill ef-
fects. Cagle (1950:41) found that eggs of Pseudemys scripta
could not withstand temperatures of 10 degrees for two weeks nor
would they survive if incubated at 40 degrees. Cunningham ( 1939)
reported that eggs of Malaclemys terrapin could not survive pro-
longed exposure to temperatures of 35 to 40.6 degrees but tolerated
temporary exposure to temperatures as high as 46 degrees.

In the summer of 1955, a clutch of three eggs, all of which con-
tained nearly full-term embryos, was placed in a refrigerator for
48 hours. The temperature in the refrigerator was maintained
at approximately 4.5 degrees; maximum and minimum temperatures
for the 48 hour period were 2.8 and 9.5 degrees, respectively. When
the eggs were removed from the refrigerator they showed gains in

Natural History of the Ornate Box Turtle 563

weight and increases in size comparable to eggs, containing em-
bryos of the same age, used as controls. The experimental eggs
began to hatch two days after they were removed to normal tem-
peratures— approximately 24 hours later than the controls.

In the late stages of incubation, the outer layer of the shell be-
comes brittle and is covered with a mosaic of fine cracks or is
raised into small welts. Several days before hatching, movements
of the embryo disturb the surface of the shell and cause the outer
layer to crumble away, especially where the head and forequarters
of the embryo lie against the shell. Some embryos could be seen
spasmodically thrusting the head and neck dorsally against the
shell.

The role of the caruncle in opening the shell seems to vary
among diflPerent species of turtles. Cagle (1950:41) reported that
it was used only occasionally by Pseudemys scripta; Allard (1935:
332) thought that it was not used by Terrapene Carolina; and, the
observations of Booth (1958:262) and Grant (1936:228) indicate
that embryos of Gopherus agassizi use the caruncle at least in the
initial rupturing of the shell.

In the three instances in which hatching was closely observed
in T. ornata, the caruncle made the initial opening in the shell;
claws of the forefeet may have torn shells in other hatchings that
were not so closely observed. In all observed instances, the shell
was first opened at a point opposite the anterior end of the embryo.
The initial opening had the appearance of a three-cornered tear.
A quantity of albuminous fluid oozed from eggs as soon as the shells
were punctured.

The initial tear is enlarged by lateral movements of the front
feet, and later the hind feet reach forward and lengthen the tear
farther posteriorly. In many instances a tear develops on each
side and the egg has the appearance of being cleft longitudinally.
The young turtle emerges from the anterior end of the shell or backs
out of the shell through a lateral tear.

The process of hatching, from rupture of shell to completion of
emergence, extended over three to four days in the laboratory.
Many hatchlings from time to time crawled back into the shell over
a period of several days after hatching was completed. In a clutch
of eggs kept in a pail of earth, by William R. Brecheisen, eight days
elapsed between onset of hatching and appearance of the first
hatchling at the surface.

A nest in an outdoor pen at the Reservation was discovered in

564

UNivEiisiTY OF Kansas Publs., Mus. Nat. Hist,

Table 2. — CoMPARATrvE Rates of
Fertility and Prenatal Mortaltty
FOR Eggs Dissected from Oviducts
AND FOR Eggs That Were Laid in
THE Laboratory and Subsequently
Removed to Incubation Dishes.

early October. The cap had been recently perforated and the
hatchlinGis had escaped. One of them, judged to be approximately
two weeks old, was found in a burrow nearby. The cavity of the
nest appeared to have been enlarged by the young. The eggs were
probably laid in early July. Emergence of young from the nest
had been delayed for a time after hatching, until rain softened the
ground in late September and early October.

Fertility and Prenatal Mortality

Eggs were incubated in the laboratory at more nearly optimum
temperature and humidity than were eggs in natural nests. Per-
centage of prenatal mortality probably was lower in laboratory-
incubated eggs than in those
under natural conditions.

Of sixty eggs studied in the
laboratory, 45 (75 per cent)
were fertile; 36 (80 per cent)
of the fertile eggs (those in
which the blastodisc was at some
time discernible by transmitted
light) hatched successfully. In
six clutches all the eggs were
fertile and five of these clutches
hatched with 100 per cent suc-
cess. One clutch contained eggs
that were all infertile and an-
other clutch had four infertile
eggs and two fertile eggs that
failed to hatch. Among nine
fertile eggs that failed to survive,
four casualties occurred in the
late stages of incubation or after
hatching had begun, indicating
that these are probably critical periods.

Fertility of eggs was not correlated with size or age of female,
with size of clutch, or with size of egg. Eggs laid in the laboratory
had higher rates of infertility and prenatal mortality than did eggs
dissected from oviducts. Handling of eggs in removing them from
nests to incubation dishes, after embryonic development had be-
gun, might have been responsible for reduced viability (Table 2).

Number

or

Percentage

>- o

on

O

-3 3

o >

^ o

Number of eggs examined

22

38

Percentage of fertile eggs

64

82

Percentage of fertile
eggs hatclicd

50

94

Percentage of eggs hatched

32

76

Natural History of the Ornate Box Turtle 565

Reproductive Potential

Assuming that 4.7 eggs are laid per season, that all eggs are
fertile and all hatch, that all young survive to maturity, that half
the hatchlings are females, and that females first lay eggs in the
eleventh year, the progeny of a single mature female would num-
ber 699 after twenty years. Considering that infertility and pre-
natal mortality ehminate approximately 40 per cent of eggs laid
( according to laboratory findings ) the average number of surviving
young per clutch would be 2.8 and the total progeny, after 20
years, would be 270, provided that only one clutch of eggs was
laid per year. But it is thought that, on the average, one third
of the female population produces two clutches of eggs in a single
season. If the second clutch contains 3.5 eggs (resulting in 2.1
surviving young when factors of infertility and prenatal mortality
are considered), the progeny of a single female, after 20 years,
would number approximately 380. Postnatal mortality reduces
the progeny to a still smaller number.

The small number of eggs laid each year and the long period
required to reach sexual maturity make the reproductive potential
of T. ornata smaller than that of the other turtles of the Great
Plains, and much smaller than nearly any of the non-chelonian
reptiles of the same region.

Number of Reproductive Years

The total span of reproductive years is difficult to determine;
I am unable to ascertain the age of a turtle that has stopped grow-
ing. No clearly defined external characteristics of seniHty were
discovered in the populations studied. A male that I examined
had one atrophied testis. In another male both testes were
shrunken and discolored and appeared to be encased by fibrous
tissue. Both males were large, well past the age of regular growth,
and had smoothly worn shells. Several old females had seemingly
inactive ovaries. Reproductive processes probably continue
throughout life in most members of the population, although pos-
sibly at a somewhat reduced rate in later life.

GROWTH AND DEVELOPMENT

Initiation of Growth

Young box turtles became active and alert as soon as they
hatched, and remained so until low temperatures induced quies-
cence. If sand or soil was available, hatchlings soon burrowed into

566 University of Kansas Publs., Mus. Nat. Hist.

it and became inactive. Covering containers with clamp cotton
also induced inactivity; the hatchlings usually made no attempt to
burrow through the confining layer. Desire to feed varied in
hatchlings of the same brood and seemed not to be correlated with
retraction of the yolk sac or retention of the caruncle. Some
hatchlings actively pursued mealworms; on subsequent feedings
they learned to associate my presence with food and eagerly took
mealworms from forceps or from my hand. Meat, vegetables, and
most other motionless but edible objects were ignored by hatchlings
but some individuals learned to eat meat after several forced feed-
ings. Hatchlings that regularly took food in the first month of life
ordinarily grew faster than hatchlings that did not eat. Many of
the hatchlings in the laboratory showed no areas of new epidermal
growth on the shell in the time between hatching and first ( induced )
hibernation.

Size and Appearance at Hatching

The proportions of the shell change somewhat in the first few
weeks of life. At hatching the shell may be misshapen as a result
of confinement in the egg. Early changes in proportions of the
shell result from expansion — widening and, to a lesser degree,
lengthening of the carapace — immediately after hatching. Subse-
quent retraction or rupture of the yolk sac and closure of the navel
are accompanied by a decrease in height of shell and slight, further
widening of the carapace.

The yolk sac retracts mainly between the time when the egg
shell is first punctured and the time when the turtle actually emerges
from the shell. When hatching is completed, the yolk sac usually
protrudes no more tlian two millimeters, but in some individuals
it is large and retracts slowly over a period of several days.

One individual began hatching on November 11 and was com-
pletely out of the egg shell next day; the yolk sac was 15 millimeters
in diameter, protruded six millimeters from the umbilical opening,
and hindered the hatchling's movements. The sac broke two days
later, smearing the bottom of the turtle's dish with semifluid yolk.
The hatchling then became more active. Twenty-six days later the
turtle was still in good condition and its navel was nearly closed.
A turtle that hatched with a large yolk sac in a natural nest possibly
would benefit, through increased ease of mobility, if the yolk sac
ruptured.

A recently hatched turtle was found at the Reservation in Oc-

Natural History of the Ornate Box Turtle 567

tober, 1954, and was kept in a moist terrarium in the laboratory
where it died the following May. The turtle was sluggish and ate
only five or six mealworms while in captivity; no growth was de-
tectable on the laminae of the shell. Autopsy revealed a vestige
of the retracted yolk sac, approximately one millimeter in diameter,
on the small intestine.

The navel ("umbilical scar") of captive hatchlings ordinarily
closed by the end of the second month but in three instances re-
mained open more than 99 days. The position of the navel is
marked by a crescent-shaped crease, on the abdominal lamina,
that persists until the plastron
is worn down in later years
(PI. 24, Fig. 1).

The caruncle ("egg tooth")

(Fig. 7) remains attached to

the horny maxillary beak for a

variable length of time; 93 per

cent of the live hatchlings kept

in the laboratory retained the Fig. 7. A hatchling of T. o. ornata

1 j-t- J. ii- J •fi ( X 2 ) that still retains the caruncle

caruncle on the tenth day, 71 ("egg tooth"). A distinct boss will re-

per cent on the twentieth day, main on the maxillary beak after the

1 1 irv . .1 carvmcle is shed,

and only 10 per cent on the

thirtieth day of hfe. Few individuals retained the caruncle when
they entered hibernation late in November, and none retained it
upon emergence from hibernation. Activities in the first few days
or weeks of life influence the length of time that the caruncle is
retained; turtles that begin feeding soon after hatching probably
lose the caruncle more quickly than do those that remain quiescent.
The caruncles of some laboratory specimens became worn before
finally dropping off. Almost every caruncle present after 50 days
could be flicked off easily with a probe or fingernail. The initiation
of growth of the horny maxillary beak probably causes some loosen-
ing of the caruncle. The caruncle may aid hatchlings in escaping
from the nest.

After the caruncle falls off, a distinct boss remains, marking its
former place on the homy beak ( PI. 25, Fig. 1 ) ; this boss is gradually
obhterated over a period of weeks by wear and by differential
growth, and is seldom visible in turtles that have begun their first
full year of growth. The "first full year of growth" is here consid-
ered to be the period of growth beginning in the spring after
hatching.

568 University of Kansas Publs., Mus. Nat. Hist.

Growth of Epidermal Laminae

Growth of ornate box turtles was studied by measuring recaptured
turtles in the field, by periodically measuring captive hatchlings
and juveniles, and by measuring growth-rings on the epidermal
laminae of preserved specimens. Studies of growth-rings provided
by far the greatest volume of information on growth, not only for
the years in which field work was done, but for the entire life of
each specimen examined.

It was necessary to determine the physical nature of growth-
rings and the manner in which they were formed before growth
could be analyzed. Examination of epidermal laminae on the shell
of a box turtle reveals that each has a series of grooves — growth-
rings — on its surface. The deeper grooves are major growth-rings;
they occur at varying distances from one another and run parallel
to the growing borders of the lamina. Major growth-rings vary
in number from one to 14 or more, depending on the age of the
turtle (PI. 22). In juvenal turtles and in young adults, major
growth-rings are distinct and deep. Other grooves on the shell —
minor growth-rings — have the same relationship to the borders
of the laminae but are shallower and less distinct than major
growth-rings. One to several minor growth-rings usually occur
on each smooth area of epidermis between major growth-rings.
As the shell of an adult turtle becomes worn, the minor growth-
rings disappear and the major rings become less distinct. Both
sets of rings may be completely obliterated in old turtles but the
major rings usually remain visible until several years after puberty.

In cross section, major growth-rings are V- or U-shaped. The
inner wall of each groove is the peripheral edge of the part of the
scute last formed whereas the outer wall represents the inner edge
of the next new area of epidermal growth. The gap produced
on the surface of the lamina ( the open part of the groove ) results
from cessation of growth at the onset of hibcration. Minor growth-
rings are shallow and barely discernible in cross-section (Fig. 8).
It may therefore be understood that growth-rings are compound
in origin; each ring is formed in part at the beginning of hiberna-
tion and in part at the beginning of the following growing season.

The few publications discussing growth in turtles express con-
flicting views as to the exact mode of growth of epidermal laminae.
Carr (1952:22) briefly discussed growth of turtle scutes in gen-
eral and stated that eccentric growth results from an entirely new

Natural History of the Ornate Box Turtle 569

laminal layer forming beneath, and projecting past the edges of the
existing lamina. Ewing (1939) found the scutes of T. Carolina to
be the thickest at the areola and successively thinner in the fol-
lowing eight annual zones of growth; parts of scutes formed sub-
sequent to the ninth year varied irregularly in thickness. He
stated that epidermal growth took place at the margins of the
laminae rather than over their entire undersurfaces.

It is evident that the mode of scutular growth described by
Carr (loc. cit.) applies to emyid turtles that shed the epidermal
laminae more or less regularly (for example, Chrysemys and
Fsettdemys). In these aquatic emyids a layer of the scute, the
older portion, periodically becomes loose and exfoliates usually
in one thin, micalike piece; since the loosened portion of the scute
corresponds in size to the scute below, it must be concluded that
a layer of epidermis is shed from the entire upper surface of the
scute, including the area of new epidermal growth. Box turtles
ordinarily do not shed the older parts of their scutes; the areola
and successively younger portions of the lamina remain attached
to the shell until worn off. The appearance of a single unworn
scute, especially one of the centrals or the posterior laterals, closely
resembles a low, lopsided pyramid.

Examination of parasagittal sections of scutes revealed that they
were composed of layers, the number of layers varying with the
age of the scute. A scute from a hatchling consists of one layer. A
scute that shows a single season of growth has two layers; a new
layer is added in each subsequent season of growth. Stratification
is most evident in the part of the scute that was formed in the first
three or four seasons and becomes increasingly less distinct in newer
parts of the scute. It may further be understood that scutes grow
in the manner described by Carr ( loc. cit.).

When the epidermal laminae are removed, a sheet of tough, pale
grayish tissue remains firmly attached to the bones of the shell be-
neath. This layer probably includes, or consists of, germinal epi-
thelium. Contrasting pale and dark areas of the germinal layer
correspond to the pattern of markings on the scute removed.

Growth of epidermal laminae is presumably stimulated by growth
of the bony shell. As the bone grows, the germinal layer of the
epidermis grows with it. When growth ceases at the beginning of
hibernation, the thin edges of the scutes are slightly down-turned
where they enter the interlaminal seams (Fig. 8). When growth
is resumed in spring, the germinal layer of the epidermis, rather
than continuing to add to the edge of the existing scute, forms an

570

University of Kansas Publs., Mus. Nat. Hist.

'_o

Fig. 8. The second central scute
from a juvenal T. o. ornata (KU
16133) in its third full season of
growth. A ) Entire scute from above
( X 2J2); dashed line shows portion
removed in parasagittal section. B )
Diagonal view of section removed
from scute in "A" ( X 4/8, thickness
greatly exaggerated) showing lay-
ers of epidermis formed in succes-
sive seasons of growth. Each layer
ends at a major growth-ring (M
1-3) that was formed during hiber-
nation; minor growth-rings (m),
formed in the course of the grow-
ing season, do not result from the
formation of a new layer of epi-
dermis. Note the granular texture
of the areola (a); the smooth zone
between the areola and Ml shows
amount of growth in the season of
hatching.

entirely new layer of epidermis. The new layer is thin and indis-
tinct under the oldest part of the scute but becomes more distinct
toward its periphery. Immediately proximal to the edge of the
scute, the new layer becomes greatly thickened, and, where it
passes under the edge, it bulges upward, recurving the free edge of
the scute above. At this time the formation of a major growth-ring
is completed. The newly-formed epidermis, projecting from under
the edges of the scute, is paler and softer than the older parts of the
scute; the presence or absence of areas of newly formed epidermis

Natural, History of the Ornate Box Turtle 571

enables one to determine quickly whether a turtle is growing in the
season in which it is captured. There is little actual increase in
thickness of the scute after the first three or four years of growth.
The epidermal laminae are therefore like low pyramids only in ap-
pearance. This appearance of thickness is enhanced by the con-
tours of bony shell which correspond to the contours of the scutes.

Minor growth-rings difiEer from major growth-rings in appear-
ance and in origin. Ewing ( op. cit.: 91 ) recognized the difference
in appearance and referred to minor growth-rings as "pseudoannual
growth zones." Minor growth-rings result from temporary cessa-
tions of growth that occur in the course of the growing season, not
at the onset of hibernation. They are mere dips or depressions in
the surface of the scute. The occurrence of minor growth-rings
indicates that interruptions in growth of short duration do not re-
sult in the formation of a new layer of epidermis. Slowing of
growth or its temporary cessation may be caused by injuries, peri-
ods of quiescence due to dry, hot, or cold weather, lack of food,
and possibly by physiological stress, especially in females, in the
season of reproduction. Minor growth-rings that lie immediately
proximal to major growth-rings (PI. 22, Fig. 2), are the result of
temporary dormancy in a period of cold weather at the end of a
growing season, followed by nearly normal activity in a warmer
period before winter-long hibernation is begun. Cagle (1946:
699) stated that sliders {Pseudemys scripta elegans) remaining
several weeks in a pond that had become barren of food would stop
growing and develop a growth-ring on the epidermal laminae; he
did not indicate, however, whether these growth-rings differ from
those formed during hibernation.

In species that periodically shed scutes a zone of fracture de-
velops between the old and new layers of the scute as each new
layer of epidermis is formed, and the old layer is shed. Considering
reptiles as a group, skin shedding is of general occurrence; the
process in Pseudemys and Chrysemys differs in no basic respect
from that in most reptiles. Retention of scutes in terrestrial emyids
and in testudinids is one of many specializations for existence on
land. Retention of scutes protects the shell of terrestrial chelonians
against wear. Some box turtles were observed to have several
scutes of the carapace in the process of exfoliation but no exfoha-
tion was observed on the plastron. Exfoliation ordinarily occurred
on the scutes of the carapace that were the least worn; the ex-
foliating portion included the areola and the three or four oldest
(first formed) layers of the scute. The layer of scute exposed

572 Umversity of Kansas Publs., Mus. Nat. Hist.

was smooth and had yellow markings that were only slightly less
distinct than those on the portion that was e.xfoliating.

Wear on the shell of a bo.x turtle reduces the thickness of scutes,
as does the shedding of scutes in the aquatic emyids mentioned.
It is noteworthy that any of the layers in the scute of a box turtle
can form the cornified surface of the scute when the layers above it
wear away or are shed.

It is uncertain whether turtles that have ceased to grow at a
measurable rate continue to elaborate a new layer of epidermis
at the beginning of each season. Greatly worn shells of ornate
box turtles, particularly those of the subspecies liiteola, have only
a thin layer of epidermis through which the bones of the shell and
the sutures betvveen the bones are visible. I suspect that, in these
old individuals, the germinal layer of the epidermis does not be-
come active each year but retains the capacity to elaborate new
epidermis if the shell becomes worn thin enough to expose and
endanger the bone beneath it. The germinal layer of old turtles
loses the capacity to produce color.

Major growth-rings constitute a valuable and accurate history
of growth that can be studied at any time in the life of the turtle
if they have not been obliterated. They are accurate indicators
of age only as long as regular growth continues but may be used
to study early years of growth even in turtles that are no longer
growing. Minor growth-rings, if properly interpreted, provide
additional information on growing conditions in the course of each
growing season.

Nichols (19o9a: 16-17) found that the number of growth-rings
formed in marked individuals of T. Carolina did not correspond to
the number of growing seasons elapsed; he concluded that growth-
rings were unreliable as indicators of age and that box turtles fre-
quently skipped seasons of growth. Woodbury and Hardy (1948:
166-167) and Miller (1955:114) came to approximately the same
conclusion concerning Goplxerus agassizi. It is significant that these
workers were studying turtles of all sizes and ages, some of which
were past the age of regular, annual growth. Cagle's review of
the literature concerning growth-rings in turtles (1946) suggests
that, in most of the species studied, growth-rings are formed reg-
ularly in individuals that have not attained sexual maturity but are
formed irregularly after puberty.

Cagle's (op. cit.) careful studies of free-living populations of
Psciidcmijs scripta showed that growth-rings, once formed, did not
change in size, that the area between any two major growth-rings

Natural History of the Ornate Box Turtle

573

represented one season of growth, and that growth-rings were re-
liable indicators of age as long as the impression of the areola
remained on the scutes studied. Cagle noted decreasing distinct-
ness of growth-rings after each molt.

The relative lengths of the abdominal lamina and the plastron
remain approximately the same throughout life in T. ornata.
Measurements were made of the plastron, carapace, and abdominal
lamina in 103 specimens of T. o. ornata from Kansas and neighbor-
ing states. The series of specimens was divided into five nearly
equal groups according to length of cai-apace. Table 3 summarizes
the relationship of abdominal length to plastral length, and of
carapace length to plastral length. The matliematical mean of the
ratio, abdominal length/plastral length, in each of the four groups
of larger-sized turtles, was not significantly different from the same
ratio in the hatchling group. The relative lengths of carapace and

Table 3. — The Relationship of Length of Abdominal Scute to Plastral
Length, and of Plastral Length to Length of Carapace, in 103 Speci-
mens OF T. o. ornata Arranged in Five Groups According to Length of
Carapace. The Relative Lengths of Abdominal Scute and Plastron
are not Significantly Different in the Five Groups. The Plastron
Tends to be Longer than the Carapace in Specimens of Adult or

Nearly Adult Size.

Length
OF Carapace

Number

of
specimens

Length of abdominal
as a percentage of
length of plastron

Individuals having
plastron longer
than carapace

Mean =fc am

Extremes

Number

Percentage

Less than
50 mm.
(Juveniles)

23

18. 3 ±.498

13.7-20.3

7

38.5

50 to 69 mm.

(Juveniles)

20

17.8±.303

15.2-20.2

8

40.0

70 to 100 mm.
(Subadults)

20

17.9±.445

14.3-20.6

15

75.0

More than
100 mm.
(Adult males)

20

17.8±.236

10.4-20.6

13

65.0

More than

100 mm.
(Adult females)

20

18.8±.510

15.1-25.7

19

1

95.0!

574 University of Kansas Publs., Mus. Nat. Hist.

plastron are not so constant; the carapace is usually longer than the
plastron in hatchlings and juveniles, but shorter than the plastron
in adults, especially adult females.

The length of any growth-ring on the abdominal lamina can be
used to determine the approximate length of the plastron at the
time the growth-ring was formed. Actual and relative increases
in length of the plastron can be determined in a hke manner.
For example, a seven-year-old juvenile (KU 3283) with a plastron
74.0 millimeters long had abdominal growth-rings (beginning with
areola and ending with the actual length of the abdominal) 5.9,
7.8, 9.5, 10.7, 12.0, 12.5, 14.3, and 14.9 millimeters long. Using the

proportion.

3 ABn

rx J'

rAB ABi .

where AB is the abdominal length, PL the
PL "' ■

plastral length, AB^ the length of any given growth-ring, and X
the plastral length at the time growth-ring AB^ was formed, the
plastral length of this individual was 29.3 millimeters at hatching,
38.8 at the end of the first full season of growth, and 47.2, 53.2,
59.6, 62.1, and 71.0 millimeters at the end of the first, second, third,
fourth, fifth, and sixth seasons of growth, respectively. The present
length of the abdominal ( 14.9 mm. ) indicates an increment of three
milHmeters in plastral length in the seventh season, up to the time
the turtle was killed (June 25). This method of studying growth
in turtles was first used by Sergeev (1937) and later more ex-
tensively used by Cagle (1946 and 1948) in his researches on
Pseiidemijs scripta. Because the plastron is curved, no straight-line
measurement of it or its parts can express true length. Cagle ( 1946
and 1948) minimized error by expressing plastral length as the
sum of the laminal (or growth-ring) lengths. This method was
not possible in the present study because growth-rings on parts of
one or more laminae (chiefly the gulars and anals) were usually
obliterated by wear, even in young specimens. It was necessary
to express plastral length as the sum of the lengths of forelobe and
hind lobe.

The abdominal lamina was selected for study because of its
length (second longest lamina of plastron), greater symmetry, and
flattened form. Although the abdominal is probably subject to
greater, over-all wear than any other lamina of the shell, wear is
even, not localized as it is on the gulars and anals.

In instances where some of the growth-rings on an abdominal
lamina were worn but other rings remained distinct, reference to

Natural History of the Ornate Box Turtle 575

other, less worn lamina permitted a correct interpretation of in-
distinct rings.

Abdominal laminae were measured at the interlaminal seam;
since the laminae frequently did not meet perfectly along the
midline (and were of unequal length), the right abdominal was
measured in all specimens. Growth-rings on the abdominal laminae
were measured in the manner shown in Plate 22.

Data were obtained for an aggregate of 1272 seasons of growth
in 154 specimens (67 females, 48 males, and 39 of undetermined
sex, chiefly juveniles). Averages of calculated plastral length were
computed in each year of growth for specimens of known sex
(Figs. 9 and 10) and again for all specimens examined. Annual
increment in plastral length was expressed as a percentage of plas-
tral length at the end of the previous growing season (Fig. 11).
Increment in plastral length for the first season of growth was
expressed as a percentage of original plastral length because of
variabihty of growth in the season of hatching; growth increments
in the season following hatching are, therefore, not so great as
indicated in Figure 11.

Growth of Juveniles

Areas of new laminal growth were discernible on laboratory
hatchhngs soon after they ate regularly. Hatchlings that refused
to eat or that were experimentally starved did not grow. The first
zone of epidermis was separated from the areola by an indistinct
growth-ring (resembling a minor growth-ring) in most hatchlings,
but in a few specimens the new epidermis appeared to be a con-
tinuation of the areola. Major growth-rings never formed before
the onset of the first hibernation.

Growth in the season of hatching seems to depend on early
hatching and early emergence from the nest. Under favorable
conditions hatchlings would be able to feed and grow eight weeks
or more before hibernation. Hatchlings that emerge in late autumn
or that remain in the nest until spring are probably unable to find
enough food to sustain growth.

Sixty-four (42 per cent) of the 154 specimens examined showed
measurable growth in the season of hatching. The amount of in-
crement was determined in 36 specimens having a first growth-ring
and an areola that could be measured accurately. The average
increment of plastral length was 17.5 per cent (extremes, 1.8-66.0
per cent) of the original plastral length. Ten individuals showed

4—773

576 University of Kansas Publs., Mus. Nat. Hist.

E
E

c
o

110

100

90

80

70

o

o.

^ 60

o

C7>

I 50

40

30

20

67

67

66

67

65

46

47

48

48

46

63

38

K^

W

If

w

If

w

W

II |^

II IF

J ' I I I

i 2 3 4 5 6

age

Fig. 9. See legend for Fig. 10.

Natural History of the Ornate Box Turtle

577

130

120

I 110

c
o

■S 100

c

90

80

70

1

8

9 10

age

II

12

Fig. 10. The relationship of size to age in T. o. ornata, based on studies of
growth-rings in 115 specimens of known sex (67 females and 48 males)
from eastern Kansas. Size is expressed as plastral length at the end of each
growing season (excluding the year of hatching) through the twelfth and
thirteenth years (for males and females, respectively) of life. Vertical and
horizontal hnes represent, respectively, the range and mean. Open and
solid rectangles represent one standard deviation and two standard errors
of the mean, respectively. Age is expressed in years.

an increment of more than 20 per cent; the majority of these indi-
viduals ( 8 ) were hatched in the years 1947-50, inclusive.

Some hatchlings that grow rapidly before the first winter are as
large as one- or two-year-old turtles, or even larger, by the follow-
ing summer. Individuals that grew rapidly in the season of hatch-
ing tended also to grow more rapidly than usual in subsequent
seasons; 80 per cent of the individuals that increased in plastral
length by 20 per cent or more in the season of hatching, grew faster

578 University of Kansas Publs., Mus. Nat. Hist.

than average in the two seasons following hatching. Early hatch-
ing and precocious development presumably confer an advantage
on the individual, since turtles that grow rapidly are able better to
compete with smaller individuals of the same age. Theoretically,
turtles growing more rapidly than usual in the first two or three
years of life, even if they grew subsequently at an average rate,
would attain adult size and sexual maturity one or more years be-
fore other turtles of the same age. A few turtles (chiefly males)
attain adult size ( and presumably become sexually mature ) by the
end of the fifth full season of growth (Figs. 9 and 10). These indi-
viduals, reaching adult size some three to four years sooner than
the average age, were precocious also in the earlier stages of post-
natal development.

Young box turtles reared in the laboratory grew more slowly
than turtles of comparable ages under natural conditions; this was
especially evident in hatchlings and one-year-old specimens.
Slower growth of captives was caused probably by the unnatural
environment of the laboratory. Captive juveniles showed a steady
increase in weight ( average, .52 grams per ten days ) as they grew
whereas captive hatchlings tended to lose weight whether they
grew or not.

Growth in Later Life

After the first year growth is variable and size is of little value
as an indicator of age. Although in the turtles sampled variation
in size was great in those of the same age, average size was suc-
cessively greater in each year up to the twelfth and thirteenth years
(for males and females, respectively), after which the samples
were too small to consider mathematically.

Increments in plastral length averaged 68.1 per cent in the year
after hatching, 28.6 per cent in the second year and 18.1 per cent
in the third year. From the fourth to the fourteenth year the growth-
rate slowed gradually from 13.3 to about three per cent (Fig. 11).
These averages are based on all the specimens examined (with
no distinction as to sex); they give a general, over-all picture
of growth rate but do not reflect the changes that occur in growth
rate at puberty (as shown in Figs. 9 and 10).

Rate of growth and, ultimately, size are influenced by the at-
tainment of sexual maturity. Adult females grow larger than adult
males. Males, nevertheless, grow faster than females and become
sexually mature when smaller and younger. Examination of go-
nads showed 17 per cent of the males to be mature at plastial
lengths of 90 to 99 millimeters, 76 per cent at 100 to 109 milli-

Natural History of the Ornate Box Turtle

579

meters, and 100 per cent at 110 millimeters, whereas the corres-
ponding percentages of mature females in tlie same size groups
were: zero per cent, 47 per cent, and 66 per cent. Of the females,
97 per cent were mature at 120 to 129 millimeters and all were
mature at 130 millimeters (Fig, 13). Because growth slows per-
ceptibly at sexual maturity, it is possible, by examination of growth-
rings, to estimate the age of puberty in mature specimens.

Attainment of sexual maturity, in the population studied, was

— 1 — 1 — 1 — 1 — 1 — 1 — 1 — 1 1

-1 — 1 — r— 1 — 1 — r-

70

(100)

-

sz _

^60

/ \

-

c

a>

~ 50

/ \

-

o

>-.

w 40

- / \

»

o

Q.

/ \(I32)

.£ 30

-

a>
S 20

/ \i'^^*

o

' • •\(123)

1 10

(36) T» •--il^!^ '^'^^

(119)* • •<.

_- <47) (19) .,, "

o

(92)

^ 0

1 1 1 1 1 1 1 1 1

<56) (3.) (•p* .
1 1 1 1 1 1

HI 2345678

9 10 II 12 13 14

age in years

Fig. 11. Average increment in plastral length (expressed as a per-
centage of plastral length at the end of the previous season of growth)
in the season of hatching (H) and in each of the following 14 years
of Hfe, based on 1073 growth-rings. The number of specimens
examined for each year of growth is shown in parentheses. Records
for males and females are combined.

more closely correlated with size than with age. For example,
nearly all males were mature when the plastron was 100 to 110
millimeters long, regardless of the age at which this size was at-
tained. The smallest mature male had a plastral length of 99 milH-
meters; according to the data presented in Figures 9 and 10, there-
fore, a few males reach sexual maturity in the fourth year, and
increasingly larger portions of the population become mature in
the fifth, sixth, and seventh years. The majority become mature
in the eighth and ninth years. Likewise, females ( smallest mature
specimen, 107 mm.) may be sexually mature at the end of the
sixth year but most of them mature in the tenth and eleventh
years.

580 University of Kansas Publs., Mus. Nat. Hist.

Annual Period of Growth

In growing individuals, narrow zones of new epidermis form
on the laminae in spring. Nearly all the growing individuals col-
lected in May of 1954 and 1955 had zones of new epidermis on
the shell but those collected in April did not. Activity in the first
week or two after spring emergence is sporadic and regular feed-
ing may not begin until early May. Once begun, growth is more
or less continuous as long as environmental conditions permit for-
aging. The formation of minor growth-rings and adjacent growth-
zones in autumn, provddes evidence that growth commonly con-
tinues up to the time of hibernation. The number of growing days
per year varies, of course, with the favorableness of environmental
conditions. The length of time (162 days) given by Fitch (1956b:
438 ) as the average annual period of activity for T. ornata is a good
estimate of the number of growing days per season.

Environmental Factors Influencing Growth

Zones of epidermis formed in some years are wider or narrower
than the zones bordering them (PI. 22). Zones notably narrower
or wider than the average, formed in certain years, constituted dis-
tinct landmarks in the growth-histories of nearly all specimens; for
example, turtles of all ages grew faster than average in 1954 and
zones of epidermis formed in this year were always wider than those
formed in 1953 and 1955.

An index to the relative success of growth in each calendar year
was derived. Records of growth for all specimens in each age
group were averaged; the figure obtained was used to represent
"normal" or average growth rate in each year of life (Fig. 12).
The over-all averages for the various age groups were then com-
pared with records of growth attained by individuals of correspond-
ing age in each calendar year, growth in a particular year being
expressed as a percentage of the over-all average. The percent-
ages of average growth for all ages in each calendar year were then
averaged; the mean expresssed the departure from normal rate
of growth for all turtles growing in a particular calendar year. For
example, the over-all average increment in plastral length in the
fifth year of life was 12.1 per cent, the increment in the sixth year
was 10 per cent, and so on (Fig. 11). In 1953, turtles in their
fifth and sixth years increased in plastral length by 11.4 and 9.1
per cent, or grew at 94.2 and 91.0 per cent of the normal rate, re-
spectively. The percentages of normal growth rate for these age
groups averaged with percentages of the other age groups in 1953

Natural History of the Ornate Box Turtle 581

revealed that turtles grew at approximately 86 per cent of the nor-
mal rate in 1953.

Growth rates were computed for the twelve-year period, 1943-
1954, because of the concentration of records in these years. Scat-
tered records also were available for many of the years from 1901-
1942. Records for individuals in the season of hatching and the
first full season of growth were not considered.

Direct correlation exists between growth rate and average
monthly precipitation in the season of growth (April to September)
(Fig. 12). In nine of eleven years, the curve for growth rate fol-
lowed the trend of the curve for precipitation; but because other
chmatic conditions also influenced growth, the fluctuations in the
two curves were not proportional to one another.

Grasshoppers form an important element in the diet of box tur-
tles. Smith (1954) traced the relative abundance of grasshoppers
over a period of 100 years in Kansas, and this information is of
significance for comparison with data concerning growth of box
turtles. In general, the growth index was higher when favorable
weather and large populations of grasshoppers occurred in the
same year.

In the following summary, the numbers (1 to 5) used to express
the relative abundance of grasshoppers are from Smith (op. cit.).
Maxima and minima refer to the twelve-year period, 1943-1954.
The growth index for each year (shown as a graph in Fig. 12) ap-
pears in brackets and indicates the percentage of normal growth
attained by all turtles in that year.

Years Favorable for Growth

1954 [126.3]: Growth was better than average for turtles of all
ages. Grasshopper populations were highest (4-f-) since 1948.
Continuously warm weather, beginning in the last few days of
March, permitted emergence in the first week of April; thereafter
conditions were more or less continuously favorable for activity
until late October. Although there was less than an inch of
precipitation in September, precipitation in August and October
was approximately twice normal and more or less evenly dis-
tributed. Warm weather in early November permitted an addi-
tional tsvo weeks of activity.

1945 [125.5]: This was the second most favorable year for growth
and the second wettest year. Records of growth are all from young
turtles (one to four years old), all of which grew more than aver-
age. Daily maximum temperatures higher than 60 degrees Fahren-

582 University of Kansas Publs., Mus. Nat. Hist.

heit on 18 of the last 19 days of March, combined with twice the
normal amount of precipitation in the same period, stimulated
early emergence. August and October were both dry (each with
less than one inch of precipitation) but diurnal temperatures re-
mained warm through the first week in November and probably
prolonged activity of box turtles at least until then. Grasshoppers
were more abundant (3.7) than normal.

Years Unfavorable for Growth

1944 [83.1]: Tliis was the poorest growing year for the period
considered. The lack of a continuously warm, wet period in early
spring probably delayed emergence until the last week in April.
Temperatures remained warm enough for activity until early No-
vember, but dry weather in September and October probably cur-
tailed activity for inducing long periods of quiescence; most of the
precipitation that occurred in the latter two months fell in a one-
week period beginning in the last few days of September. Grass-
hopper populations were higher (4.0) than normal.

1953 [85.6]: This was the second poorest growing year and the
driest year in the period considered. Intermittently cold weather
in spring delayed emergence until the last week in April when
nearly an inch of rail fell. Temperatures were higher than normal
from June to October. The period from September to the end of
October was dry and the small amount of precipitation that oc-
curred was concentrated chiefly at the beginning and end of that
period. Temperatures in late October and early November were
lower than normal. Grasshopper populations were low (2.2).

1952 [88.3]: Environmental conditions were poor for growth and
much like the conditions described for 1953. In both years growth
was much less than normal in turtles of all ages except for one
group (adults that were 10 and 11 years old in 1952 and 1953,
respectively) that was slightly below normal in 1952 and slightly
above normal in 1953.

The small number of records for 1955 were not considered in
Figure 12. Warm weather in the last half of March lengthened the
growing season, and environmental conditions, as in 1954, were
more or less favorable throughout the rest of the summer; 1955
probably ranks with 1954 as an exceptionally good year for growth
of box turtles.

Although the number of records available for turtles hatched
in the period from 1950 to 1954 is small, a few records are available
for all these years except 1951. In general, small samples of turtles

Natural History of the Ornate Box Turtle

583

hatched in these years reflect only the difficulty of collecting
hatchlings and juveniles. In 1951, conditions for incubation and
hatching were poor and the lack of records for that year actually
represents a high rate of prenatal and postnatal mortality. Rainfall
in the nesting season was two to three times normal and tempera-
tures were below normal. Flooding occurred in low areas of Doug-
las County and many eggs may have been destroyed when nests
were inundated. Cold weather probably increased the time of
incubation for surviving eggs so that only a few turtles could hatch
before winter. Flooding and cold, wet weather in the season of
growth and reproduction, affecting primarily eggs and hatchlings,
may act as checks on populations of T. ornata in certain years.

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Fig. 12. The relation of growth rate in Terrapene o. ornata (solid line)
to precipitation ( dotted line ) in eastern Kansas. "Normal" rate of growth
was determined by averaging records of increase in length of plastron for
turtles in each age group. The growth index is expressed as a percentage
of normal growth and is the mean departure from normal of all age groups
in each calendar year. Precipitation is for the period, April to September
(inclusive), at Lawrence, Douglas Co., Kansas. The means for precipita-
tion (4.3) and growth index (100) are indicated by horizontal lines at

the right of the graph.

The environmental factors governing activity of terrestrial turtles
seem to differ at least in respect to threshold, from the factors in-
fluencing the activity of aquatic turtles. A single month that was
drier or cooler than normal probably would not noticeably affect

584 Univei^sity of Kansas Publs., Mus. Nat. Hist.

growth and activity of aquatic emyids in northeast Kansas, but
might greatly curtail growth of box turtles.

Cagle (1948:202) found that growth of shder turtles (Pseudemys
scripta) in Illinois paralleled the growth of bass and bluegills in
the same lake; in the two years in which the fish grew rapidly, the
turtles did also, owing, he thought to "lessened total population
pressure" and "reduced competition for food." Growth of five-
lined skinks (Eumeces fasciatus) on the Natural History Reserva-
tion paralleled growth of box turtles, probably because at least
some of the same environmental factors influence the growth of
both species. Calculations of departure from normal grovvi:h in E.
fasciatus, made by me from Fitch's graph (1954:84, Fig. 13), show
that relative success of growth in the period he considered can be
ranked by year, in descending order, as: 1951, 1949, 1948, 1950,
1952. This corresponds closely to the sequence, 1951, 1948, 1949,
1950, 1952, for T. ornata.

Number of Growing Years

Grovvi:h almost stops after the thirteenth year in females and
after the eleventh or twelfth year in males, approximately three
years, on the average, after sexual maturity is attained. The oldest
individuals in which plastral length had increased measurably in
the season of capture were females 14 (2 specimens) and 15 (1)
years old. The age of the oldest growing male was 13 years.

The germinal layer of the epidermis probably remains semi-
active throughout life but functions chiefly as a repair mechanism
in adults that are no longer growing. Growiih-rings continue to
form irregularly in some older adults. Growth-rings formed after
the period of regular growth are so closely approximated that they
are unmeasurable and frequently indistinguishable to the unaided
eye. If the continued formation of growth-rings is not accompa-
nied by wear at the edges of the laminae, the laminae meeting at
an interlaminal seam descend, like steps, into the seam (PI. 22,
Fig. 2). Interlaminal seams of the plastron deepen with advancing
age in most individuals.

Some individuals that are well past the age of regular growth
show measurable increments in years when conditions are especially
favorable. The three oldest growing females were collected in 1954
— an exceptionally good year for growth. Allowing some latitude
for irregular periods of growth in favorable years subsequent to
the period of regular, more or less steady growth, 15 to 20 years is
a tenable estimate of the total growing period.

Natural History of the Ornate Box Turtle

585

Longevity

Practically nothing is known about longevity in T. omata or in
other species of Terrapene although the several plausible records
of ages of 80 to more than 100 years for T. Carolina (Oliver, 1955:
295-6) would indicate that box turtles, as a group, are long-lived.
There is no known way to determine accurately the age of an adult
turtle after it has stopped growing. It was possible occasionally
to determine ages of 20 to 30 years with fair accuracy by counting
all growth-rings (including those crowded into the interabdominal
seam) of specimens having unworn shells. Without the presence
of newly formed epidermis as a landmark, however, it was never
certain how many years had passed since the last ring was formed.

90-99 100-109 110-119 120-129
length of plastron (mm)

130-139

Fig. 13. The relationship of sexual maturity to size in 164 specimens
(94 females and 70 males) of Terrapene o. omata, expressed as the
percentage of mature individuals in each of five groups arranged ac-
cording to plastral length. Sexual maturity was determined by examina-
tion of gonads. Solid bars are for males and open bars for females.
The bar for males in the largest group is based on assumption since no
males in the sample were so long as 130 mm. Males mature at a smal-
ler size and lesser age (see also Figs. 9 and 10) than females. Plastral
lengths of the smallest sexually mature male and female in the sample
were, respectively, 99 and 107 mm.

Mattox ( 1936 ) studied annual rings in the long bones of painted
turtles (Chrysemys picta) and found fewer rings in younger than
in older individuals but, beyond this, reached no important con-

5S6 University of Kansas Publs,, Mus. Nat. Hist.

elusion. In the present study, thin seetions were ground from the
humeri and femurs of box turtles of various ages and sizes; the
results of this investigation were negative. Distinct rings were
present in the compact bony tissue but it appeared that, after the
first year or two, the rings were destroyed by encroachment of the
marrow cavity at about the same rate at which they were formed
peripherally.

The only methods that I know of to determine successfully the lon-
gevity of long-lived reptiles would be to keep individuals under
observation for long periods of time or to study populations of
marked individuals. Both methods have the obvious disadvantage
of requiring somewhat more than a human lifetime to carry them
to completion. Restudy, after one or more decades, of the popu-
lations of turtles marked by Fitch and myself may provide valu-
able data on the average and maximum age reached by T. ornata.

Ornate box turtles probably live at least twice as long as the
total period of growing years. An estimated longevity of 50 years
would seem to agree with present scant information on age. Con-
sidering environmental hazards, it would be unusual for an individ-
ual to survive as long as 100 years in the wild.

Weight

Weights of ornate box turtles varied so much that no attempt
was made to correlate weight with size. Absolute weights have
little significance since weight is affected to a large extent by the
amount of fluid in the body. Turtles that had recently imbibed
were naturally heavier than those that had not; turtles brought to
the laboratory and kept there for several days lost weight by evap-
oration and by voiding water. Weights of 22 adult females (53
records) and 10 adult males (22 records) averaged 391 and 353
grams respectively, in the period from September, 1954, to October,
1956. Females characteristically gained weight in spring and
early summer and were lighter after nesting. Turtles of both sexes
gained weight in September and October.

Bony Shell

Fontanelles

At the time of hatching, fontanelles remain where bones of the

shell have not yet articulated with their neighbors. In general, the

fontanelles of the shell are closed by the time sexual maturity is

attained, but some remain open a year or two longer.

The fontanelles of the shell are classified as follows (see Figs.
14 to 16 and 18 to 19):

Natural History of the Ornate Box Turtle

587

Plastron

1.) Anteromedian. Rhomboidal; limited anteriorly by hyoplastral
bones and posteriorly by hypoplastral bones; posterior tip of ento-
plastral bone may project into this fontanelle.
2. ) Posteromedian. Limited anteriorly by hypoplastral bones and
posteriorly by xiphyplastral bones (since hypoplastral bones do
not articulate medially in hatchlings, anteromedian and postero-
median fontanelles form a single, more or less dumbbell-shaped
opening ) .

)ntanelles

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40 60 80 100

length of plostron

Fig. 14. Extent of closure of the costoperipheral fonta-
nelles in relation to length of plastron in 17 skeletons of
T. o. ornata from eastern Kansas. Extent of closure is ex-
pressed as an estimated percentage of total closure of all
the costoperipheral fontanelles, even though some of them
close sooner than others. Closure is usually complete by
the time sexual maturity is attained.

Carapace

1.) Costoperipheral. Openings between the free ends of develop-
ing ribs, between nuchal bone and first rib, and, between pygal
bone and last rib; limited laterally by peripheral bones; variable
in shape.

2.) Costoneural. Triangular openings on either side of middorsal
line between proximal ends of costal plates and developing neural
plates.

The costoneural fontanelles are nearly closed in individuals

588 Untv'ersity of Kansas Publs., Mus. Nat. Hist.

of the 70 millimeter (plastron length) class and seldom remain
open after a length of 80 milhmeters is attained (Fig. 14). Of
the costoperipheral fontanelles, the anterior one (between first
rib and nuchal bone) closes first and the posterior one (between
last rib and pygal bone) last. It remains open in some turtles in
which the plastron is longer than 100 millimeters. The remaining
costoperipheral fontanelles close in varying sequence but those
in the area of the bridge (nos. 2 to 5), where there is presumably
greater stress on the shell, close sooner than the others.

The plastral fontanelles are closed in most specimens of the
90 millimeter (plastron length) class; the anteromedian fontanelle
closes first.

The meager covering of the fontanelles makes juvenal turtles
more susceptible than adults to many kinds of injuries and to
predation.

Movable Parts of the Shell

Parts of the shell that are more or less movable upon one another
and that function in closing the shell are found in several famihes
of Recent turtles. African side-necked terrapins of the genus
Pelusios have a movable forelobe on the plastron. Kinostemids
have one or two flexible transverse hinges on the plastron. In the
Testudinidae the African Kiiiixys has a movable hinge on the
posterior part of the carapace and Pyxis arachnoides of Madagascar
has a short, hinged, anterior plastral lobe. Certain trionychid
turtles, such as Lissemys, utilize the flexible flaps of the carapace
(the flaps of some species are reinforced with peripheral bones)
to close the shell.

Movable shell-parts of turtles are, in general, protective in func-
tion; they cover parts of the soft anatomy that would otherwise
be exposed.

A hinged plastron, capable of wholly or partly closing the shell,
occurs in six genera of the family Emyidae (see introduction).
In these emyids the plastron is divided into two lobes, which are
joined to each other by ligamentous tissue at the junction of the
hyoplastral and hypoplastral bones; externally, the hinge occurs
along the seam between the pectoral and abdominal laminae. This
junction forms a more or less freely movable hinge in adults. The
plastron is attached to the carapace by ligamentous tissue. Both
lobes of the plastron or only the buttresses of the hind lobe may
articulate with the carapace. The former condition obtains in
Ei7iys and Emydoidea; the latter more specialized condition is
found in Terrapene.

Naturai. History of the Ornate Box Turtle 589

In generalized emyid turtles such as Clemmys there are no mov-
able shell parts. The plastron is joined to the carapace by the su-
tures of the bridge. A long stout process, the axillary buttress,
arises on each side from the hyoplastron and articulates with the
tip of the first costal. A similar process, the inguinal buttress, arises
from the anterior part of each of the hypoplastral elements and
meets the sixth costal on each side. The buttresses form the an-
terior and posterior margins of the bridge. It is clear that move-
ment of the plastron in many emyids is mechanically impossible
because of the bracing efiFect of the buttresses.

In Terrapene the movable articulations of the shell are neither
structurally nor functionally developed in juveniles. Adults of T.
omata have highly modified bony buttresses on the plastron that
are homologous with those in more generalized emyids. The in-
guinal buttresses are low and wide, and have a sheer lateral sur-
face forming a sliding articulation with the fifth and sixth peri-
pheral bones of the carapace. The axillary buttresses are reduced
to mere bony points near the posterolateral comers of the forelobe
and do not articulate directly with the carapace (Figs. 15 and 16).

The fifth peripheral bone, constituting the lowest point of the
carapace, has a medial projection that acts as a pivoting point for
both lobes of the plastron; the roughened anterior corners of the
hind lobe articulate with these processes. The roughened posterior
corners of the forelobe of the plastron likewise articulate with these
processes. The posterior process or "tail" of the entoplastron ex-
tends to, or nearly to, the bony transverse hinge.

In juveniles that have been cleared and stained, the homologues
of the parts that are movable in adults are easily identifiable; the
proportions of these parts and their relations to one another are,
however, much different.

In juveniles (Figs. 18 and 19) the buttresses are relatively longer
and narrower, and are distinct — more nearly like those of general-
ized emyids than those of adult T. ornata. The buttresses enclose
a large open space, which in adults is filled by the fifth peripheral.
The hyoplastral and hypoplastral bones are in contact only later-
ally. They are firmly joined by bony processes; the interdigitating
nature of this articulation contrasts with its homologue in the adult,
the point where the roughened corners of the forelobes and hind
lobes meet. The fifth peripheral in juveniles (Fig. 19) lies dorsal
to this articulation. The position of the future transverse hinge
corresponds to a line passing through the articulations of the hyo-

590

University of Kansas Publs., Mus. Nat. Hist.

Fig. 15. Lateral view of adult shell
(XS), showing movable parts with
anterior portion at left. (Abbrevia-
tions are as follows: ab, axillary butt-
ress; hp, hypoplastron; hy, hyoplas-
tron; ib, inguinal buttress; p5, fifth
peripheral bone; th, transverse hinge).

Fig. 16. Medial view of adult shell
(X^), showing movable parts with
anterior portion at left. (Abbrevia-
tions as in fig. 15).

Fig. 17. Lateral view of adult shell
(X^), showing scutellation of mov-
able parts with anterior portion at
left. (Abbreviations are as follows:
, spical scale; ax, axillary scale; m5,
th marginal scale; pi, pectoral
lamina. )

plastra and hypoplastra. The tail of the entoplastron ordinarily
extends posterior to this hne in juveniles.

The external scutellation of the plastral hinge in adults also dif-
fers from that in juveniles. In adults ( Fig. 17 and PI. 8 ) the trans-
verse hinge is marked by ligamentous tissue between the pectoral
and abdominal laminae; the forelobe of the plastron is distinctly
narrower than the hind lobe. Two small scales lie near the corner
of the hinge on each side. The larger and more anterior of these
scales is the axillary; it is present in box turtles of all ages. The
smaller scale (Fig. 17), to my knowledge, has never been named
or mentioned in the literature; it is herein termed the apical scale.
It is a constant feature in adults but is always lacking in hatchlings
and small juveniles. Other scales, much smaller than the axillary

Natxjral History of the Ornate Box Turtle

591

and apical, occur on the ligamentous tissue of the hinge of some
adults.

In juveniles (Fig. 20) the pectoroabdominal seam contains no
ligamentous tissue and is like the other interlaminal seams of the
plastron. A lateral apex of the pectoral lamina projects upward
behind the axillary scale on each side, in the position occupied by

Fig. 18. Plastron of hatchling (x2), cleared and stained to show bony
structure. (Abbreviations not listed in legend for Fig. 15 are as follows: at,

anteromedian fontanelle; ep, epiplastron; pf, posteromedian fontanelle. )

Fig. 19. Carapace of hatchling (x 1^), cleared and stained to show bony
structure; lateral view; anterior end at left. (Abbreviations as in Fig. 15.)

Fig. 20. Lateral view of hatchling ( X 1 ) ; note the lateral process of the
pectoral lamina (pi) extending posterior to the axillary scale (ax) in a posi-
tion corresponding to the apical scale of adults. There is no external indi-
cation of the transverse hinge in young individuals. The yolk sac of this
individual has been retracted but the umbilicus (umb) has not yet closed.

the apical scale of adults. Examination of a large series of speci-
mens revealed that the apical scale of adults becomes separated
from the lateral apex of the pectoral lamina at approximately the
time when the hinge becomes functional as such.

Ontogenetic changes in the shell can be summarized as follows:

5—773

592 University of Kansas Publs., Mus. Nat. Hist.

1) Buttresses become less distinct in the first two years of life
(plastral lengths of 40 to 55 mm.); 2) Interdigitating processes of
the forelobes and hind lobes become relatively shorter and wider,
the entoplastron no longer projects posterior to the hinge, the lat-
eral apex of the pectoral lamina becomes creased, and some move-
ment of the plastron can take place between the second and third
years (plastral lengths of 55 to 65 mm.); 3) Plastral lobes become
freely movable upon one another and upon the carapace by the end
of the fourth year (plastral length approximately 70 mm.) in most
individuals.

The plastron of a juvenal box turtle is not completely immovable.
The bones of the shell are flexible for a time after hatching and
allow some movement of the plastron; but the relatively greater
bulk of the body in young box turtles would prevent complete clos-
ure of the shell even if a functional hinge were present. Hatchlings
can withdraw the head and forelegs only to a line running between
the anterior edges of the shell. To do so the rear half of the
shell is opened and the hind legs are extended. When the head
and forelegs are retracted to the maximum, the elbow-joints are
pressed against the tympanic region or behind the head; the fore-
limbs cannot be drawn part way across the snout, as in adults.
Hatchlings can elevate the plastron to an angle of approximately
nine degrees; the plastron of an adult, with shell closed, is elevated
about 50 degrees. Hatchlings flex the plastron chiefly in the
region of the humeropectoral seam, rather than at the anlage of
the transverse hinge.

Adult box turtles, when walking, characteristically carry the
forelobe of the plastron slightly flexed. This flexion of the plastron,
combined with its naturally up-turned anterior edge, cause it to
function in the manner of a sled runner when the turtle is moving
forward. A movable plastron, therefore, in addition to its pri-
marily protective function, seems to aid the turtle in traveling
through tall grass or over uneven ground. The gular scutes, on
the anterior edge of the forelobe, become worn long before other
plastral laminae do.

An adult female from Richland County, Illinois, had an abnormal
but functional hinge on the humeropectoral seam in addition to a
normal hinge on the pectoroabdominal seam. The abnormal hinge
resulted from a transverse break in which ligamentous tissue later
developed. The muscles closing the plastron moved the more
anterior of the two hinges; the normal hinge was not functional.

Natural History of the Ornate Box Turtle 593

Color and Markings

The markings of the shell change first when postnatal growth
begins and again when sexual maturity is attained. They are
modified gradually thereafter as the shell becomes worn.

In hatchlings the ground color ordinarily is dark brown but in
some individuals is paler brown or tan. Markings on the dark
background are pale yellow. Markings on the central and lateral
scutes vary from a regularly arranged series of well defined spots
and a middorsal stripe to a general scattering of small flecks. In
some specimens the pale markings of the carapace are faint or
wanting. Lateral parts of marginal scutes are always pale yellow
and form a border around the carapace.

Close examination of the carapace of any hatchling shows the
following basic arrangement of markings: each lateral scute has
a centrally placed pale spot and four to seven smaller pale marks
arranged around the edge of the scute; each central scute has a
central, longitudinal mark and several (usually two, four, or six)
smaller pale marks arranged around the edge of the scute, chiefly
the lateral edges (PI. 23). Variations in pattern result when some
or all of the markings divide into two or more parts.

By the end of the first full season of growth, the markings have
a radial pattern. At this stage, the markings of the areola, with the
exception of the central spot, are obscure. The radial marks,
sharply defined and straight-sided, appear only on tlie newly
formed parts of the epidermal laminae. Each radial mark orig-
inates opposite one of the peripheral marks of the areola. Other
radial marks are developed later by bifurcation of the original
radiations.

The ground color of the plastron of hatchlings is cream-yellow,
or less often, bright yellow. The solid, dark brown markings on the
medial part of each lamina form a central dark area that contrasts
sharply with the pale background (PI. 24). The soft tissue of the
navel is pale yellow or cream; when the navel closes, the dark cen-
tral mark of the plastron is unbroken except for thin, pale lines
along the interlaminal seams.

When growth begins, the areas of newly formed epidermal tissue
on the anterior and medial borders of each areolar scute are pale.
Wide, dark radial marks, usually three per scute, appear on the
newly formed tissue. Subsequently, finer dark radiations appear
between the three original radiations. The wdde radiations later
bifurcate. By the time adult or subadult size is reached, the plas-

594 University of Kansas Publs., Mus. Nat. Hist.

tron appears to have a pattern of pale radiations on a dark back-
ground. In general, the markings of the plastron are less sharply
defined than the markings of the carapace (PI. 24).

There is a tendency for the dark markings of the plastron to
encroach on the lighter markings, if no wear on the shell occurs.
However, as the plastron becomes worn, the pale areas become
more extensive and the dark markings become broken and rounded.
Severely worn plastra of some old individuals lack dark markings.
Wear on the carapace produces the same general efiFect; but mark-
ings of the carapace, although they may become blotched, are never
obliterated in Terrapene o. omata.

The top of the head in most hatchlings is dark brown, approxi-
mately the same shade as the ground color of the carapace; the
part anterior to the eyes is usually unmarked but a few individuals
have a semicircle of small pale spots over each eye or similar spots
on much of the head. The posterior part of the head is ordinarily
flecked with yellow. The skin on the top of the head, particularly
between the eyes, is roughened. The granular skin of the neck is
grayish brown to cream-yellow. There are one or two large pale
spots behind the eye and another pale spot at the corner of the
mouth. Smaller, irregularly arranged pale markings on the necks
of some specimens form, with the post-orbital and post-rictal spots,
one or two short, ragged stripes. The gular region is pale.

In juveniles, the yellow markings of the head and neck are larger
and contrast more sharply with the dark ground color than in hatch-
lings. Markings above the eyes, if present, fuse to form two pale,
semicircular stripes. In some older juveniles yellow marks on top
of the head blend with the dark background to produce an amber
color. The top of the neck darkens or develops blotches of darker
color that produce a mottled effect. Spots and stripes on the side
of the neck remain well defined. The skin on top of the head be-
comes smooth and shiny.

Adult females tend to retain the color and pattern of juveniles
on the head and neck although slight general darkening occurs
with age. Many adult females have the top of the head marked
with bright yellow spots. In adult males, the top and sides of the
head, anterior to the tympanum, are uniformly grayish green or
bluish green; the mandibular and maxillary beaks are brighter,
yellowish green. Markin<!;s on the head and neck of most adult
males arc obscure (PI. 25) but the sides of the neck remain mottled
in some individuals.

The antebrachium has large imbricated scales and is distinctly

Natural History of the Ornate Box Turtle 595

set oflF from the proximal part of the foreleg which is covered with
granular skin. The antebrachial scales of hatchhngs are pale yel-
low; each scale is bordered with darker color. General darkening
of the antebrachium occurs at puberty. In adult females each scale
on the anterior surface of the antebrachium is dark brown and has
a contrasting yellow, amber, or pale orange center. The anterior
antebrachial scales of adult males are dark brown to nearly black
and have bright orange or red centers. Old males have thickened
antebrachial scales.

The iris of hatchlings and juveniles is flecked with yellow and
brown; the blending of these colors makes the eye appear yellow,
golden, or light brown when viewed without magnification. Adult
females retain the juvenal coloration of the eye; the iris of adult
males is bright orange or red. The work of Evans (1952) on T.
Carolina suggests that eye color in box turtles is under hormonal
control.

Wear

Presence or absence of areolae on laminae of the shell indicated
degree and sequence of wear. The anterior edges of carapace
and plastron, and the slightly elevated middorsal hne ( Pi. 23 ) wear
smooth in some individuals before the first period of hibematioii.
Subsequent wear on the carapace proceeds posteriorly. For ex-
ample, turtles that retained the areola of the third central lamina,
retained also the areolae of the fourth and fifth centrals; when
only one central areola remained, it was the fifth. Lateral laminae
wear in the same general sequence. The areola of the fifth central
lamina, because of its protected position, persists in adult turtles
that are well past the age of regular growth. Areolae that are re-
tained in some older turtles are shed along with the epidermal
layers formed in the first year or two of life. Wear on the shell is
probably correlated with the habits of the individual turtle;
smoothly-worn specimens varied in size and age but were usually
larger, older individuals. No smoothly worn individual was still
growing.

Wear on the plastron is more evenly distributed than wear on
the carapace; wear is greatest on the lowest points of the plastron
(the gular laminae, the anterior portions of the anal laminae, and
the lateral edge of the tranverse hinge ) .

The claws and the homy covering of the jaws are subject to
greater wear than any other part of the epidermis; presumably
they continue to grow throughout Hfe. The occasional examples
of hypertrophied beaks and claws that were observed, chiefly in

596

University of Kansas Publs., Mus. Nat. Hist.

juveniles, were thought to result from a continuous diet of soft
food or prolonged activity on a soft substrate. Ditmars (1934:44,
Fig. 41) illustrated a specimen of T. Carolina, with hypertrophied
maxillary beak and abnormally elongate claws, that had been kept
in a house for 27 years.

The conformation of the maxillary beak in all species of Terra-
pene is influenced to a large extent by wear and is of limited value
as a taxonomic character. The beak of T. ornata is slightly notched
in most individuals at the time of hatching and remains so through-
out life. The underlying premaxillary bone is always notched or
bicuspidate. The sides of the beak are more heavily developed
than the relatively thin central part. Normal wear on the beak
maintains the notch (or deepens it) in the form of an inverted
U or V, much in the manner of the bicrenate cutting edge on the
grooved incisors of certain rodents. In a series of 34 specimens of
r. ornata from Kansas, selected at random from the K. U. col-
lections, 92 per cent had beaks that were "notched" to varying
degrees, four per cent had hooked (unnotched) beaks, and four
per cent had beaks that were flat at the tip (neither hooked nor
notched ) .

Fig. 21. Plantar views of right hind foot (male at left,
female at right) of T. o. ornata ( X 1), showing sexual
dimorphism in the shape and position of the first toe.
The widened, thickened, and intumed tenninal phahmx
on the first toe of the male is used to grasp the female
before and during coitus.

SEXUAL DIMORPHISM

Difl^crences between adult males and females of T. ornata have
been mentioned in several places in the preceding discussion of
growth and development. Several sexual characteristics — greater

Natural History of the Ornate Box Turtle

597

preanal length, thickened base of the tail, slightly concave plastron,
and smaller bulk — are found also in males of many other kinds
of emyid turtles. From females, males of T. ornata are most easily
distinguished by the bright colors of their eyes, heads, and ante-
brachial scales. An additional, distinctive characteristic of males
is the highly modified hind foot. The first toe is greatly thickened
and widened; when the foot is extended, the first toe is held in
a horizontal plane nearly at right angles to the medial edge of the
plantar surface ( Fig. 21 ) . The hind foot of females is unmodified
in this respect. Males tend to have heavier, more muscular hind
legs than females.

The bright colors of males are maintained throughout the year
and do not become more intense in the breeding season. Males
of T. o. hiteola become melanistic in old age whereas males of tlie
subspecies ornata do not. In old males of luteola the skin be-

Table 4. — A Summary of Sexual Dimorphism in Terrapene ornata.

Character

Males

Females

Head .... . . . .

Snout truncate in lateral
profile, top of head and front
of maxilliary beak forming an
angle of nearly 90°; head j^el-
lowish green to bluish green;
markings on head and neck
reduced; head never spotted
dorsally (PI. 11, Figs. 7 and 8).

Snout relatively round iu

lateral profile; front of maxil-
lary beak not forming right
angle with top of head; head
dark brown, distinct pale
markings on head and neck;
head commonly spotted dor-
sally (PI. 25, Figs. 5 and 6).

Iris

Red

Yellowish brown

Hind legs

Heavy and muscular; first
toe turned in, thickened, and
widened (Fig. 21).

Not especially heavy or
muscular; first toe, if turned
in, never thickened or widened
(Fig. 21).

Forelegs

Centers of antebrachial
scales bright orange or red.

Centers of antebrachial
scales yellow, pale orange, or
brown.

Carapace

Relatively lower, length con-
tained in height (48 specimens)
.58 times (±.005<Tm, range,
.50 to .69).

Relatively higher, length
contained in height (94 speci-
mens) .50 times (±.005(Tm,
range .44 to .60).

Plastron

(hind lobe)

Ordinarily slightly concave.

Flat or convex, never con-
cave.

598 University of Kansas Publs., Mus. Nat. Hist.

comes dark gray, bluish, or nearly black and much of the bright
orange or red of the antebrachial scales and the green of the head
is obliterated; the iris may also darken but in most specimens it
retains some red. Females of luteola tend also to darken some-
what in old age but not so much as males; females of ornata do
not. Table 4 summarizes the more important secondary sexual
characters of T. ornata.

TEMPERATURE RELATIONSHIPS

Tolerances to environmental temperatures, and reactions to
thermal stimuH influence the behavior of ectothermal animals to a
large extent. Terrapene ornata, like other terrestrial reptiles in-
habitating open grassland, is especially subject to the vicissitudes
of environmental temperature. Other species of turtles Hving in
the same area are more nearly aquatic and therefore live in a micro-
habitat that is more stable as regards temperature.

Approximately 500 temperature readings in the field and many
others in the laboratory were obtained from enough individuals to
permit interpretation of reactions involved in basking, in seeking
cover, and in emerging from temporary periods of quiescence at
various times of the day.

Box turtles commonly used open places such as cow paths,
ravines, and wallows, for basking as well as for feeding and as routes
of travel. Burrows, dens beneatli rocks, and forms, were used as
shelter from high and low temperatures as well as from predators.
Determining whether a turtle was truly active ( moving about freely,
feeding, or copulating), was basking, or was seeking shelter was
diflScult because the turtle sometimes reacted to the observer; for
instance, basking turtles, whose body temperatures were still sub-
optimum, might take cover when surprised, and truly active turtles
might remain motionless and appear to be basking. By scanning
open areas from a distance with binoculars, an observer frequently
could determine what turtles were doing without disturbing them.
In the final analysis of data, temperature records accompanied by
data insufiicient to determine correctly the state of activity of the
turtle, were discarded, as were temperature records of injured
turtles and turtles in livetraps.

Cowles and Bogert (1944:275-276) and Woodbury and Hardy
(1948:177) emphasized the influence of soil temperatures on body
temperatures. It is thought that air temperatures played a more
important role than soil temperatures in influencing body tempera-

Natural History of the Ornate Box Turtle 599

tures of T. ornata. Soil temperatures were taken in the present
study only when the turtle was in a form, hibernaculum, or den.

Optimum Temperature

Cowles and Bogert (1944:277) determined optimum levels of
body temperature of desert reptiles by averaging body temperatures
falling within the range of normal activity; they defined this range
as, ". . . extending from the resumption of ordinaiy routine
[activity] ... to ... a point just below the level at
which high temperatures drive the animal to shelter." Fitch
(1956b: 439) considered optimum body temperature in the several
species that he studied to be near the temperature recorded most
frequently for "active" individuals; he found {loc. cit.) that of body
temperatures of 55 active T. ornata, 66 per cent were between 24
and 30 degrees, and that the temperatures 27 and 28 occurred most
frequently. Fitch concluded (op. cji.:473) that the probable op-
timum body temperature of T. ornata was 28 degrees and that
temperatures from 24 to 30 degrees were preferred. Although Fitch
treated all non-torpid individuals that were abroad in daytime as
"active" and did not consider the phenomenon of basking, his
observations on optimum body temperature agree closely with my
own.

Body temperatures of 153 box turtles that were known definitely
to be active, ranged from 15.3 to 35.3 degrees. The mean body
temperature for active turtles was 28.8 degrees ( ±: 3.78(t) (Fig. 22).
Ninety-two per cent of the temperatures were between 24 and 30
degrees and 50 per cent were between 28 and 32; temperatures of
29 and 30 degrees occurred most frequently (22 and 21 times,
respectively). The ten body temperatures below 24 degrees all
were recorded before 9 A. M. on overcast days when the air was
cool and humid. It is notewortliy that two of these low tempera-
tures (18.8° and 19.0°) were from a copulating pair of turtles; two
others (21.8° and 22.0°) were from individuals that were eating.
The highest temperature (35.3°) was from a large female that was
feeding at mid-morning in a partly shaded area.

The mean body temperature for active individuals (Fig. 22)
is probably somewhat below the ecological optimum, because a
few temperatures were abnormally low. The large number of
body temperatures in the range of 29 to 31 degrees indicates an
optimum closer to 30 degrees. Optimum body temperatures may
vary somewhat with the size, sex, or individual preference of the
turtle concerned.

600 Univebsity of Kansas Publs., Mus. Nat. Hist.

Basking

Although basking is common in terrestrial turtles, only a few
authors have mentioned it, Woodbury and Hardy (1948:177-178)
did not use the term in their account of thermal relationships in
Gopliertis agossizi; their discussion indicates, however, that the
tortoises move alternately from sunny to shady areas to regulate
body temperature. Desert tortoises removed from hibernacula and
placed in the sun warmed to approximately 29.5 degrees before
they became active, although a few did so at temperatures as low
as 15 degrees. According to Cagle (1950:45), Sergeev (1939)
studied body temperature and activity in the Asiatic tortoise,
Testiido horsefieldi, and found that individuals basked for as
much as two hours in the morning before beginning the first ac-
tivity of the day (feeding), but that tortoises did not bask after a
period of quiescense from late morning to late afternoon, during
which body temperatures were seemingly maintained nearer the
optimum than they were during nocturnal rest; body temperatures
rose to approximately 30 degrees before the tortoises became ac-
tive. Since body temperatures of 23 to 24 degrees were main-
tained at night, the basking range of Testndo horsefieldi may be
considered to be approximately 23 to 32 degrees.

Ornate box turtles basked chiefly between sunrise and 10 or
11 A. M. Body temperatures of 60 basking turtles ranged from
17.3 to 31.4 degrees (mean, 25.5 ± 3.0Sa). More than two-thirds
(42) of these body temperatures were higher than the air tem-
perature near the turtle, indicating probably that body temperature
rises rapidly once basking is begun. In tlie instances where body
temperature was below air temperature, the turtles had recently
begun to bask ( many were known to have just emerged from forms
or other cover where they had spent the night) or were warming
up more slowly because of reduced sunlight. On cloudy days bask-
ing began later than on clear days and body temperatures usually
remained at a suboptimum level. Turtles that basked on days
that were cloudy and windy, or cold and windy, did so in sheltered
places, usually on the leeward sides of windbreaks such as lime-
stone rocks, rock fences, or ravine banks. It was evident in these
instances that the turtles either sought such shelter from the wind
or remained ensconced in the more complete shelter of a form
or burrow, not emerging at all.

Open areas of various kinds were used as basking sites. Level
ground — such as on roads, cattle pathways, and bare areas sur-

Natural History of the Ornate Box Turtle 601

rounding farm ponds — having unobstructed morning sunlight,
nearby dense vegetation, and choice opportunities for feeding
(cow dung, mullberry trees) was preferred. Basking was fre-
quently combined with feeding; in several instances box turtles
were noted early in the morning at suboptimum body temperatures
eating grasshoppers, berries, or dung insects. The predilection of
box turtles for open areas is probably important in permitting ex-
tended activity at suboptimum temperatures. T. ornata probably
carries on more nearly normal activity on cool days than do reptihan
species with more sharply delimited thermal tolerances. Collared
lizards (Crotaphytus collaris), for example, are chiefly inactive on
days when the sky is overcast, although a few individuals having
suboptimum body temperatures can be found in open situations
(Fitch, 1956a: 229 and 1956b: 442).

Toleration of Thermal Maxima and Minima

The foregoing remarks on basking indicate the approximate,
normal, thermal tolerances of ornate box turtles. Many additional
records of body temperature were taken from turtles that were
found under cover. Turtles under cover in daylight were usually

under cover

(low temperature) N= 78

basking N = 60

^

normally active N = I53

^

under cover (high temperoture) N=64

•+-

■ '■■'■' I ■ I ■ I I I I ■ I

■•■'*'

0 5 10 15 20 25 30 35

BODY TEMPERATURE

Fig. 22. The relationship of body temperature (Centigrade) and kind
of activity in T. o. ornata, compiled from 355 field observations. Vertical
and horizontal lines represent, respectively, the range and mean. Open
and solid rectangles represent one standard deviation and two standard
errors of the mean, respectively.

602 UNivERsrri' of Kansas Publs., Mus. Nat. Hist.

seeking protection from either below-optimum or above-optimum
temperatures. In avoiding low temperatures, turtles usually chose
more complete and permanent cover than in avoiding high tem-
peratures.

Body temperatures of 64 box turtles that were seeking cover or
that were under cover because of high temperatinres ranged from
28.9 to 35.8 degrees (mean, 31.9 ± 1.55o-). Fifty-nine of these tem-
peratures (92 per cent) were 30 degrees or higher. Figure 22
shows this range to overlap broadly with the temperature range of
active turtles and the means of the two groups are close to each
other. Body temperatures below 30 degrees (5) were all recorded
late in the morning on hot summer days when the air temperature
was well above 30 degrees; they are somewhat misleading because
they are from turtles that were under cover long enough to lower
body temperature to the range of activity although the turtles re-
mained under cover because of hazardous environmental tempera-
tures.

The commonest retreats used by box turtles to escape heat were
burrows of other animals and small dens under thick limestone
rocks, where the air remained cool, even in late afternoon. Most
of the burrows and dens on the Damm Farm were known to me
and could be checked each day. Turtles seeking temporary refuge
from high temperatures characteristically rested just inside the
opening of a den or burrow. Less frequently, turtles burrowed into
ravine banks or just under the sod on level ground. A number of
individuals with above-optimum body temperatures were found in
the shade of trees or high weeds in early afternoon on hot days.
Mulberry trees provided ample shade for such activity and, in
June and July, when ripe mulberries were abundant on the ground,
turtles frequently fed on them at times of the day when tempera-
tures were more hazardous in other areas.

Several turtles were found buried in mud or immersed in water
at the edges of ponds in the hottest part of the day; they were dis-
covered at first by accident and, on subsequent field trips by sys-
tematic probing. Ordinarily the turtles were covered with mud or
muddy water and remained motionless, except for periodically
raising the head to the surface to breath. There was little vegeta-
tion near the edges of ponds and by late morning on hot days the
temperature of the shallowest water was as high as the air tempera-
ture or higher. Correspondingly, turtles found resting in mud and
water had body temperatures much higher than turtles in dens,
burrows, or forms at the same time of day. Box turtles that retreat

Natural History of the Ornate Box Turtle 603

to mud or shallow water cool themselves less efficiently than they
would in drier, better protected microhabitats. I found no evidence
that turtles went into deeper water to cool themselves.

The length of time spent under cover varied; most turtles had
two daily periods of activity, the second beginning in late afternoon.
Some ti-irtles moved from shelter to shelter in the time between
periods of activity. Several turtles were known to remain quiescent
continuously for several days in the hottest part of the summer.

The maximum temperature that a reptile can tolerate physio-
logically is ordinarily higher than the maximum temperature tol-
erated voluntarily (Cowles and Bogert, 1944:277); but, the two
maxima may be separated by only a few degrees. Most poikilo-
thormous vertebrates neither tolerate nor long survive body tem-
peratures exceeding 40 degrees (Cowles and Bogert, op. cit.:269).

It is evident (Fig. 22) that ornate box turtles do not often toler-
ate body temperatures above 33 degrees and that temperatures in
excess of 35 degrees are probably never tolerated under natural
conditions. At 9:15 A. M. on July 5, 1955, an adult female emerged
from mud where she had spent the night (body temperature 28.4°,
mud 28.4°, air 30^^). After foraging for 40 minutes in bright sun-
light on a grassy hillside she had moved approximately 100 feet
and her temperature had reached 34.6 degrees (air 33.0°). At
9:56 A. M. she moved rapidly and directly to a den under a rock
nearby; 15 minutes later her body temperature had not changed
but after 65 minutes it had dropped to 33.4 degrees. The tempera-
ture of air in the den was 31 degrees. This female began her activi-
ties at nearly optimum body temperature relatively late in the
morning and, by foraging intensively for less than one hour, prob-
ably was able nearly to satisfy her daily food requirements; by for-
aging near suitable cover she could remain active until her body
temperature reached a critical threshold, and she thereby saved
time otherwise required for finding cover or making a form.

The following observations, extracted from field notes, indicate
that body temperatures near 40 degrees are the approximate lethal
maximum and are well above those temperatures voluntarily tol-
erated by T. ornata. On July 4, 1955, a subadult female was in the
water at the edge of a pond. The temperatures of the air, water,
and turtle were 32.0, 30.6, an 30.2 degrees, respectively. At 11
A. M. the turtle was tethered in direct sunlight on the hard-baked
clay of the pond embankment (temperature of air 33.4°). The
turtle's response to steadily rising body temperature over a period
of 31 minutes is illustrated by the following notes.

Time (A.M.)

Body
temperature

11:00

33.0

11:05

34.6

11:09

36.5

11:27

39.4

11:30

39.6

11:31

604 Uxn'ERSiTY OF Kansas Publs., Mus. Nat. Hist.

Remarks

Tethered on slope.

Strains at tether in several directions.

Tries frantically to get away; draws in limbs and
head rapidly and moncntaril>- at any movement on
my part, and hisses loudly.

11:13 37.5 Mouth held open slightly; turtle overturns in effort

to escape; frantic scrambling resumed a few sec-
onds alter I right turtle.

11:17 38.2 Mouth now held open most of the time; white

froth begins to appear around mouth.

11:20 38.6 Stops activities every 10 seconds or so, rests chin

on ground and gapes widely; will still pull into
shell when prodded with stick.

11:23 39.2 Still wildly active; continues to gape widely every

few seconds.

Frothing at mouth profusely.

Attempts to escape are now in short feeble bursts.

Turtle released; crawls toward me and immediately
seeks shade of my body; when I move off, turtle
seeks shade of small isolated weed on pond em-
bankment; turtle removed to damp earth at edge
of pond.

11:35 39.5 Attempts to burrow into mud at edge of pond.

11:36 Enters shallow water and moves slowly back to

shore.

11:37 38.8 Turtle thrown into center of pond where it re-

mains motionless and drifts with wind to opposite
shore; remains inactive in mud and shallow water
at edge of pond; temperature of water near turtle
35.5.

11:57 35.0 Moves 50 ft. up slope to shade of low vegetation.

1:55 P.M. 32.5 Turtle has not moved.

The overheating may have incapacitated the turtle since it
moved only 50 feet in the next two days; its body temperatures
on the two days subsequent to the experiment were 26.8 and
20.6, respectively.

The mentioned gaping, as in higher vertebrates generally, cools
the animal by evaporation from the moist surfaces of the mouth
and pharynx. By keeping the mouth open for more than a few
minutes at a time in hot dry weather, a turtle would surely lose
body water in amounts that could not always be easily replaced.
Ornate box turtles seem to utilize evaporation for cooling only in
emergencies and rely for the most part on radiation and conduc-

Natural History of the Ornate Box Turtle 605

tion to lower body temperature after reaching a relatively cool,
dark retreat.

Box turtles were never active at body temperatures below 15
degrees and were seldom active at temperatures below 24 degrees.
The two lowest temperatures (15.3° and 16.3°) were taken from
individuals crossing roads on overcast days in early May.

In 78 box turtles that were under cover because their environ-
mental temperatures were low, the body temperatures ranged from
2.7 to 30.6 degrees (mean 19.8 =h 6.38o-). The range of body tem-
peratures in this group is greater than in the other groups shown in
Figure 22 because low body temperatures were studied over a wide
range of conditions, including hibernation.

Box turtles actually seek cover because of low temperatures only
in fall and spring and on occasional unseasonable days in summer
when temperatures drop rapidly. Retreat to cover, in the normal
cycle of daily activity, is governed usually by high temperatures
at midday or by darkness at the end of the day. Turtles in dens,
burrows, and grass forms, tended to burrow if temperatures re-
mained low for more than a few hours.

Box turtles under cover where they cannot bask have little con-
trol over the lower range of body temperatures. The freezing
temperatures of winter can be escaped by burrowing deeper into
the ground. Temperatures approaching the lethal minimum, how-
ever, seldom occur during the season of normal activity. By re-
maining hidden in a burrow or den therefore, box turtles are fairly
well protected from predators but are at a thermal disadvantage.

A number of turtles that had wet mud on their shells were
found basking in early morning near ditches, ponds, and marshy
areas; several others were partly buried in mud, shortly after day-
break, and another v/as at the edge of a pond after dark.

Eight adults, located just as they emerged from cover in early
morning on sunny days, had body temperatures of 19.7, 21.9, 24.2,
24.5, 25.8, 26.6, 28.7, and 29.5 degrees. In five emerging from earth
forms, body temperatures were at least a degree or two below the
temperature of the air; the other three came from mud or shallow
water and had body temperatures higher than the air tempera-
ture.

Temperature is probably the primary stimulus governing emer-
gence after temporary periods of quiescence. Turtles in earthen
forms are usually completely covered or are head downward witli
only the hind quarters exposed. Obviously, the more thoroughly

606 University of Kansas Publs., Mus. Nat. Hist.

a turtle protects itself (beneath the insulating cover of a form,
burrow, or den) against unfavorable temperatures, the longer it
will take for favorable temperatures to bring about normal ac-
tivity again. Turtles in forms and deep burrows have a minimum
of contact with the outer environment; but in dens beneath rocks
and in shallow burrows light and air can enter freely. Turtles
might be influenced in their activities to some extent by the in-
tensity of light at the opening of a burrow or den; they are surely
stimulated by changes in the temperature and humidity of air
coming through the opening. Shallow retreats that a turtle can
enter and leave with the least efiFort therefore seem most efiBcient
for purposes of thermocontrol, especially when they provide earthen
surfaces into which the turtles can burrow more deeply if more
severe environmental conditions develop.

In October, 1955, nine T. ornata of various sizes, collected in
Douglas County, Kansas, were brought to the laboratory for ob-
servation under conditions of controlled temperature. They were
kept at room temperature for several days and were fed regularly,
with the exception of one hatchling that was fed nothing in this
period. On October 22 the turtles were placed in a room where
the temperature was maintained constantly at zero degrees. One of
the nine turtles, an adult female, was killed with chloroform im-
mediately prior to its removal to the cold room. A list of the turtles
used in this experiment is given below.

Age Carapace Weight

class length in mm. in grams

1) Hatchling 33.1 8.4

2) Hatchling * 29.9 6.7

3) Juvenile 52.5 29.3

4) Juvenile 50.2 26.1

5) Adult 5 125 376

6) Adult 9 118 400

7) Adult S 119 386

8) Adult 9 110 325

9) Adult 9 115

* Starved.

Turtles were kept in the cold room for periods of 100 minutes
(hatchlings and juveniles) and 200 minutes (adults). The entire
experiment, including the time in which the turtles were allowed
to warm after they were taken from the cold room, covered a
period of nearly six hours (375 minutes) during which the turtles
were under constant observation. Individual body temperatures
were taken continuously in this period (39 for each juvenile and
24 for each adult) in the order that the turtles were numbered;
gaps between records of the body temperature of a given indi-

Natural History of the Ornate Box Turtle

607

vidual therefore represent the time required to record temperatures
for the rest of the turtles in the group. The rates of rise and fall
of temperature for each of the nine turtles considered are shown
as a graph in Figure 23. Rate of temperature change was inversely
proportional to bulk; hatchlings, for example, cooled and warmed
a httle more than twice as rapidly as did adults. Rate of tempera-
ture change was intermediate in juveniles but was more nearly like
that of adults in the warming phase and closer to that of hatchhngs
in the cooling phase ( Table 5 ) .

Considering that hatchling no. 2 was smaller than no. 1, the rate
of change in its temperature did not seem to be significantly al-
tered by starvation. The adult males showed a tendency to change
temperature faster than adult females even though both males were
larger than any of the females. The slight difiFerence in rate of
temperature change between the sexes (Fig. 23) may have been
fortuitous.

Table 5. — Average Rate of Change in Temperatube (Expressed in De-
grees PER MINUTE ) FOR FOUR GrOUPS OF TURTLES SUBJECTED TO TEMPERATURE

OF Zero Degrees and then Allowed to Warm at 27 Degrees (Centigrade).

Group

Hatchlings
Juveniles . ,
Adult c? . .
Adult 9 . .
Adult (all)

Number

2

2

2
3
5

Cooling
phase

.282
.264
.122
.119
.120

Warming

phase

(to 25")

.310

.180

.152

.130*

.138

* None of the females reached a temperature of 25° before the experiment was terminated.

One hatchling ( No. 1 ) , when its temperature dropped below one
degree, fully extended all four limbs and the body was elevated
and only the anterior edge of the plastron was in contact with the
confining glass dish. Raising the body from an uncomfortably cold
or hot substrate is a well known phenomenon in many lizards and
in crocodihans, but to my knowledge has not been reported for
turtles.

Hibernating turtles and those experimentally chilled were usually
comatose but were almost never completely incapacitated, even at
temperatures at or near zero degrees. Experimental pinching,
probing, and pulling revealed that muscles operating the neck, the

6—773

608

U.vivERsiTi' OF Kansas Publs., Mus. Nat. Hist.

time

in

60

30 1 — I— J — I — 1—1 ' ' ' » ' I

120

I I I

Fig. 23. Changes in temperature of the body of four juvenal (nos. 1 to 4)
and five adult individuals of T. o. ornata (nos. 5 to 9) exposed to a constant
air temperature of zero degrees Centigrade for periods of 100 and 200 minutes,
respectively. The vertical arrows indicate when the turtles were removed to

Natural. History of the Ornate Box Turtle

609

J I L

240
j_J-

minutes

300

360

I I

Jl L

I

J i I L

J i— L

<^v^

an air temperature of 27 degrees. Sizes and weights of the turtles used are
given in the text. Turtle number nine, a female, was kiUed by means of
chloroform before experiment began. Rate of change in temperature in speci-
mens was inversely proportional to size. All turtles survived the experiment.

610 University of Kansas Publs., Mus. Nat. Hist.

limbs, and the lobes of the plastron could be controlled by the turtle
at low temperatures; hissing, resulting from rapid expulsion of air
through tlie mouth and nostrils ( when the head and limbs are drawn
in reflexively ) occurred at all body temperatures but was sometimes
barely audible in the coldest turtles. Of all living turtles observed,
only two (hatchlings 1 and 2 in coldroom experiment) were com-
pletely immobile at low temperatures, failing to respond even to
pinpricks at body temperatures of 0.8 and 1.7 degrees, respectively,
although other turtles, under the same experimental conditions,
consistently gave at least some response to the same stimulation.

Turtles chilled experimentally continued to move about volun-
tarily, albeit sluggishly, at temperatures much lower (2.5° for each
of four adults; 10.0° and 6.2° for two juveniles) than those at which
locomotion was resumed in the warming phase ( 13° for the adults,
21.7° and 20.1° for the juveniles). Hatchlings chilled so rapidly
that it was difiRcult to ascertain accurately the temperature at which
inactivity was induced. Juveniles became active gradually, moving
slowly about when the body temperature reached approximately 20
degrees but not attempting more strenuous activities such as
climbing the walls of enclosures, until body temperatures of 22 to
25 degrees were attained. Adults, on the other hand, exhibited
**normar activity as soon as they became voluntarily active.

The ability of ornate box turtles to move about when the body
temperature is near the lethal minimum probably enables those
caught in the open by a sudden drop in environmental temperature
to find cover that keeps them from freezing to death. Prolonged
chilling, on the other hand, seems to create a physiologically dif-
ferent situation; the temperature at which activity is resumed is
higher and subject to less variation.

Juveniles were more rapidly aflFected by environmental tempera-
tures, were subject to different thresholds, and were inactive over
a wider range than were the adults. Indeed, the rate of chilling,
rather than absolute body temperature alone, might in large measure
influence the reactions of turtles to environmental temperatures.
If this be so, smaller turtles, having a narrower thermal range of
normal activity, must lose at least some of the advantages gained
by their ability to warm up more rapidly.

Hatchlings and juveniles at the Damm Farm were always active
on days when at least some adults were also active. Fitch
(1956b:466) found that, in northeastern Kansas, species of small
reptiles and amphibians are active earlier in the season than larger

Natural History of the Ornate Box Turtle 611

species and that the young of certain species become active earher
than adults. Fitch stated, ". . . small size confers a distinct
advantage in permitting rapid rise in body temperature by contact
with warmed soil, rock or air, until the threshold of activity is at-
tained"; he pointed out also tliat young animals, if able to emerge
earlier than adults, would benefit from a longer growing season.
Hatchlings and juveniles of T. ornata would benefit greatly from an
extra period of activity of say, one or two weeks in spring and a
similar period in autumn, especially if food were plentiful. The
extra growth realized from such a "bonus" period of feeding would
significantly increase the chance of the individual turtle to survive
in the following season of growth and activity.

Ornate box turtles are active within a narrower range of tempera-
tures than are aquatic turtles in nearby ponds and streams of the
same region. Observations by William R. Brecheisen and myself
on winter activity of aquatic turtles indicate that, in Anderson
County, Kansas, the commoner species {Chelydra serpentina,
Chrysemys picta, and Pseudemys scripta) are more or less active
throughout the year; although they usually do not eat in winter,
they are able to swim about slowly and in some instances (P.
scripta) even to carry on sexual activity at body temperatures only
one or two degrees above freezing. But, ornate box turtles hiber-
nating in the ground a few yards away are incapable of purposeful
movement at such low body temperatures.

HIBERNATION

In northeastern Kansas ornate box turtles are dormant from late
October to mid-April — approximately five and one half months of
the year. Individuals may be intermittently active for short periods
at the beginning and end of the season, however. Once a perma-
nent hibernaculum is selected dormancy continues until spring;
unseasonably warm weather between mid-November and March
does not stimulate temporary emergence. There is little movement
during dormancy except for the deepening or horizontal extension
of the hibernaculum.

Woodbury and Hardy (1948:171) found desert tortoises {Go-
pherus agassizi) in dormancy from mid-October to mid- April in
southwestern Utah; some tortoises became temporarily active on
warm days in winter. Cahn (1937:102) was able to compare hiber-
nation in several individuals each of T. ornata and T. Carolina, kept
under the same conditions in Illinois. Individuals of T. ornata bur-
rowed into the ground in October, two weeks before those of T. car-

612 University of Kansas Publs., Mus. Nat. Hist.

olina did, and continued to burrow to a maximum depth of 223i
inches. Some individuals of T. Carolina spent the entire winter in
the mud bottom of a puddle and became semiactive on warm winter
days. Other individuals of T. Carolina burrowed nearly as deeply as
did T. ornata. Individuals of T. ornata emerged from hibernation
one or two weeks later in the spring than did those of T. Carolina.
There are some indications that populations of T. Carolina in eastern
Kansas are dormant for a shorter period of time than those of T. or-
nata but comparative studies are needed to verify this. Richard B,
Loomis gave me a large female of T. Carolina that he found active
beside a highway in Johnson County, Kansas, on November 23,
1954; on that date most individuals of T. ornata under my observa-
tion had already begun permanent hibernation but a few at the
Reservation were still semiactive.

Fitch (1956b: 438) listed earliest and latest dates on which box
turtles were active at the Reservation in the years 1950 to 1954;
in the five year period box turtles were active an average of 162
days per year (range, 140-187) or approximately 5.3 months of
the year. It is significant that 1954, having the most days of ac-
tivity was, according to my studies of growth-rings, an exceptionally
good year for growth. Fitch's data indicate the approximate sea-
son of growth and reproduction but not of total activity, since he
did not take into account the sporadic movements of box turtles in
late fall and early spring.

Activity in autumn is characterized by movement into ravines and
low areas; many turtles move into wooded strips along the edges
of fields or small streams. Sites protected from wind, providing
places for basking and for burrowing, are sought. Burrows of
other animals, along the banks of ravines, were often used for tem-
porary shelter; overhanging sod at the lips of ravine-banks provided
cover beneath which turtles could easily burrow. After mid-Octo-
ber progressively fewer box turtles were found in open places and
activity was restricted to a few hours in the warmest part of the
day.

Low air temperature probably is the primary stimulus for hiber-
nation. Autumn rains are usually followed by a decrease in gen-
eral activity. Rain probably hastens burrowing by softening the
ground.

Ornate box turtles more often than not excavate their own hiber-
nacula. Digging begins with the excavation of a shallow form
which is deepened or extended horizontally over a period of days
or weeks. Such hibernacula arc sometimes begun at the edges of

Natural History of the Ornate Box Turtle 613

rocks or logs; the overhanging edge of an unyielding object acts as
a fulcrum on the shell and hastens digging. Ornate box turtles
are slow but efficient burrowers.

Forms in open grassy areas are begun at an angle of 30 to 40
degrees; an adult box turtle requires approximately one hour to
burrow far enough beneath the sod to conceal itself but can dig into
soft, bare earth much more rapidly. Once a hibernaculum is
begun, all four feet are used for its excavation, the front feet doing
most of the digging and the hind feet pushing loose earth to the
rear.

Several turtles were seen entering burrows and dens in late au-
tumn and trailing records showed that some individuals visited
several of these shelters in the course of a single day.

By means of systematic probing of known hibernacula it was
found that they are deepened gradually in the course of the winter.
Depth seems to be governed by the temperature of the soil. Hiber-
nacula in wooded or sheltered areas were ordinarily shallower than
hibernacula in open grassland.

In the autumn of 1953-54 two pens were constructed at the Res-
ervation in order to study hibernation; one pen was on a wooded
hillside and the other was on open grassland. Turtles in the grass-
land pen were in newly excavated hibernacula, just beneath the
sod, on October 25 and did not emerge for the remainder of the
winter, whereas turtles in the woodland pen were intermittently
active until November 10. Correspondingly, turtles in the grass-
land pen descended to depths of eight and one half and 11/2 inches,
respectively, whereas those in the woodland pen were covered by
a scant six inches of loose earth and leaf litter. In 1954 four turtles
were traced (by means of trailing threads) to hibernacula on
wooded slopes at the Reservation; two entered permanent hiber-
nacula on November 13 and two remained semiactive until some-
time after November 20. All four turtles spent the winter in hiber-
nacula that were not more than six inches deep. Temperatures of
the soil at a detph of nine inches were usually slightly lower at the
grassland pen than at the woodland pen on a given date. It is
probably significant that individuals with trailing devices and indi-
viduals in experimental pens furnish the latest records for autumn
activity. The unnatural conditions created by confining the turtles
in pens restricted the number of hibernation sites that were avail-
able to them; although trailing devices did not a£Fect the normal
movements of box turtles on the surface of the ground these de-
vices certainly hampered the turtles somewhat in digging. How-

614 University of Kansas Publs., Mus. Nat. Hist.

ever, it is noteworthy that box turtles are able to move about after
mid-November, whether this is of general occurrence under more
natural conditions or not. Depths of hibemacula at the Damm
Farm were also influenced by amount of vegetation or other cover.
Maximum depth of hibernacula in more or less open situations
ranged from seven to 18 inches whereas a female hibernating in a
ditch that was covered with a thick mat of dead grasses was four
inches beneath the surface of the soil, and another female was only
two and one half inches below the floor of a den.

Several T. ornata kept by William R. Brecheisen in a soil-filled
stock tank on his farm in the winter of 1955-56, burrowed to maxi-
mum depths of seven to eight inches in the course of the winter.
A layer of straw covered the soil. All the turtles were alive the
following spring except for one juvenile, found frozen at a depth of
one inch on December 30 (the lowest air temperature up to this
time was approximately -12°). Three adult and 24 juvenal T.
ornata hibernating in the earth of an outdoor cage at the University
of Kansas in the winter of 1955-56, were all dead on December 3
after air temperatures had reached a low of -12 degrees.

Ornate box turtles are usually solitary when hibernating; in the
rare instances in which more than one turtle is found in the same
hibemaculum, the association has no social significance and is sim-
ply a reflection of the availability and suitability of the hibemacu-
lum. The only communal hibemaculum — the "Tree Den" — at the
Damm Farm was discovered on October 16, 1955, after a turtle was
traced to it by means of a trailing thread. The flask-shaped cavity,
approximately two and one-half feet deep, in the north-facing bank
of a narrow ravine, had an entrance one foot wide and nine inches
high, nearly flush with the bottom of the ravine. Grasses on the
bank of the ravine hung over the entrance and nearly concealed it.
The steep sides of the ravine protected the entrance from wind.

Seven turtles were in the den when it was discovered, and on
each of five subsequent visits from October 20, 1955, to March 6,
1956, fewer turtles were found in the den. Figure 24 shows the
approximate length of stay of each known occupant of the den.
Only one of the turtles ( an adult female ) that left the den returned.
Turtles found in the den on three visits in October were more or
less torpid and were seen easily from the entrance but on Novem-
ber 6 the two remaining individuals had burrowed into the sides
and floor of the den.

Three turtles (one female, one male, and one juvenile) were
found in separate form-hibemacula within a few inches of one

Natural History of the Ornate Box Turtle

615

another on November 6, 1955 (PI. 21, Fig. 2). The common en-
trance to all three hibernacula was a shallow depression that resulted
from an old post-hole. Soil in the depression was loose and moist
and ideal for burrowing. The three hibernating turtles were situ-
ated, in a vertical plane, at depths of 18 ( ^J ), 12 (juvenile), and
seven ( 5 ) inches. One of the turtles hibernating at this place on
November 6 was basking on October 30 in the shelter of some tall
weeds a few feet from the hibernaculum.

16 Oct 20 Oct 30 Oct 6 Nov. 26 Dec 6 March 15 May

A 17.5

A 19.0

A 103

A 10.6

A 6.4

A 73

B 18.2

B 17.6

B 14.2

B 127

B 5.4

B 83

Fig. 24. The approximate length of stay of each known occupant
of a den that was examined six times in the winter of 1955-1956 at
the Damm Farm. Most of the occupants used the den as a temporary
shelter and sought permanent hibernacula elsewhere. One turtle
left the den for approximately two weeks and then returned to it for
the rest of the winter. The temperature of the air outside the den
(A) and the average body temperature of turtles in the den (B) are
given at the bottom of the diagram for each date the den was ex-
amined. The symbol "J" represents a juvenal turtle.

In general, body temperatures approximated the temperature of
the soil around the turtle. Body temperatmres tended to be slightly
higher than soil temperatures in November and December but were
slightly lower than soil temperatures in the months of February and
March. The lowest body temperature recorded for any turtle that

616 University of Kansas Publs., Mus. Nat. Hist.

survived a winter was 2.7 degrees, taken from an adult female on
December 26, 1955. Body temperatures one to three degrees
higher were common in the coldest part of the winter. Turtles in
shallow hibemacula, like those observed in wooded areas at the
Reservation, are probably subjected to freezing temperatures at
least for short periods but I have no records of body temperatures
this low, except where they were induced experimentally. Turtles
exposed to temperatures of zero degrees or sHghtly lower would
retain enough heat to survive without freezing for a period of
several hours or even a day if well insulated. A temperature
gradient exists within the body; cloacal temperatures, for example,
differ from temperatures deep in the colon and temperatures in the
dorsal and ventral parts of the body cavity (taken by manipulating
the bulb of the thermometer while it was in the colon) differ from
one another. Probably, therefore, some parts of some turtles —
probably the top of the shell or the extremities— freeze in winter
without causing the death of the turtle. Ewing (1939:91) found a
female of T. Carolina, just emerging from hibernation, that had lost
some scutes from its carapace; he found the missing scutes in the
hibernaculum and attributed their loss to severe temperatures in
the winter of 1933-34.

The incidence of mortality due to freezing is unknown for most
species of reptiles. The observations of Bailey (1948) on DeKay
snakes (Storeria dekatji) and Legler and Fitch (1957) on collared
lizards suggest that rates of mortality are high in dormant reptiles.
Bailey {op. cit.) suggested that winter mortality might act as a
natural check on snake populations. Neill (1948a:114) thought
more box turtles (T. Carolina) were killed in Georgia by cold
weather in late autumn than ". . . by all other factors together,"
and that this winter mortality acted as an effective check on popula-
tion levels. Neill reported that many turtles left their burrows in
late autumn and began to forage; if the temperature dropped
suddenly, the turtles became ". . . too torpid to dig" and froze.

If ornate box turtles are occasionally caught in the open by a
sudden cooling of air temperature, it would occur at a time of year
when temperatures would approximate freezing but would drop
not far below this level; laboratory and field records show that
adults could probably survive these low temperatures overnight
and warm up sufficiently on the following day to seek adequate
shelter. Box turtles deepening their burrows in winter do so at
body temperatures somewhat lower than 10 degrees (near the
minimum temperature at which co-ordinated activity was observed

Natural History of the Ornate Box Turtle 617

in the laboratory); turtles found in the open in late October were
known to biurow into the ground at body temperatures of approxi-
mately 15 degrees.

Emergence from hibernation usually occurs in April but in some
years a few turtles may emerge as early as the first week of March.
Emergence is stimulated by temperature and humidity. Fitch
(1956b: 438) stated that emergence was delayed until ". . .
the ground has been sufficiently moistened and until air tempera-
tures have reached at least 26°." Box turtles at the Reservation
emerged on April 21 in 1954 and from April 16 to 17 in 1955.
William R. Brecheisen found recently emerged box turtles in
Anderson County on April 2, 1955, and March 6, 1956.

Turtles were found facing upward in their hibemacula in early
March. As the temperature of the soil rises, they move slowly
upward, usually following the route by which they entered. They
remain just below the surface of the soil for a week or two before
actually emerging; tliis final phase of emergence is probably has-
tened by spring rains that soften the soil. Activity may be sporadic
after emergence if the weather is cold.

A number of box turtles at the Reservation emerged in a cold
rain in 1954 when the temperatures of the air and ground were
16 and 13 degrees, respectively, but remained inactive for several
days afterward. In 1955 the air and ground temperatures were
higher (28° and 17°, respectively) on the day of emergence and
box turtles became active almost immediately.

DIET

Published information on the food of T. ornata consists of a
few miscellaneous observations. Cahn (1937:103) opened five
stomachs that contained partly digested vegetable matter but no
insects or other animal food: Ortenburger and Freeman (1930:
187) noted that grasshoppers were a main part of the diet of T.
ornata in Oklahoma and that turtles displayed unsuspected agility
in catching them. Those authors also saw turtles eating caterpillars
and robber flies. Strecker (1908:79) stated that "The natural diet
of this species consists of vegetable matter and earthworms." Nor-
ris and Zweifel (1950:3) observed the feeding habits of captive
T. 0. luteoJa. Coyote melon (Cuctirbita foetidissima) was eaten
viath reluctance but a collared lizard (Crotaphytm collaris) was
quickly devoured. Tadpoles of Scaphiopus hammondi were caught
in a small pool and eaten. Adults of the same species were re-
jected after being caught; box turtles were seen wiping their mouths

618 University of Kansas Publs., Mus. Nat. Hist.

after rejecting adult toads. The authors suggested that T. o. luteola
is an important predator of Scaphiopus hammondi, since the two
species occur together in many areas and the emergence of both
is controlled to a large extent by rainfall. One individual of
luteola was seen eating a dead box turtle on a road.

Captive individuals of T. ornata, observed in the present study,
ate nearly every kind of animal and vegetable food given to them.
Table scraps, consisting chiefly of greens, various fruits and vege-
tables, meat, and cooked potatoes, formed the main diet of turtles
kept in outdoor cages.

A number of persons have told me of ornate box turtles eating
the succulent stems and leaves, and the fruits of various garden
plants; similar incidents probably occur in areas of native vegeta-
tion. J. Knox Jones told me he saw an individual of T. ornata
eating a spiderwort ( Tradescantia sp. ) in Cherry County, Nebraska.

Sight-records of foods eaten by box turtles at the Damm Farm
(excluding the many records of individuals foraging in dung or
eating mulberries) were for grasshoppers, caterpillars, and various
kinds of carrion. Box turtles were often seen eating grasshoppers
on roads in early morning; Sophia Damm told me of frequently
seeing individuals catching grasshoppers in her garden. Ralph J.
Donahue told me that on his farm in Bates County, Missouri, an
individual of T. ornata made a circuit of the lawn each morning in
summer and ate all the cicadas (Magicicada septendecim) found.

Vertebrate remains found in the stomachs of box turtles seem
to result chiefly from the ingestion of carrion. One box turtle ate
a white egg ( unidentified ) that had fallen from a nest and another
was seen with a blue down feather clinging to its mouth. Several
colleagues have told me of box turtles eating small mammals caught
in snap-traps and Marr (1944:489) reported a similar incident.
J. Knox Jones told me he once found an ornate box turtle in the
nest of a blue-winged teal in Cherry County, Nebraska; the three
eggs in the nest had been broken. The only authentic record of an
ornate box tiirtle preying on a vertebrate under natural conditions
was one supplied by Ralph J. Donahue who saw an adult catch and
eat one of a brood of bobwhite quail. In many areas where box
turtles are abundant, it is the opinion of local residents that the
turtles decimate populations of upland game birds by eating the
eggs and young of these birds; these opinions result probably from
rare encounters such as the one described by Donahue. I believe
that box turtles at the Damm Farm were sometimes able to catch
young frogs and tadpoles (chiefly Rana catesbeiana and R. pipiens)

NATxmAX, History of the Ornate Box Turtle 619

at the margins of ponds. In autumn literally thousands of young
Rana were present in these places.

Ornate box turtles ordinarily attempt to catch and, without
further examination, to eat, small objects moving on the ground, but
are more critical of stationary objects. Captive turtles, for example,
would immediately chase and seize a grape that was pulled or
rolled slowly across a floor but a stationary grape was examined
and then smelled before it was eaten. Similar observations were
made a number of times with hving and dead insects in the field
and in the laboratory. A turtle discovering an object that is of
possible value as food, approaches it closely, turns the head from
side to side (presumably using the eyes alternately to examine the
object), and then, with head cocked at a slight angle, momentarily
presses the nostrils against the object (PI. 28, Fig. 4). If acceptable
as food, the object is then swallowed whole or taken into the mouth
with a series of bites; large insects are usually broken into several
pieces in the process of being bitten and swallowed. Larger ob-
jects, such as dead vertebrates, are torn to pieces with the beak
and forefeet before they are swallowed. Hatchlings, when fed for
the first time, ignored inanimate foods but eagerly chased meal-
worms, catching them usually by the anterior end. The tendency
of the young of certain species of turtles (especially captives) to
be more carnivorous than adults is probably due to the association
of movement with food; recognition of inanimate objects as food is
presumably learned by older individuals.

Mulberries (Morns rubra), when they are abundant, constitute
all or an important part of the diet of ornate box turtles. On June
4, 1955, William R. Brecheisen and I drove along a road in Ander-
son County, Kansas, and stopped at each mulberry tree that we
saw beside the road; we found at least one specimen of T. ornata
under nearly every tree. Approximately twenty box turtles were
collected in this manner in a little more than one hour. The heads
and necks of most were stained dark-red from the fruit and, in
some, nearly the entire shell was stained. Dissection of these
turtles revealed that their stomachs were distended to two or three
times normal size with mulberries; no other kinds of food were
found in the stomachs. Some of the turtles voided purplish-black
fluid from the cloaca when we handled them; the color of the
fluid presumably resulted from mulberries.

Several turtles were observed through binoculars as they foraged.
Individuals snapped or lunged periodically at objects on the ground
along the route of travel. Upon reaching an area where cow dung

620 UxrvERSiTi' of Kansas Publs., Mus. Nat. Hist.

was abundant, a turtle would move directly to a pile of dung and
begin tearing it apart with the forelegs or burrowing into it.
Turtles most often foraged in cow dung that had a superficial,
dried crust. The invertebrate fauna of older dung was probably
greater than that of fresh dung. Adult and larval insects were
eaten, along with quantities of dung, as they were uncovered.
Sometimes box turtles chased and caught larger insects that ran
a foot or more away from the pile of dung; the turtles could cover
the distance of one foot with three or four quick steps. Depres-
sions made by box turtles in cow dung, as well as drier cow dung
that had been more completely dissected, were regarded as charac-
teristic "sign" of T. ornata at the Damm Farm and in other areas
studied (Pi. 26). Several persons have told me of box turtles
"eating cow dung"; these reports, most of them made by competent
observers, probably result from observations of box turtles ingest-
ing cow dung incidentally, along with some unseen item of food.

Contents of stomachs were analyzed. Scats and contents of
lower digestive tracts, although obtained in large quantity, were
unsuitable for analysis because of the fragmentary nature of the
foods they contained. Relative amounts of various kinds of foods
in stomachs were estimated; volume was determined by displace-
ment of water or fine shot.

Twenty-three stomachs of adults were selected at random (ex-
cept for the fact that empty stomachs were discarded) from more
than a hundred specimens collected in Douglas County, Kansas,
in the period from June, 1954, to June, 1957; the sample included
stomachs obtained in nearly all the months of the season of activity.
Kinds of foods in stomachs did not differ significantly in regard to
the sex of the turtles or to time of year. The stomach of each of
two juveniles (included in Table 6) contained a greater variety of
animal food than did the stomach of any adult, but no kind of
animal was eaten by the juveniles exclusively.

Each of the 23 stomachs contained animal matter and, in addi-
tion, all but two contained at least some plant material from dung,
which constituted up to 20 per cent of total stomach contents.

Insects were present in each of the 23 stomachs and constituted
the bulk of the animal matter; beetles, caterpillars, and grass-
hoppers (ranked in descending order) were the kinds occurring
most frequently and constituting the largest average percentages
of total stomach-contents. Most of the beetles were scarabaeids
and carabids; the bulk of the caterpillars were noctuids and arctiids.
Grasshoppers, with one exception, were of a single species, Mel-

Natural History of the Ornate Box Turtle

621

Table 6. — Kinds of Animals Found in the Stomachs of 25 Terrapene o.

omata of Both Sexes (23 adults, 2 juveniles) from Douglas County,

Kansas. Frequency of Occurrence (number of stomachs in which

foiwd) is Given for Each Item Listed.

Frequency of Occurrence

Adults

Larvae

Total

Gastropoda

Helisoma sp

Succinia sp

Polygyra sp

Retinella sp

Crustacea

Procambaris gracilis. .
Armadillidium vulgar e

1

4

Orthoptera (Locustidae)

Locustinae {Melanoplus differentialis) .
Oedipodinae

13
1

Lepidoptera (unspecified) ,

Arctiidae

Noctuidae

Pyralidae

Sphingidae

Diptera (Sarcophagidae)

Coleoptera (unspecified)

Cantharidae

Carabidae (unspecified)

(Eumolops colossus)

Cerambycidae {Prionus fissicomis)

Chrysomelidae (Diabotrica 12-punctata).

Curculionidae {Calendra parvulus)

Lampyridae {Photinus pyralis)

(Photuris sp.)

Phengodidae

Scarabaeidae

Hymenoptera (Formicidae) .

6
1
1
1
3
2

11

2

1
9
10
1
1

1
4

13
1

1
9
10
1
1

1

3

1
6
1
1
1
3
2
1
1
11

Phalangida

Araneida (Epeira) .
Diplopoda

Vertebrata (carrion) ,

622

University of Kansas Publs., Mus. Nat. Hist.

Table 7. — Occxjrrence of Insects, by Frequency and Volume, in Stomachs

OF 23 Terrapene ornata from Douglas County, Kansas. Relative Volume

IS Based on Total Amount of Food Material Present, Excluding Stones

AND Vegetable Material Contained in Dung.

Insects
(all)

Orthoptera

Lepidoptera
(larvae)

Coleoptera

Average

volumetric percentage. . .

88.6

28.7

26.9

32.5

Range

(volumetric percentage) . .

trace
to 100

0 to 100

0 to 100

Oto 100

Frequency of occurrence
(percentage of total
stomachs in which found)

100

52

65

74

anoplus differentialis. It is noteworthy that two of the kinds of
insects frequently eaten ( diflFerential grasshoppers and noctuid
caterpillars) are of economic importance in that tliey damage
crops.

Snails, sowbugs, and the one individual of crayfish found in
stomachs were kinds that could be expected to occur in moist
grassland or in wooded stream courses. Mulberries were present
in one stomach and fragments of bird's-nest fungi (Cyathtis stria-
tus) were present in another. Carrion consisted of remains of mam-
mals and birds; the only identifiable items were bones of the eastern
cottontail (Sylvilagus floridanus) and a chicken. Stones up to
seven millimeters in diameter were found in many stomachs; stones
constituted as much as half of total stomach-contents. Presumably
the stones were accidentally swallowed when food was taken from
the ground.

The few adequate reports on dietary habits of T. Carolina (Al-
lard, 1935:325-326; Carr, 1952:147, 150, 152, 153; Stickel, 1950:361;
Surface, 1908:175-177) indicate that the species is omnivorous but
that individuals tend to be herbivorous or carnivorous at certain
times. Ornate bo.x turtles resemble T. Carolina in being opportun-
istic feeders but rely on insects as a staple part of the diet. In this
respect the ornate box turtle seems to differ from all other kinds
of box turtles in the United States and it is probably unique in its
habitual utilization of dung communities as a source of food.

Natural History of the Ornate Box Turtle 623

POPULATIONS

Ornate box turdes were probably more numerous on the Damm
Farm than any other kinds of reptiles, excepting skinks {Eumeces
fasciatus and E. obsoletus), and were by far the most conspicuous
element of the reptilian fauna.

The 194 box turtles that were marked at the Damm Farm were
captured a total of 437 times. Seventy-nine (41 per cent) individ-
uals were recaptured at least once, 49 (25 per cent) twice, 29 (15
per cent) three times, and 20 (10 per cent) were recaptured at least
four times. Only three individuals were recaptured more than eight
times. The greatest number of recaptures for a single individual,
an old female, was 23.

In all, 185 turtles (95 per cent of total recorded at Damm Farm)
were captured on the pasture. Of these, 73 were in the northwest
comer area, 44 in the house pond area, and 35 in the southern ra-
vine area. The density of the population at the Damm Farm, con-
sidering the entire area, was .88 turtles per acre; for the woodland
area alone, density was .41 turtles per acre and for the pasture alone,
density was 1.49. Acreage and population density in the northwest
comer, house pond, and southern ravine areas were respectively, 28
acres with 2.6 turtles per acre, 7 acres with 6.3 turtles per acre, and,
17 acres with 2.6 turtles per acre. The densities noted above for the
wooded area and for the entire Damm Farm are low as a result of
incomplete sampling in the wooded area. Estimates of population
density for the subdivisions of the pasture seem more closely to
approach the true population density in areas of favorable habitat.

Fewer unmarked turtles were captured as the study progressed,
but they were still being captured occasionally when field work was
terminated. In order to estimate the number of turtles in the pop-
ulation at the Damm Farm the "Lincoln Index" (Lincoln, 1930)
was used to compare the ratio of marked individuals to total num-
ber of individuals (17:56) in collections for June, 1956, to the ratio
of marked individuals as of July 31, 1955 (87) to total individuals
in the population; the result was 286.

Fitch (1958:78) estimated the population of T. ornata in one area
of the Reservation (including woodland and ungrazed pasture) to
be .076 turtles per acre. Stickel (1950:373) estimated the popula-
tion of adult T. Carolina to be four to five turtles per acre in favor-
able habitat at the Patuxent Research Refuge, Laurel, Maryland;
juveniles comprised less than ten per cent of the population.

Of the 194 turtles marked at the Damm Farm, 103 (53 per cent)
were adult or subadult females, 61 (31 per cent) were mature
7—773

624 University of Kansas Publs., Mus. Nat. Hist.

males, and 30 (16 per cent) were juveniles of undetermined sex.
The ratio of males to females was then, 1.00 to 1.69, and the ratio
of juveniles to adults was, 1.00 to 6.47. Eighteen of the 194 indi-
viduals were juveniles less than 90 millimeters in plastral length
and only six had plastra less than 60 millimeters long (Fig. 25).
The unbalanced ratio between males and females may result, in
part, from sexual difiFerences in habits. The studies of Carr (1952:
9), Fitch (1954:140), Forbes (1940:132), Legler (1954:138), and
Risley (1933:690), have shown, however, that unbalanced sex ra-
tios, with females outnumbering males, are found in several spe-
cies of reptiles, especially in turtles.

Records for 540 adult T. ornata collected at the Damm Farm, the
Reservation, and on roads in eastern Kansas, show that females out-
number males just before and during the nesting season and again
in late autumn (Fig. 26). The high incidence of females in May,
June, and July, can be explained by their more extensive movements
associated with nesting in these montlis. I have no explanation for
the increased number of females captured in late autumn. In April
and August, the only two months in which males were more abun-
dant than females, the samples were small. The number of juveniles
collected was too small to allow any trustworthy conclusions con-
cerning their seasonal incidence; a few juveniles were taken in
nearly all the periods in which adults were active.

Risley (1933:690), studying Sternotherus odoratus in Michigan,
found an over-all sex ratio of 1.0 male to 2.3 females; the percentage
of females in collections ranged from 50 to 71 per cent in April and
most of May and rose to 83 and 85 per cent in late May and mid-
June, respectively.

The infrequency with which hatchlings and small juveniles of
ornate box turtles are observed is well known to naturalists. Several
of my colleagues who are expert field observers and who have lived
in areas where ornate box turtles are abundant, have never seen
hatchlings; many other persons have seen only one or two. Rodeck
(1949:33), noting the abundance of coleopterous insects in the scats
of captives and the rarity of individuals of all age groups during dry
periods in Colorado, commented, "It is possible that the young are
even more subterranean tlian the adults. Perhaps they spend their
early years in rodent or other burrows where there is a fairly abun-
dant insect fauna. Increasing size might force them to the surface
for feeding, with a daily return to a burrow for resting and pro-
tection."

Natural History of the Ornate Box Turtle

625

My own experience in the field has shown that small examples of
T. ornata are not so rare as previous workers have beUeved. Small
box turtles occupy the same microhabitat as do the adults and seem
not to be more aquatic or subterranean in habits. Juveniles are
found in burrows, in marshy areas, and in other sheltered places, but
so are adults. Most of the juveniles that I found were in open
situations where adults were abundant, sometimes within several
inches of a place where an adult was feeding or basking. Nearly
every one of the smaller turtles was discovered when I was closely
scrutinizing some other object on the ground; sometimes juveniles
were actually touched before being seen. Most juveniles were
covered with cow dung or mud and blended so well with the sub-

50

_

1

40

-

v>

1 §

■

o

•>

^^H

= 30

-

ff =6

^H

:§

5 .£

^H

■o

o

^^^^1

c

E £

3

H

"5 20

-

o» i

H

w

•^ e

^^^

0)

•g —

^^^^M

f

1 *

■ ■

i 10

ri

'^

+

II

0k

O O) 0) o>

' ■ ^^m ^^m ^ i^MB~

o> o> 100-109 110-119 120-129 130-139

lO

* in «o h-

oo o>

1 • I 1

• 1

o

o o o o
^ in u> to

o o length of plastron (mn^)

Fig. 25. Composition of the population of T. o. ornata at the Damm
Farm based on the 194 individuals marked there in the years 1954 to
1956. Individuals smaller than 100 mm. ordinarily could not be sexed
accurately and are shown as open bars. Open bars in the groups larger
than 100 mm. are for females, whereas solid bars are for males.

strate that they were detected only when they moved. It is likely
that only a small number of the young box turtles present in an area
is ever actually observed. Young are more vulnerable to predation
and injury because of their small size, soft shells, and immovable
plastra. They evidently rely, to a large extent, on inconspicuousness
for protection.

626

Unts'ersity of Kansas Publs., Mus. Nat. Hist.

Fig. 26. The seasonal abundance of females of T. o. omata based
on 540 adults captured at the Damm Farm, the Reservation, and
on roads in eastern Kansas, in the years 1954 to 1956. Records
are grouped in periods of 30 days, the periods beginning with the
dates shown at the bottoms of the bars. Juveniles are not con-
sidered. Numbers at the top of each bar indicate the size of the
sample (both sexes) and give an approximate indication of relative
seasonal abundance of adults, except for August, when little field

work was done.

MOVEMENTS

The only previous study of movements of T. omata is that of
Fitch (1958:99-101). He recovered 14 marked T. omata at the
Reservation a total of 30 times, the period between recaptures vary-
ing from one to seven years. He reported that the average radius
of home range was 274 feet (for an area of approximately 5.4
acres), excluding a single (presumably gravid) female that moved
1830 feet in 53 days.

Although published information on T. omata is scant, a consid-
erable amount of information is available concerning its congener,
T. Carolina. The classic studies of Stickel (1950) on it constitute
the most complete account of populations and movements for any
reptile or amphibian, and probably, for any vertebrate. She found
the average home range of adults to be 350 feet in diameter. Home
ranges were not defended as territories and nearly all individuals
were socially tolerant of one another. Movements (studied by
means of a thread-trailing device) were characterized by frequent

Natural History of the Ornate Box Turtle 627

travel over the same routes within the home range. Some turtles
concentrated their activities in only one part of the home range,
moving subsequently to another part, and some turtles had two
ranges between which they traveled at varying intervals. Females
ordinarily left their home ranges to nest.

Other noteworthy, but less detailed, studies of populations of
T. Carolina are those of Breder ( 1927 ) who found evidence of home
range and homing behavior, and of Nichols (1939b) who, after
observing a marked population on Long Island over a period of
twenty years, found evidence of homing behavior and estimated
normal home range to be approximately 250 yards in diameter.
Numerous shorter papers such as those of Schneck (1886) and
Medsger ( 1919 ) document the tendency of T. Carolina to remain in
restricted areas over long periods.

Important studies that indicate the presence of home range and
homing behavior in other chelonians are those of Cagle (1944) on
Pseudemys scripta and Chrysemys picta, and of Woodbury and
Hardy (1948) on Gopherus agassizi. Grant (1936) and Bogert
(1937) have also indicated that movements of individuals of Go-
pherus agassizi are restricted to limited areas.

Locomotion

Ornate box turtles moving forward over even terrain hold the
plastron a quarter to a half inch above the ground and keep the
head and neck lowered and extended. Each foreleg is brought
forward and the humerus points nearly straight ahead when the
foot touches the ground. Nearly all of the palmar surface is ini-
tially in contact with the ground but as the body is brought forward
and the humerus swings outward, only the claws, and finally, only
the two inner claws are in contact with the ground. Of the hind
feet, the medial surfaces are the principal parts that touch the
ground but some traction is derived from the hind claws at the be-
ginning of each cycle of the hind leg. Under normal conditions,
box turtles move slowly and pause to rest and examine their sur-
roundings every few feet. When resting, the plastron is in contact
with the ground, the legs relaxed, and the head and neck are ex-
tended upward. Some turtles seeking shelter from the heat of sun-
shine walk rapidly for a hundred feet or more without pausing.

Turtles seen feeding under natural conditions displayed remark-
able agility in making lunges, consisting of one or two short steps
and a thrust of the head, at moving objects. Turtles kept in my

628 University of Kansas Publs., Mus. Nat. Hist.

home were able, after being conditioned to hand-feeding, quickly
to intercept a grape rolled slowly across a linoleum-covered floor.

Frederick R. Gehlbach told me that, of several species of captive
turtles observed by him, T. ornata characteristically walked with
the plastron held well above the substrate, as did Gapherus herlan-
dieri, but that T. Carolina (specimens from the northeastern U. S.)
dragged their shells as they walked. Apparently T. Carolina in
Kansas (currently referred to the subspecies triunguis) differs
somewhat in gait from populations in the eastern part of the range;
several individuals of T. Carolina from Kansas that I observed in
captivity, kept their plastra raised well above the smooth, hard
substrate over which they walked.

Box turtles at the Damm Farm were able easily to climb ravine
banks that sloped at an angle of 45 degrees and, with some diffi-
culty, could climb banks as steep as 65 degrees. Most individuals,
however, were reluctant to walk directly downward on banks as
steep as 45 degrees. Several individuals were seen to lose footing
when climbing up or down a steep bank and to roll or slide to the
bottom. Ordinarily, T. ornata is able to climb over a sheer surface
as high as its shell is long, provided the surface is rough enough to
give some traction to tlie foreclaws. The claws of first one, then
the other forefoot are placed over the top of the barrier and then
a hind foot, extended as far forward as possible, secures a hold as
the turtle goes over the barrier.

A number of observations on speed were made in the field
where distance traveled and time elapsed were known approxi-
mately. Speeds ranged from 20 to 100 feet per hour in the course
of foraging. Higher speeds (400 or more feet in one hour) were
for turtles moving along pathways or seeking shelter. Gould
(1957:346) observed somewhat faster speeds in T. Carolina (192
feet per hour in cloudy weather and 348 feet per hour in sunny
weather); he observed individuals that had been removed from
their normal home ranges.

Individuals of T. ornata that were placed in water swam moder-
ately well but were clumsy in comparison to individuals of more
aquatic emyids such as Pseudemys and Chrtjsemys. Box turtles
were never observed to swim voluntarily, although they were fre-
quently found in shallow water. On several occasions I confronted
individuals at the edge of a pond so that the only unblocked route
for their escape was through deeper water; nearly always these
individuals attempted to crawl past me, to crawl away in shallow

Natural History of the Ornate Box Turtle 629

water parallel to the shore, or to hide in soft mud at the edge of
the water. Box turtles floated high in the water with the dorsal
side upward and had httle difficulty in righting themselves when
turned over. The head and neck are extended and submerged
when the turtle is swimming; forward progress is interrupted every
few moments to elevate the head, presumably for purposes of
breathing and orientation. The shell is never submerged. The
swimming of T. ornata is in general like that of Pseudemys or
Chrysemys that have become dehydrated after long periods out
of water and cannot submerge. These more aquatic turtles, how-
ever, quickly overcome their bouyancy, whereas examples of T.
ornata, even if left in water for several days, are unable to sub-
merge. Clarke (1950) saw an ornate box turtle swim a 60-foot-
wide stream in Osage County, Kansas; his description of swimming
agrees with that given above.

The meager swimming abihty of T. ornata is of apparent sur-
vival value under unusual conditions and enables T. ornata to
traverse bodies of water that would act as geographic barriers to
completely terrestrial reptiles; however, swimming is a mode of
locomotion seldom used under ordinary circumstances.

Gehlbach (1956:366) and Norris and Zweifel (1950:2) observed
individuals of T. o. luteola swimming in temporary rain pools and
small ponds in New Mexico; the two authors last named saw an
individual quickly enter a pond and dive beneath the water after
being startled on the bank. Several of my colleagues, in conver-
sation, have also reported seeing T. o. luteola in small bodies of
water in the southwestern United States.

Daily Cycle of Activity

The daily cycle of T. ornata consists basically of periods of bask-
ing, foraging, and rest that vary in length depending upon en-
vironmental conditions. Turtles emerge from burrows, forms, and
other places of concealment soon after dawn and ordinarily bask
for at least a few minutes before beginning to forage; foraging is
combined sometimes with basking, especially in open areas that
are suitable for both kinds of activity. Foraging usually continues
until shelter is sought sometime between mid-morning and noon.
Turtles remain under cover (or continue to forage in shaded
areas) until mid-afternoon or late afternoon when they again be-
come active. They forage in both morning and afternoon. Study
of travel records of a few of the turtles equipped with trailers

630 Untv'ersity of Kansas Publs., Mus. Nat. Hist.

suggests that, under normal conditions, activity is slightly greater
in forenoon tlian in afternoon, but that the converse is true of
gravid females seeking nesting sites. Strecker (1908:79) reported
that captive T. omata, after developing a feeding reflex, ate and
retired until feeding time next day.

As environmental temperatures rise in summer, the period of
mid-day quiescence is lengthened. In the hottest part of the year,
some turtles remain under cover for several days at a time. In
periods of clear, cool weather at the beginning and end of the
growing season, some turtles remain abroad and bask for most of
the day.

Examination of thread trails showed that activity of all individ-
uals except nesting females was terminated at dusk. Breder
(1927:236), Allard (1935:336), and Stickel (1950:358) reported a
corresponding lack of nocturnal activity in T. Carolina. Terrapene
o. ornata in Kansas, and T. o. luteola in New Mexico (Norris and
Zweifel, 1950:2) — unlike desert tortoises, Gopherus agassizi, which
are active at night in hot weather (Woodbury and Hardy, 1948:
186) — do not utihze the hours of darkness for foraging, even in the
hottest part of the year.

Seasonal Cycle of Activity

Data obtained by mapping the movements of turtles that were
equipped with trailing devices made it possible to compare dis-
tances traveled in the course of daily activities at different times of
the year. Some of these data are expressed graphically in Figure
27. It should be noted that movement at all times in the season
of activity was uneven; that is to say, an individual would move
several hundred feet each day for a period of several days, and
then, for an interval of one to several days, move only a few feet
from one shelter to another, or not move at all. Such periods of
rest could not be correlated definitely with environmental condi-
tions; some individuals were inactive on days that were probably
ideal (in terms of moderately warm temperatures and high hu-
midity) for activity of box turtles. Analagous rest periods were
noted in T. Carolina by Stickel ( 1950:358).

Two males of T. ornata that had been removed by me from their
normal home ranges traveled the longest average distance per
day (429 feet). Gravid females in June traveled the next longest
average distance per day (363 feet). The average distances trav-
eled per day by non-gravid females in June (226 feet) and July

Natural History of the Ornate Box Turtle

631

(260 feet) and by males (within their known home ranges) in June
(289 feet) were thought to approximate normal amount of move-
ment under average environmental conditions. Average distance
traveled per day by females in October (152 feet) was shortest
because of frequent and extended rest periods. Nevertheless, in
October actual distances traveled on days of activity tended to be
longer than in any other month. A gravid female traveled farther
in a single day than any otlier individual of T. ornata observed;

^1000

a>

>« 800

a>
o.

600

■o

>
o

a, 400
u

c
o

(A

200

O-

Fig. 27. Average distances traveled per day by males and fe-
males at different times of tlie year, determinea by mapping of
thread trails at the Damm Farm. The diagram for "homing males"
represents tlae distances traveled by two males removed from their
normal home ranges to test homing abihty. The data presented
are for an aggregate of 136 days of trailing. Vertical and hori-
zontal lines represent, respectively, the range and mean. Open
and solid rectangles represent one standard deviation and two
standard errors of the mean, respectively.

she moved along a rock fence for approximately 700 feet, then
left the study area and moved, in a nearly straight line, 1,200 feet
across a cultivated field. Then the thread on her trailer was ex-
pended. The total distance moved, therefore, was at least 1,900
feet and probably more.

An adult male at the Reservation traveled 2,240 feet in the 36-day
period from October 16 to November 20, 1954, mostly on a wooded

632 University of Kansas Publs., Mus. Nat. Hist.

hillside. Eleven forms found along the route of the turtle's travels
indicated that movement took place on roughly one out of three
days in the elapsed period and demonstrated the sporadic nature
of movements in autumn. The turtle remained active for an un-
determined time after November 20.

Home Range

Data obtained from trailing and various methods of recapture
at the Damm Farm indicated that each individual used only a small
part of the total study area in the course of daily activities and
tended to remain within a restricted area for a long time.

The number of recaptures of no individual was great enough to
permit application of refined calculations of size of home range as
described by Odum and Kuenzler (1955). For individuals that
were recaptured six or more times, or individuals for wliich ade-
quate traihng records were available, the area enclosed by a Hne
joining the peripheral points of capture was considered adequately
representative of the home range of that individual, unless re-
captures were all within a few feet of each other or lay in an ap-
proximately straight hne. If less than six records of recapture were
available, home range was estimated, in the manner described by
Fitch (1958:73), by averaging the distance between successive
points of recapture and letting this average represent the radius of
home range; the actual area of home range was determined by the
formula, 7r(R)^, for the area of a circle.

Size of home ranges of males and females did not differ signi-
ficantly and data for the two sexes were combined in tlie final
analysis. The average radius of the home ranges of 44 adults (cap-
tured a total of 146 times) was 278 feet (extremes, 71 to 913) when
computed by measuring the distance between successive captures;
the average area of these home ranges was 5.6 acres. Data from
10 turtles that had been recaptured only once were combined with
data from 34 turtles tliat had been recaptured more than once when
it was found tliat the average size of home range in these two groups
did not differ significantly. Data concerning the home ranges of
eight of the 44 individuals were sufficient to permit actual measure-
ment of home ranges with a planimeter; home ranges of these eight
individuals had an average area of five acres (extremes, 1.2 to 10.2).

A minimum home range could theoretically consist of the smallest
area in which adequate food and shelter were available. Under
favorable conditions a turtle could stay in an area ten to twenty
feet in diameter. Although several such favorable small areas

Natural History of the Ornate Box Turtle 633

existed on the Damm Farm, box turtles seldom stayed in one for
more than a day or two. Seemingly, therefore, factors additional to
food and shelter influence size of home range. At the Damm Farm
these additional factors seemed to be: rock fences that acted as
physical barriers; areas that were cultivated, barren, or otherwise
unfavorable, acting as ecological barriers; and, cowpaths and ravines
that offered relatively unobstructed routes along which box turtles
tended to move.

One subdivision of the main pasture, the northwest comer area,
is an example of a relatively small natiural area in which many in-
dividual box turtles had home ranges. This tract of 28 acres was
roughly triangular and was bordered on two sides by rock fences
that contained no gates or other passageways. On its third (south-
eastern) side the area sloped into a deep ravine. Habitat in this
subdivision of the pasture (as well as in the other two subdivisions)
was especially favorable for box turtles because of peiTnanent water,
rocky slopes, ravines, and several fruit trees. Box turtles usually
foraged near the rock fences and the ravine (where dung was more
abundant than in other parts of the area), and tended, as they
foraged, to move parallel to these barriers. Turtles crossing the
area eventually came eidier to one of the fences or the ravine.
Therefore, most of the turtles in the northwest corner area
eventually completed a circuit of the area. Turtles that came to
the ravine tended to move along its bottom or sides. Several turtles
were known to cross the ravine and to forage in the grassy area on
its southeastern side. These turtles usually re-entered the ravine
by way of smaller side-ravines. Of 22 box turtles known to have
home ranges in the northwest corner area, only two individuals
(both gravid females) were known to leave the area in the period
in which observations were made.

Two other subdivisions of the main pasture — the house pond
area and the southern ravine area — although not so distinct as the
northwest corner area in terms of hmiting barriers, nevertheless
constituted separate areas of favorable habitat, each of which con-
tained a number of individual home ranges. Although the two
areas were not far apart, but Httle movement was observed of turtles
from one area to the other. The home range of only one turtle, an
adult female, was known to include parts of both areas.

Unbroken expanses of tall grass seem not to be optimum habitat.
The crest of tlie hill at the Damm Farm (PI. 17, Fig. 1) was an area
of more or less homogeneous grassy habitat. Turtles were seldom

634 University of Kansas Publs., Mus. Nat, Hist.

found on the crest of the hill although this area was as thoroughly
searched for turtles as any other area. Known home ranges of
nearly every individual observed were on either one of the sides of
the hill but not on both sides.

At several places on the border of the pasture, turtles were able
to move freely into cultivated areas but seldom did so except for
nesting. Trailing records show that most of the turtles that entered
one of the cultivated areas returned again to the pasture.

Ornate box turtles seem to find places of shelter by trial and error
along regularly used routes of travel in their home ranges. The
individuals that I studied never returned to the same forms, and
seldom returned to the same natural burrows and dens. Probably
foraging, basking, and watering sites are found also by trial and
error.

Stickel (1950:375) placed considerable importance on the occur-
rence of transient turtles in populations of T. Carolina; in estimating
population density, she added to her study area a peripheral strip,
half as wide as the average, estimated home range, to account for
turtles that had home ranges only partly within the study area. The
study area used by Stickel had no natural boundaries, as habitat
conditions on all sides were essentially the same as tliose of the
study area itself. The pasture at the Damm Farm, on the contrary,
is a relatively isolated area of natural grassland, bordered by rock
fences and cultivated fields. I believe that most of the box turtles
found on the pasture were permanent residents there. Individual
box turtles at the Damm Farm seemingly occupied but one home
range and it did not change from year to year. Populations of T.
ornata in areas less isolated than the Damm Farm, like the popula-
tions of T. Carolina studied by Stickel {loc. cit.), could be expected
to have a higher percentage of transient individuals and individuals
with multiple or changing home ranges. Henry S. Fitch told me
that he considered most of the individuals of T. ornata tliat were
captured only once at the Reservation were transients.

Several females at the Damm Farm traveled long distances from
their home ranges to nest but other females nested within their
known or estimated home ranges. Seemingly a complex of environ-
mental factors, including soil texture, weather, availability of water,
and possibly the urge for random wandering in the breeding season,
governs the distances traveled by gravid females and the ultimate
selection of a satisfactory nesting site. Females, because of tlieir
more extensive travels in the nesting season, seem more likely than

Natural History of the Ornate Box Turtle

635

males to have multiple or changing home ranges. Males of T. ornata
did not noticeably alter the extent or pattern of their movements in
the breeding season. Hibemacula, unlike nesting sites, were within
the known or estimated home ranges of all individuals studied.

Fig. 28. The movements of an adult (nongravid) female of T. o. ornata in
the house pond area at the Damm Farm during a period of 24 days in July,
1955 (solid line), and a period of three days (broken line) in July, 1956.
Solid dots represent the points where the turtle was found as her thread trail
was mapped; hollow symbols represent points of recapture when no trailing
thread was attached to the turtle.

The actual home range of almost every individual studied, even
of those individuals for which the most data were available, prob-
ably differed at least slightly from the observed or estimated home
range. One adult female, for example, was captured six times in

636 University of Kansas Publs., Mus. Nat. Hist.

10 Ju

100 feet
I I

scale

5 June

Fig. 29. The movements of a gravid female of T. o. omata in the southern
ravine area at the Damm Farm in a period of ten days in June, 1956. Her
movements were, for the most part, in and around several ravines (shown
on map by broken lines) where she was searching for a nesting site. For
explanation of symbols see legend for Fig. 28.

two years within a radius of approximately 50 feet. Another fe-
male was found 2780 feet from her last point of capture. These last
two records were regarded as unusual; when they were grouped
with records of the 44 individuals mentioned above, the average
radius of home range for the entire group was much larger (327
feet).

Homing Behavior

Gould (1957) reported that 22 of 43 T. Carolina moved in a home-
ward direction when they were released in open fields up to 5.8
miles from their original points of capture. Turtles oriented them-
selves by the sun; homeward headings were inaccurate or lacking
on overcast days and, light reflected from a mirror caused turtles
to alter their courses. Seven of ten turtles released more than 150
miles from home headed in directions that corresponded most nearly
to the headings last taken (at release-points near home base) and
did not necessarily correspond to the direction of home. Gould's
studies point out that box turtles perhaps practice a kind of "solar
navigation." His work raises the question of whether the move-
ments of box turtles are guided by the sighting of local landmarks
or whether such landmarks alter the course of movement only when
acting as barriers.

Natural History of the Ornate Box Turtle 637

In the present study two experiments were made to determine
the homing abihty of T. ornata. An adult male, taken from his nor-
mal home range in the house pond area and released 1200 feet
away in the southern ravine area, traveled a generally northward
course (not northeastward in the direction of home) for five days,
moving a distance of approximately 1900 feet. His detached trailer
was recovered several days later 740 feet southeast of the last
known point in his travels (a distance that could have been cov-
ered in two days ) and 150 feet from the point of original capture;
he had returned to his home range by a circuitous route in a period
of approximately seven days. Another adult male, captured in
the southern ravine area, and released in the house pond area
1900 feet away, traveled on a course that bore approximately 25
degrees north of true homeward direction; after five days he was
approximately 600 feet north of the original capture point. He
then began a northeastward course that took him back to the
house pond area where he remained for several days; no further
data are available for this individual. It is significant that the
homing males discussed above traveled greater average distances
per day (based on records for nine days of trailing) than any
of the other turtles studied (Fig. 27). Fitch (1958:101) released
an individual one half mile from where he captured it and, one
year later, recovered the tmtle near the point of release.

Social Relationships

Ornate box turtles are soUtary except during periods of mating.
Meetings with other individuals in the course of foraging, basking,
or seeking shelter, are fortuitous and have no social significance.
A broad overlapping of home ranges of both sexes at the Damm
Farm suggests that box turtles do not intimidate other individuals
in the home range or exclude them from it. No instances of fighting
were observed.

Allard (1935:336), Penn and Pottharst (1940:26), and Latham
(1917) recorded instances of fights between individuals of T. Caro-
lina; in the latter two instances fights were between males. Stickel
(1950:362) observed an incident between two males that may have
been a fight; however, she was of the opinion that fights rarely
occur in nature and that box turtles do not defend territories.
Evans (1954:23-25) considered the behavior of T. Carolina re-
ported by Penn and Pottharst ( loc. cit. ) to represent "territoriahty."
He found "... a true hierarchy . . ." existing between

638 University of Kansas Publs., Mus. Nat. Hist.

four captive males of T. Carolina and another between three captive
females of the same species; young individuals in the group raised
their social level in the hierarchy after receiving experimental
doses of male hormone. Evans {op. cit.:25) pointed out that true
tortoises (family Testudinidae ) have a more complex pattern of
social behavior than do emyid turtles.

Observations made with binoculars from the vantage point of a
blind provide the only information that I have concerning the
reactions of box turtles to one another under natural conditions.
Turtles foraging in a bare area were not startled by the approach
of other turtles, and turtles moving across the area seemed to take
no notice of turtles already there, regardless of whether these
tiutles were moving or not. Adults and subadults behaved in
approximately the same manner.

Individuals traveling or foraging in rough terrain or in grassy
areas probably are unable to see each other even when they are
close to one another. Conversely, box turtles can see each other
and are surely aware of each other's presence in bare, flat areas.
These facts suggest that no social hierarchy exists in T. omata. On
one occasion an adult male and a juvenile (hatched the previous
autumn) were found foraging next to one another on the same
pile of cow dung.

When an individual became motionless in an attitude of wariness
after having detected me in my bhnd, its behavior evoked no
response on the part of other turtles, a few feet away.

INJURIES

Fire, freezing, molestation by predators, and trampling by cattle
or native ungulates are only a few natural sources of injury to
which box turtles have always been exposed. Man's civilization
in the Great Plains, chiefly his automobile and other machines,
have compounded the total of environmental hazaids. Automo-
biles now constitute a major cause of death and serious injury to
box turtles. Each year thousands are struck on Kansas highways
alone, not to mention the many casualties resulting from mowing
machines, combines, and other farm machinery.

Although grass fires usually occur in early spring or late fall
when box turtles are underground, some turtles are surely killed
by fires and many are injured. In early April of 1955 the pasture
at the Damm Farm was burned. Similar burnings, I discovered,
had occurred both intentionally and accidently in past years at

Natural History of the Ornate Box Turtle 639

irregular intervals. No deaths or injuries, attributable to fire were
discovered in the course of intensive field work in the spring and
summer of 1955, when the new grass was short and conditions for
finding and marking box turtles were ideal. Badly burned indi-
viduals, if any, may have secreted themselves until their wounds
had healed. In June, 1957, an adult female, that had been burned
severely, was taken from a small puddle in a ravine on the Damm
Farm. The soft parts of her body, excepting her head and neck,
were a nearly solid mass of smooth scar tissue, the scales and
rugosities of the skin being practically obliterated. The tail was
reduced to a mere knob surrounding the anus and dead, exposed
bone was visible on most of the dorsal part of the carapace. Pos-
sibly this female was burned in the fire of 1955. Lack of injury
to the head and neck can probably be accounted for by the addi-
tional protection afforded these parts by the folded forelegs when
the turtle was withdrawn in the shell.

Turtles that are smashed flat on the highway, of course, have
no chance of survival. Highway fatalities are usually the result
either of "direct hits," where the tire of a vehicle passes directly
over the turtle, or of repeated pummeHng by subsequently passing
vehicles. The writer, while driving behind other cars that struck
turtles or by sitting beside roads, has observed numerous turtle
casualties. Most are struck a glancing blow by a tire and are pro-
pelled some distance through the air or on the surface of the pave-
ment, often to the side of the road. Such a blow is usually suffi-
cient to crack or chip the shell, or at least to scuff away parts of
the epidermal covering. Turtles, so injured, usually survive.

Parts of the shell do not break away easily, even when several
deep cracks are present, and only a little bleeding occurs. A
common injury inflicted on the highway is tlie wrenching and sub-
sequent dislocation of the caiapaco-plastral articulation. In such
instances the ligamentous tissue joining the two parts is torn ex-
tensively. Under these circumstances the movable shell parts seem
to act as a safety device, giving way under presssure that would
crack the shell of a turtle with rigid, fixed buttresses. Dislocations
of the carapaco-plastral articulation that have healed are charac-
terized by abnormally heavy development of ligamentous tissue
which may elaborate a horny, scutehke substance on its outer sur-
face.

The extent to which serious injury incapacitates a turtle is not
known. Surely open wounds are susceptible to infection and to

8—773

640 University of Kansas Publs., Mus. Nat. Hist.

various kinds of secondary injury; normal activity is probably in-
terrupted by a period of quiescence, at least in the period of initial
healing.

An injured female had a hole, slightly more than one inch in
diameter, in the right side of the carapace at the level of the second
lateral lamina. A tight, thin membrane stretched between the
broken edges of the opening; this membrane contained no bone
and was covered externally by scar tissue. It was obvious that this
turtle had recovered, at least in part, from a serious injury ( inflicted
probably by a piece of heavy farm machinery ) .

Minor chips, scratches, and abrasions on the shell result from
a variety of sources, some of them mentioned above. Small rounded
pits in the bony shell (shell pitting) due to causes other than
mechanical injury, are found in nearly all kinds of turtles according
to Carpenter (1956), Hunt (1957), and my personal observation.
In T. ornata, however, the condition is less common than in the
specimens of T. Carolina described by Carpenter and in the remain-
ing species of Terrapene that I have examined.

Carpenter (1956:86) came to no conclusion as to the cause of
shell pitting in Terrapene Carolina but suggested that a variety of
factors including parasitic fungi, parasitic invertebrates, and simple
shell erosion, might be responsible.

According to my own observations on turtles in the University
of Kansas collections, shell pits range in size and shape from shal-
low, barely discernible depressions to deep borings; I suspect that
shell pitting for turtles in general has many causes, some of which
may be of more frequent occurrence in one species than in another.

Hunt (1957:20) presumably was referring to shell pitting by a
more suitable name when he wrote of, ". . . necrosis . . .
of mycotic origin." Hunt (loc. cit.) stated that "Of those cases
which have been recently examined, the author found all were due
to the invasion of Mucorales beneath the plates of the epidermal
laminae. This disease is of extremely common occurrence and has
been found in all members of the order but is seldom found in
marine species. Mycosis more frequently occurs on the plastron
than on the carapace." Hunt presented no evidence to support his
statement regarding invasion of the shell by Mucorales.

Evidence that injury to the soft parts of the body is also fairly
common is seen in the many T. ornata with missing feet and legs.
Stumps resulting from amputations are covered with tough, cal-
loused skin and sometimes by horny tissue similar to that of the

Natural History of the Ornate Box Turtle 641

antebrachial scales. Amputees are incapacitated only slightly in
normal locomotion if a functional stump remains; probably a crip-
ple is somewhat handicapped in other functions, such as burrowing,
nest digging ( females ) , and copulation ( males ) . Causes of ampu-
tation are discussed in the section on predators.

Fractures of the limb bones are common. A female from StaflFord
County, Kansas (PI. 29, Fig. 4), showed a typical case of fracture
and subsequent repair; the right fibula had been broken and the
ends dislocated; a great mass of bone joined the repaired break to
the middle of the tibia, giving the entire skeleton of the leg the
appearance of the letter "H." The fibula, shortened by the dislo-
cation, no longer articulated by its proximal end with the femur;
the tibia probably bore the entire load in the period of repair and
the transverse connection that formed between the bones later
took over the function of the fibula.

There is little doubt that ornate box turtles are stepped on or
trampled by cattle, at least occasionally, but I never observed such
an incident; the predilection of ornate turtles for dung insects and
for moving along cattle pathways brings them to close quarters
with cattle and probably did likewise with native ungulates. A
steer, stepping on a box turtle, could inflict superficial damage to
the shell or cause broken limbs but would probably not crush the
turtle unless on a hard substrate.

REPAIR OF INJURIES TO THE SHELL

Most adults and a few juveniles examined in the field and labora-
tory had one or more small injuries on the carapace that had healed
or were undergoing repair. Such injuries almost never occurred on
the plastron. In an injury that was undergoing repair, a small piece
of smooth, whitened bone was exposed where a piece of epidermis
was missing from the shell. One or more edges of the exposed bone
characteristically projected over the surrounding epidermis, making
the bone appear as though it had been driven forcefully, hke a
splinter, into the shell (PL 29, Figs. 1 and 2). Because of their
curious appearance, small areas of repair were referred to in my
notes as "splinter scars." The position and number of splinter scars
were often recorded as supplementary means of individualizing
turtles in the field.

SpHnter scars result from minor abrasions that damage a few
square millimeters of the shell. Larger areas of exposed bone were
noted in only a few specimens. Two turtles at the Damm Farm had

642 University of Kansas Publs., Mus. Nat. Hist.

bone exposed on more than one-half the surface area of the cara-
pace; both of these turtles were probably burned in the grass fire of
1955. Ordinarily, a break in the shell does not induce extensive
regeneration of tissues; when shells are damaged by crushing or
cracking, regeneration of epidermis and bone occurs only along
the hnes of fracture, unless tlie broken parts have been dislocated.
Ligamentous tissue develops in some breaks on the plastron, the
broken area remaining shghtly movable after heahng is completed
(PI. 24).

Dissection of injured shells revealed the mode of shell regenera-
tion to be tlie same whether a large or small portion of the shell had
been damaged. An abrasion may gouge out a small portion of the
shell; burning, freezing, or concussion may kill a portion of the
epidermis and a corresponding portion of bone beneath it without
actually disfiguring the shell. Dead bone and epidermis become
loosened at tlie margin of the wound. The epideiTnis sloughs oflF
soon afterward but the bone adheres to the wound. New epidermis
and new bone, growing from undamaged tissues at the edges of the
wound, encroach on the wound beneatli the layer of dead bone.
The piece of dead bone is thereby gradually isolated from the rest
of the shell and is sloughed oflF when healing is complete. The dead
bone may come off in one piece or slough off gradually at its edges
as heahng proceeds toward the center of the wound. The layer of
dead bone protects the wound during the process of regeneration
(PI. 30). Areas of exposed bone become white and shiny, nearly
enamelhke in appearance, as a result of wear on the shell.

The above conclusions, in regard to T. ornata, agree basically
with the findings of Woodbury and Hardy (1948:161-162) and
Miller (1955:116) on regeneration of the shell in desert tortoises
(Gophertis agassizi). Danini (1946:592-4, Enghsh summary) made
histological studies on regeneration of the shell in specimens of
Emys orbicularis; he found that new bone trabeculae formed on
the surfaces of undamaged trabeculae at the edge of the wound
and formed also in connective tissue at the center of the wound.
Regeneration of bone was incomplete in some instances where total
extirpation of a portion of the shell had occurred. Regenerated
epidermis was usually thicker than the original scute.

Exposed bone on the shells of turtles that have been injured in
fires, although dead, is unmarked and shows no evidence of being
burned. Exposure to fire kills the growing portions of both the
epidermis and the bone but seemingly does not actually char or dis-

Natural History of the Ornate Box Turtle 643

figure the bone (although the epidermis may be so affected) (PI.
29, Fig. 3 ) . Injuries from fire result probably from brief encounter
with the fire itself or from more prolonged contact with some surface
heated by the fire. A turtle that remained in a fire long enough to
have its shell charred would presumably have little chance of sur-
vival. Grossly disfigured shells therefore do not result directly from
burns but are due to the gnarled texture of the regenerated bone
and epidermis remaining after the dead portions of the shell have
been sloughed off. Information on injuries from fire was supple-
mented by examination of several badly burned specimens of T.
Carolina. Their shells were nearly covered with exposed bone and
regenerated epidermis. One specimen was so badly damaged that
the entire anterior rim of its carapace was loose and could be pulled
away easily to disclose a gnarled mass of regenerating bone beneath
it (PI. 29, Fig. 3). There were areas near the posterior margin of
the carapace of each specimen where regenerated epidermis was
evident but where the bone was seemingly uninjured; the re-
generated epidermis was nearly transparent.

Areas of regenerated epidermis on specimens of T. ornata were
rough in texture and slightly paler than the surrounding scutes.
Color-pattern is not reproduced in the process of regeneration but
irregularly shaped light blotches sometimes occur in the places
where radiations or other distinct markings formerly were present.
A slight depression remains on the shell after regeneration is com-
pleted. I suspect that small injuries may be repaired in the course
of a single growing season but that injuries involving a large part
of the shell may take several years to heal completely. Cagle
(1945:45) reported that a bullet wound in the shell of a painted
turtle {Chrysemys picta) healed completely in approximately 23
montlis. Danini (loc. cit.) found that regeneration of the shell in
Emys orbicularis was complete in as short a time as 225 days.
Woodbury and Hardy (loc. cit.) stated that small injuries to the
shell of Gopherus agassizi may take as long as seven years to heal.

ECTOPARASITES

Two kinds of ectoparasites were found on ornate box turtles
in the course of the present study; larvae of chigger mites ( Trom-
bictila alfreddugesi) were abundant on specimens collected in
summer and, larvae of the bot fly {Sarcophaga cistudinis) were
found on specimens throughout the season of activity, and, in a
few instances, on hibernating turtles. In general, these ectoparasites

644 University of Kansas Publs., Mus. Nat. Hist.

do little or no harm to ornate box turtles, although heavy infes-
tations may cause temporary interruption of normal activity or may
even cause occasional death.

Concerning the larvae of T. alfrcddtigesi, Loomis (1956:1260)
wrote, "In northeastern Kansas, larvae become numerous in early
June (shortly after they first appear), increase in numbers to
greatest abundance throughout late June and July, decrease
slightly in August, become markedly reduced in September, and
only a few larvae (mostly on hosts) remain in October and early
November." He considered T. alfrcddtigesi to be the most abun-
dant chigger mite in Kansas and stated {op. df.:1265) that it is
most common ". . . in open fields supporting good stands of
grasses, weeds and shrubs, and where moderate to large popula-
tions of vertebrates are present." Loomis listed ornate box turtles
(op. cif.: 1261-2) as important hosts of Tromhicidn alfreddugesi
but noted that box turtles are not so heavily infested as are certain
other reptiles. The two other species of chigger mites that Loomis
{op. ciY.:1368) found on T. ornata in Kansas (T. lipovskyana and
T. montanensis) were not found in the present study.

Box turtles were considered to have chigger infestations when
the reddish larvae could be detected with the unaided eye. No
chiggers were seen on turtles in the period from spring emergence
until June 13, 1955. On the latter date a few scattered chiggers
were noted on several individuals and it was on this same date
that the writer received his first "chigger bites" of the year. Num-
bers of chiggers increased in the latter half of June and heavily
infested turtles were noted throughout July. No chiggers were
seen on box turtles after mid-September in 1955.

Chiggers were ordinarily found only on the soft parts of tlie
turtles' bodies. Early in the season infestations were chiefly on
the head and neck. Favorite sites of attachment were the point
where the skin of the neck joins the carapace and on the skin
around the eyes. Later in the season some chiggers could be
found on nearly every part of the body where soft skin was present;
concealed areas of skin, such as the axillary and inguinal pockets,
the anal region, and the inner rim of the carapace (where it joins
the skin of the body), harbored concentrations of chiggers. Ju-
veniles were relatively more heavily infested than adults and, even
early in the season, had chiggers attached along many of the inter-
laminal scams of the shell. Broad areas of soft, newly-formed
epidermis on the shells of juveniles probably afforded a better

Natural History of the Ornate Box Turtle 645

place of attachment to chiggers than did the interlaminal seams
of adults. The interlaminal seams and transverse hinges of adults
were not infested until the height of the season of chigger activity.
Heavily infested adults, observed in early July, were literally cov-
ered with chiggers; red larvae outlined nearly all the scutes of the
shell, the anus, the mouth, and the eyes. When turtles were picked
up for examination, chiggers could be seen moving rapidly from
one interlaminal seam to another.

Box turtles kept in outdoor pens and in the laboratory did not
long maintain visible infestations of chiggers, even during the time
in summer when turtles found in the field were heavily infested.

A four-year-old juvenile was found nearly immersed in the shal-
low water of a pond on July 4, 1955; its right eye had been dam-
aged by an especially heavy concentration of chiggers. When I
released the turtle, some 50 feet from the pond, it returned to the
water and spent the next four days there. The turtle was probably
in a period of quiescence induced by the eye injury and the heavy
infestation of chiggers; immersion in water could be expected to
help free the turtle of chiggers and to relieve trauma resulting
from the injured eye. Richard B. Loomis told me that larval chig-
gers are able to survive under water for several days but that warm
water will hasten their demise.

Infestations of larval bot flies ( Sarcophaga cistudinis ) were noted
in several turtles at the Damm Farm and, upon closer scrutiny, were
found to be common in preserved specimens from other areas. Lar-
vae were always found in flask-shaped pockets (PI. 27, Fig. 2) be-
neath the skin; the pockets opened to the outside by a small hole,
the edges of which were dried and discolored. One larva some-
times protruded from the opening. The inside of the pocket is lined
with smooth, skinlike tissue. Heavily infested box turtles may have
four or five such pockets, each containing one to many larvae. The
most frequent sites of the pockets are the skin of the axillary and
inguinal regions, and the skin of the limbs and neck, especially near
the bases of these members. Subadults were more heavily infested
than older adults; no infestations of hatchlings or small juveniles
were noted.

An adult female, infested with bot fly larvae when she was re-
moved from her hibernaculum in late October, 1955, bore no trace
of larvae or of the pocket that had contained them when she was
recaptured the following June. According to Rokosky (1948), the
larvae eventually fall to earth and pupate. The individuals of T.

646 Untversity of Kansas Publs., Mus. Nat. Hist.

Carolina studied by him were not re-infested by adult bot flies; one
turtle ate some of the larvae that dropped from its body.

The manner in which box turtles are infested by bot fly larvae
is uncertain. Possibly the eggs are picked up accidentally or laid
on the skin while box turtles are foraging in dung. Belding ( 1952:
841) classifies the genus Scaropliaga as semi-host-specific, deposit-
ing eggs in open wounds.

McMullen (1940), Rodeck (1949), and Rainey (1953), described
individuals of T. ornata parasitized by S. cisttidinis. Rokosky
(1948) and Peters (1948:473) reported infestations in T. Carolina.
Infestations were the cause of death in the instances noted by Rainey
and Rokosky.

PREDATORS

Few first-hand observations on predators of T. ornata are avail-
able and I have found little direct evidence of predation in the
course of this study. In general, adults of the species seem to have
few natural enemies other than man. Several of my colleagues at
the University of Kansas have observed dogs carrying box turtles
in their mouths or chewing on them. Frank B. Cross told me his
dog caught and ate young T. ornata in Payne County, Oklahoma,
and A. B. Leonard once saw a badger carrying one in Dewy County,
Oklahoma. At the Reservation, a freshly killed juvenile was found
beneath the nest of a crow (Corvtis brachyrhynchos) and remains
of a hatchling were found in a scat of a copperhead (Agkistrodon
contortrix ) .

Dr. Fred H. Dale, Director of the Patuxent Research Refuge,
Laurel, Maryland, kindly furnished photostatic copies of cards, from
the Division of Food Habits Research of the U. S. Fish and Wild-
life Service, recording the instances in which Terrapene ornata
was listed as a food-item. In one instance the stomach of each of
two nestlings, in the same nest, of the White-necked Raven ( Corvus
crypt oleucus) in Terry County, Texas, contained remains of re-
cently hatched ornate box turtles; the remains of one turtle made
up 64 per cent of the contents of one stomach, and parts of tliree
turtles made up 80 per cent of the contents of the other stomach.
Each of two stomachs of the coyote {Canis latrans) from Quay
County, New Mexico, contained a "trace" of ornate box turtle.

Wild carnivores known to occur on the Damm Farm were rac-
coons (Procyon lotor), striped skunks (Mephitis mephitis), badgers
(Taxidea taxus), and coyotes (Canis latrans); all were suspect as
predators of ornate box turtles.

Natural History of the Ornate Box Turtle 647

On December 10, 1953, ten dead box turtles (eight adults and
two juveniles) were discovered at the top of a cut bank on the
Damm Farm, within a few feet of a burrow that was used at least
part of the time by a striped skunk. The condition of the turtles
suggested that they had lain in the open for several weeks. The
heads and legs were missing from most of the turtles and tooth
marks were discernible on several of the shells. A logical explana-
tion of this occurrence is that the turtles, using the burrow as a
hibemaculum, were ousted by a predator that also inhabited the
burrow. Turtles moving about sporadically in late autumn may
be quickly chilled by a sudden drop in temperature and therefore
be more susceptible to predation than at other times of the year.
Two of my colleagues at the Museum of Natural History informed
me that they had observed similar concentrations of dead T. ornata
in winter.

In July, 1952, H. B. Tordoff collected eight shells of juvenile T.
ornata in a dry creek bed near Sharon, Barber County, Kansas.
Some of the shells had small tooth-punctures. The stream bed
habitat and the appearance of the tooth punctures tended to in-
criminate raccoons as predators. Raccoons, more than any other
carnivore mentioned above, possess the manual dexterity necessary
to pry open the shell of a box turtle and bite away the soft parts.
Badgers and possibly coyotes are probably the only local carnivores
(excluding large dogs) that could crack open the shell of an adult
turtle by sheer force.

Adults of T. ornata, since they occasionally molest small juveniles,
must be considered in the category of predators. When captive
adults and juveniles were fed from the same container in the
laboratory, the turtles occasionally bit one another accidently. Ser-
ious injury to the young was prevented by watching the adults
closely and moving them away when they caught a smaller turtle
by the leg or head. Similar accidents presumably occur in nature;
juveniles and adults were sometimes found feeding side by side.
WiHiam R. Brecheisen told me that adults kept in a stock tank at
his farm in the summer of 1955 regularly and purposefully chased
and bit small juveniles in the same tank. Brecheisen gave me a
juvenile that had been so bitten; the right side of its head was
badly damaged (the eye gone and a portion of the bony orbit
broken ) but was partly healed. Ralph J. Donahue told me that he
saw an adult T. ornata attack a juvenal T. Carolina, and provided
a photograph of the incident. The juvenile was not injured.

648 University of Kansas Publs., Mus. Nat. Hist.

Although small box turtles may occasionally be caught and
killed by adults in nature, this seems not to constitute a major
source of predation on the young.

Other animals that may prey upon young box turtles occasionally
( and that were known to occur at the Damm Farm ) are buUsnakes
(Pituophis catenifer), red-tailed hawks (Buteo jamaicensls), marsh
hawks (Circus cyaneus), crows (Corvus hrachyrhijnchos) , and
opossums (Didelphis marsiipialis) , and domestic cats.

Nest predators probably have greater effect on populations of
T. ornata than do predators of hatchlings, juveniles, and adults.
Four robbed nests were found at the Damm Farm; in each instance,
striped skunks were thought to be the predators. E. H. Taylor
told me that he once saw a buUsnake swallow an entire clutch of
newly laid eggs before the female turtle could cover the nest.

DEFENSE

Box turtles rely for protection on the closable shell and on in-
conspicuousness; defense reactions, except in the rare instances that
biting is provoked, are purely passive.

Box turtles handled in the course of field work varied widely in
their reactions. Many struggled violently when being measured or
marked whereas others were completely passive, closing the shell
tightly and making it difficult for me to examine the soft parts of the
body. These differences in behavior did not seem to be correlated
either with sex or with age; generally lessened activity was associ-
ated with suboptimum body temperatures. All box turtles found in
the field were extremely waiy. As soon as one sighted me (some-
times at a distance of 200 feet or more), it became motionless with
shell raised from the ground and neck extended (PI. 28, Fig. 5).
Some turtles remained in this motionless stance for half an hour or
more, finally moving slowly away if I remained motionless. Turtles
made no attempt to escape until I approached them closely or until
they were in danger of being trampled by my horse; they would
then move away with remarkable rapidity. Box turtles seemed
unaware of an intruder until he could be seen or until he touched
the turtle. When a turtle was approached from the rear, whistling,
finger snapping, and normal footfalls did not attract its attention.
Latham (1917:16) observed corresponding behavior in T. Carolina.
Wever and Vernon (1956) found the ear of T. Carolina to be keenly
sensitive to sounds in the range of 100-600 cycles per second but
progressively less sensitive to sounds of higher and lower frequen-

Natural History of the Ornate Box Turtle 649

cies. Surely a predator as stealthy as a coyote could approach a
box turtle unseen and could quickly bite ofiF at least one of the
turtle's legs. Many of the mutilated box turtles that I observed may
have sui"vived such encounters with carnivores. The tendency of
some individuals, when handled, to over-extend the limbs and neck
(rather than closing the shell) in an attempt to escape, would make
them easy victims for any predator.

Ornate box turtles were kept in my home, along with several cats.
Initial behavior was characterized by mutual wariness; subsequently
the cats would follow a turtle about the house for a time, occasion-
ally pawing at an exposed limb. The turtles withdrew only when
touched or when approached from the front. After a day or two
the cats and turtles ignored each other, often eating and drinking
from the same dishes without incident. Under these circumstances
the cats, I believe, could easily have killed or injured the turtles. A
turtle would occasionally gain the respect of a cat by biting it.

The strong odor sometimes given oflF by box turtles is produced
by the secretions of four musk glands, two situated anteriorly on
each side and opening by small, nearly invisible apertures beneath
tlie fourth marginal scute. According to HofiFman (1890:9), two
other musk glands, opening beneath the eighth marginal scute on
each side, are also present in Terrapene; these posterior glands were
not found in the several specimens of T. ornata tliat I dissected.

Strong odors were produced by nearly all small juveniles until
they became accustomed to being handled. Older juveniles and
adults produced strong odors only in response to pain or injury, as,
for example, when they were killed in the laboratory prior to preser-
vation or when they were being marked in the field. Young box
turtles were capable of producing strong odors as soon as they
hatched.

Norris and Zweifel (1950:3) considered the odor produced by
T. o. luteola to issue from the ". . . concentrated, highly pungent
urine . . ." voided by individuals when they were disturbed, and
thought the production of odor to be a defense mechanism. Neill
(1948b: 130) reported that hatchlings of T. Carolina with unhealed
umbilical scars emitted a musky odoi comparable to that of the
stinkpot, Sternotherus odoratus; he thought the capacity to produce
this odor was lost at about the time that the plastral hinge became
functional.

The function of musk glands in Terrapene and, in all other turtles,
is unknown. Since biting and nuzzling of the edges of the shell is

650 Unr'ersity of Kansas Publs., Mus. Nat. Hist.

an integral part of the courtship of many turtles, odor produced by
the musk glands may well be a means of social recognition or of
sexual stimulation. Repellant odor may have a protective value in
young box turtles but it is unlikely that larger predators would be
frightened away or even discouraged by odor alone. In this respect
Neill (loc. cit.) and I concur.

DISCUSSION OF ADAPTATIONS

Most of the morphological characteristics distinguishing box
turtles from other North American emyid turtles, the most notable
of which is the movable plastron, are modifications that have
evolved as a result of selectional pressures favoring adaptation to
more or less terrestrial existence. Similar adaptations have arisen
independently in several branches of the emyid stock (see intro-
duction). The genus Terrapene seems to have departed farther
from a generalized emyid form than have other kinds of box-turtle-
like chelonians. In a morphological sense, Terrapene ornata is
clearly the most specialized member of its genus now occurring
in the United States (my own studies have revealed that popula-
tions in western Mexico now referred to as T. klauberi and T.
nelsoni are as specialized as T. ornata in some respects but more
generalized in others). The present ecological study has dem-
onstrated that T. ornata is specialized in habits as well as in struc-
ture. It is concluded that these specializations (of more gen-
eralized and perhaps more primitive conditions as, for example
in T. Carolina) constitute adaptation for terrestrial existence in
open, semiarid habitats. These adaptations in T. ornata have re-
sulted, in a few instances, in unique habits and structures; how-
ever, in most instances the adaptations have produced slight but
recognizable changes that are definable only by degree of dif-
ference from other species of box turtles.

The closable shell of box turtles is of obvious survival value
in providing protection for the soft parts of the body. In most
of the species of Terrapene, the lobes of the plastron completely
close the openings of the shell; closure is so tightly efiFected in
some individuals that it is difficult to insert the blade of a knife
between the adpressed margins of carapace and plastron. In
T. ornata nevertheless, both lobes of the plastron are deficient on
their lateral margins; four narrow openings remain when the lobes
are drawn shut. Emargination of the plastron has occurred at the
places where the limbs rub against it during locomotion. This

Naturai. History of the Ornate Box Turtle 651

reduction of the plastron permits the body to be held oflF the
ground during forward locomotion and seemingly permits a gen-
erally freer range of movement for the hmbs. The possible dis-
advantages of an imperfectly closable shell seem to be compen-
sated for by increased mobility. Reduction of the plastron is
correlated with a general lightening of the shell, probably as-
sociated with the increased vagility of this species. Lightening
of the shell is evident also in the relatively thin, loosely articulated
bony elements. Shells of adult T. ornata that are old and weath-
ered, or macerated (unless they are partly co-ossified because of
injury), can nearly always be disarticulated with ease, whereas
the bony elements in the shells of adult T, Carolina (all races) are
nearly always co-ossified or separable only after prolonged macera-
tion.

The relatively low, flattened shell of T. ornata is an adaptation
associated with the tendency to seek shelter in the limited space
of earthen forms, burrows, or small natural cavities in the course
of the warm season and to burrow more deeply into the ground
in winter. Terrapene ornata is, in fact, the only species of tlie
genus that may be considered an habitual burrower. Individuals
of T. Carolina tend to seek shelter in the warm season by making
forms in dense vegetation or by digging into yielding substrata
such as mud or humus, although they may burrow deeply into the
earth in winter. Extreme weakness or absence of the middorsal
keel of T. ornata seems to be a modification associated with bur-
rowing habits and general adaptation to terrestrial life; the keel
is similarly reduced in testudinids.

Retention of epidermal laminae (as opposed to regular exfolia-
tion of the older parts of scutes) occurs in all box turtles, in sev-
eral other groups of terrestrial emyids, and in testudinids. The
phenomenon is here considered to be a specialization of scute
shedding — developed in terrestrial and semiterrestrial chelonians
— ^that provides additional protection to the shell against wear and
minor injuries.

General shortening of digits — the result of reduction in number
of phalanges as well as in their length, and to a lesser degree the
shortening of metapodial elements — ^has occurred in several groups
of chelonians with terrestrial tendencies (the opposite — lengthen-
ing of phalanges and metapodials, and hyperphalangy — has oc-
curred in certain groups that are highly aquatic). The pes of box
turtles has remained relatively unchanged in this respect; a few

652 Untv-ersity of Kansas Publs., Mus. Nat. Hist.

phalanges on the lateral digit have been lost (especially in three-
toed forms), but little reduction in length has occurred. The chief
modification of the pes is a general narrowing brought about by
the tendency of the digits to be crowded together, one on top of
the other, ratlicr than spread in a horizontal plane. Considerably
more modification is seen in the manus of Terrapene. Phalangeal
formulae (expressing the number of phalanges from the first digit
outward) range from 2-3-3-3-2 (primitive in Terrapene) to 2-3-3-2-2
in tlie races of Carolina and have the same range in the species of
eastern Mexico. Extreme reduction in number (2-2-2-2-2) as well
as general shortening of phalanges occurs in T. omata. The for-
mula is the same in the one specimen of T. klauberi that has been
skeletonized. This modification of the forelimb in T. ornata has
produced a more rigid, stronger manus that is well adapted to the
requirements of burrowing and to locomotion over unyielding
substrata. Shortening of the manus (and, to a lesser extent, the
pes) has been accompanied by reduction and loss of interdigital
webbing. It is noteworthy that T. ornuta has achieved the same
reduction in number of phalanges as Gopherus, which displays the
extreme of specialization in this respect among North American
turtles. The manus in T. ornata is not shortened so much as in
Gopherus.

The first toe in males of T. ornata is uniquely widened, thickened,
and inturned. Males of some other species of Terrapene have
greatly enlarged rear claws, some of which turn slightly inward,
but none has the flexed first toe hooklike as it is in ornata ( a modi-
fied first toe, resembling that described for T. ornata, has been
observed in a live male of T. klauberi [now KU 51430] since the
preparation of this manuscript). In males of T. ornata the penul-
timate phalanx of the first toe has a normal, vertical articular sur-
face on its proximal end. However, the distal articular surface
(when viewed from the distal end of the phalanx) has its axis ro-
tated away from the vertical plane approximately 45 degrees in a
counterclockwise direction. As the foot is pronated and extended,
and as the digits are flexed, there is a concomitant inward rotation
of the first metatarsal at its proximal joint; this rotation, combined
with the divergent planes of the articulating surfaces on the penul-
timate phalanx, cause the ungual phalanx to be flexed at right angles
to the inner side of foot, in a plane perpendicular to that of the
other toes (Fig. 21).

The precise function of the modified first toe of males is unknown,

Natural History of the Ornate Box Turtle 653

although it is reasonably safe to assume that the modification is
closely associated with clasping during coition. In the matings
that I observed, the inturned first claw of the male secured a hold
on the female's rump or just beneath her legs, whereas the remain-
ing three toes gripped the edge of her plastron. The combined hold,
on shell and skin, clearly affords the male a more secure position
during coitus (whether the female clasps his legs with hers or not)
than would a hold on skin or shell alone. Possibly intromission can
be maintained in this position even when the female is attempting
to escape. In males the plastron is less concave in T. ornata than
in T. Carolina. Furthermore, males of T. ornata are, on the average,
smaller dian females, whereas the reverse is true in T. Carolina.
Possibly the ability of the male to secure an especially firm grip
on the female enhances the probability of small males mounting
and inseminating larger females, whereas successful matings might
otherwise be limited to pairs in which the male was the larger
member.

It is worthy of note that turtles of the genus Terrapene are seem-
ingly the only North American emyids that carry out the entire
process of mating on land; other, semiterrestrial emyids (for ex-
ample, Clemmys insculpta and Emijdoidea blandingi) return to
water for actual coition, although the precoital behavior sometimes
occurs on land.

Nearly all gradations from a fully developed zygomatic arch to
a greatly reduced arch can be observed in skulls of the various spe-
cies of Terrapene (Fig. 2) (Taylor, 1895:586, Figs. 2-7). The
highest degree of reduction is achieved in T. ornata and T. klauberi,
both of which lack the quadratojugal bone and have no zygomatic
arch whatever (except for an occasional, poorly defined anterior
vestige formed by the postfrontal, the jugal, or both). Reduction
of the zygoma clearly represents modification of a more generalized,
complete arch. As yet there is no clear evidence that reduction of
the zygomatic arch is of adaptive value. It is noteworthy, how-
ever, that similar reduction of the arch has occurred independently
in a number of emyid and testudinid groups, nearly all of which
have terrestrial or semiterrestrial habits. Although discussion of
phyletic lines in Terrapene is beyond the scope of this report, I
tentatively suggest that reduced zygomatic arches have arisen in-
dependently in more than one group of Terrapene and that similar
reduction of the arch in two species of the genus does not neces-
sarily indicate an especially close relationship of such species.

654 UNrvERSiTY OF Kansas Publs., Mus. Nat. Hist.

In a recent survey of cloacal bursae in chelonians, Smith and
James (1958:88) reported T. ornata and T. mexicana to be among
the few emyids that lacked these structures; in the opinion of the
authors (op. cit.:94) cloacal bursae evolved in chelonians that re-
quired an accessory respiratory organ for long periods of quiescence
( hibernation or aestivation ) under water, and were secondarily lost
in terrestrial forms that hibernated on land. The assumption is a
reasonable one, at least in regard to emyids and testudinids. Lack
of cloacal bursae in T. ornata and in all testudinids, can be corre-
lated with the completely terrestrial habits of those turtles. Clo-
acal bursae seem to be vestigial in the species of Terrapene pos-
sessing them and to be of little or no use as respiratory structures
( except perhaps in T. coahuila ) .

In most of the species of Terrapene the carapace has a pattern of
pale markings on a darker background; however, unicolored in-
dividuals are the rule in certain populations (for example, at the
western edge of the range of T. Carolina and in T. ornata luteola)
and occur as occasional variations in other populations (in T. yuca-
tana, T. mexicana, and, throughout the range of T. Carolina, albeit
more commonly in the southeastern part of the range). Personal
observation of interspecific and ontogenetic variation of color
patterns of box turtles has convinced me that a basic pattern of
more or less linear radiations is the one from which all other patterns
(including spots, blotches, rosettes, and the unicolored condition)
can be derived, and, that tlie radial pattern is generalized and
primitive for Terrapene (possibly for all emyids and testudinids as
well). In the light of tliis conclusion, the radial pattern of T. ornata
may be considered generalized. I suspect, however, that the pattern
of a Hving species most closely approaching that of the primitive
ancestral stock of Terrapene is the pattern of fine, wavy, dark radi-
ations (on a paler background) present in young examples of T.
coahuila.

Box turtles in general have lower reproductive potentials (as
indicated by fewer eggs and longer prepuberal period) than do
most aquatic emyids. This low potential seems to be compensated
for by a lower rate of postnatal mortaUty (especially in adults) due
to the protection ajfforded by the closable shell and the abiHty to
recover from serious injury. Terrapene o. ornata and T. c. Carolina
are the only box turtles the fife histories of which are known well
enough to permit significant comparison. The reproductive po-
tentials of T. o. ornata and T. c. Carolina seem to be much the same.

PLATE 15

9—773

PLATE 16

Fig. 1. A water-filled ravine in the northern part of the pasture at the Damm
Farm (June 28, 1958). The subdivision of the pasture referred to in te.xt as
"northwest comer area" can be seen sloping into the ravine from the west

(left background).

i

Fig. 2. A cow path leading southward away from a ravine, at the Damm

Farm (June 28, 1958). Ornate box turtles used such paths as routes of travel

in the course of their daily activities.

PLATE 17

Fig. 1. Grassland on crest of hill at Damm Farm with northeastern corner
of main pasture in background (June 29, 1958).

Fig. 2. A bare area along the rock fence at northern edge of pasture at
Damm Farm. Ornate box turtles could nearly always be found foraging
in cow dung here and in similar areas along other fences (June 28, 1958),

PLATE 18

Fig. 1. A ravine in the southern part of the pasture at the Danini Farm
(June 28, 1958). Small springs at the heads of such ravines produced
marshy conditions at their bottoms and provided drinking water, in the
form of shallow pools, for box turtles for at least part of the year. Banks
of ravines provided suitable sites for the construction of nests and forms.

^■0 **•. •

Fi(.. 2. A mulberry tree on the bank of a ra\int' near nortliern edge of

Damm pasture (June 28, 1958). Box turtles frecjuented the area beneath

the tree when fruit fell to the ground in June and July. The ravine shown

here filled with water after being dammed in June, 1956.

PLATE 19

Representative stages in the spermatogenic cycle of T. o. ornuta (all speci-
mens obtained in Douglas County, Kansas, 1955). Figs. 1 to 5, respectively,
are sections of seminiferous tubules obtained on May 17, June 14, July 15,
Aug. 31, and Oct. 4. Fig. 6: seminiferous tubule of immature male (plas-
tral length, 88 mm.), six years old, obtained on June 30. Fig. 7: section
of epididymis from mature male obtained on April 17, three days after
turtle emerged from hibernation; mature sperm form a continuous dark
mass in center of epididymis. Fig. 8: spenn in uterine portion of oviduct
of female obtained on April 18, 1954. Figs. 1 to 6 and 8 -were photo-
graphed X 430, and v^'ere enlarged 1.4 times. Fig. 7 was photographed
X 35, and was enlarged 1.4 times.

PLATE 20

Fig. 1. Left ovary of mature female, prior to ovulation, May 15, 1956
( X 1 ). Fig. 2. Fresh eorpus luteiuii, June 2, 1956 ( X 4)2). Fig. 3. Testes
of mature male, August 31, 1955 (X 1). Fig. 4. Testes of mature male,
April 14, 1956 (X 2). Fig. 5. Left ovary of subadult female (seven years
old, plastral length, 114 mm.) that would have matured in approximately
one year (x Di). Fig. 6. Left ovary of juvenal female (11 years old,

plastral length, 95 mm., X 1)«).

PLATE 21

Fig. 1. A trial nest cavity excavated by a gravid T. o. ornata at the Damm
Farm on June 8, 1956. The cavity was situated at the edge of a grassy area
on the upper rim of a ravine embankment. Twelve-inch ruler shows scale.

±M^^^

1^.':^-"v..-'ilr

Fig. 2. A depression, resulting from an old post-hole, showing the openings
made by three box turtles as they left their hibemacula in April, 1956 (photo-
graphed May 15, 1956). Twelve-inch ruler shows scale.

PLATE 22

Fig. 1. Right abdominal lamina (X 2/0 of a four-year-old juvenal male
showing method of measuring growth-rings. The last growth-ring (4) was
formed at the end of tlie 1954 growing season. Tlie first growth-ring (H)
marks the end of the season of hatching (19.50). The vunbilieal scar (U)
is faintly visible. The growth-zone for 1955 (specimen captured June 23)
is just beginning to show in interlaminal seam.

Fic. 2. Left — Right abdominal lamina (X 2) of subadult female, eight
years old. The last growth-zcme was formed in 1954. Note the relatively
small growth increments in 1952 and 1953. The growth-zone for 1955
(date of capture, May 8) is not yet visible. This specimen grew more in
the .season of hatching (1946) than the specimen shown above in Fig. 1.
Rif^ht — Interpectoral seam ( X 3) of adult male showing slowness of growth
in later life. The widest growth-zone seen here was fornu'd in tlu' tenth
year and is followed by four zones too narrow to measure accurati'ly. It
is uncertain whether this specimen was still growing in the vear it was cap-
tured (1923).

PLATE 2,3

IHICBS

r^**''*^^^*^-, 1

1 * -^ J:^-.^t^ S

#.f/^r^^m.

i|gK.^^-«r ^^^^^^fl

Ontogenetic change in color and markings of carapace. Radial markings
begin to develop at the onset of epidermal growth. Markings are sharply
defined in juveniles and young adults but may be obscured in later life
by the encroachment of dark ground color or by wear on the shell. Figures
are as follows: Upper left — Hatchling (x 1/2 ); Upper right — Juvenile
(X 1). one year old; Lower left — Juvenile (X 1), one year old; Lower
left — Female (X Vit;) showing typical adult markings; Lower right —
Adult male (X /z) showing blotched pattern resulting from wear on shell.

PLATE 24

Ontogenetic change in color and markings of plastron. Dark markings on
plastra of hatchlings are unbroken. Dark radiations appear when epidermal
growth begins. Figures are as follows: Upper left — Hatchling (Xl^*);
Ujyper rxp,ht — Juvenile (x 1); Lower left — Female (X vic) showing typical
adult markings; Lower rifiht — Adult male (X J^) showing the effect of wear
on markings. Flastra of old individuals are sometimes .solid yellow. Note
the break in the plastron that has healed and filled with ligamentous tissue.

PLATE 25

Ontogenetic change and sexual dimorphism in shape, color, and markings
of head and neck. Females retain much of the juvenal pattern of spots and
blotches. In males, the top and sides of the head become greenish or bluish
and markings are obscured. Figs. 1 and 3. Lateral and dorsal views of
hatchling (X 3^ ) ; Figs. 2 and 4. Lateral and dorsal views of juvenile
(X 2); Figs. 5 and 6. Adult female (X 1); Figs. 7 and 8. Adult male
( X 1 ) showing relatively wider and more truncated snout in this sex.

PLATE 26

Fig. 1. A foraging station next to a rock Icncc at the Dannn i''arni ( Jimc

28, 1958). TIh' box tnrtle in torcgronnd was in tin- act ot tearing apart a

pile ot partially dried cow dnng to oiitain dung inseets.

Fig. 2. A depression (X '2) made by a foraging box tnrth' in a pile ot
partially driecl eow dung (June 28, 1958). Similar "sign" of box turtles
was found in eow dung ever\ where on tln' pasture at the Dannn l''arm.

PLATE 27

Fig. 1. Thread-laying device ("trailer") taped to the carapace of an adult
female T. o. ornata. The squares of tape on the sides are to keep the bent-
over ends of the wire axle from catching on vegetation (X '2 ) •

Fig. 2. A dermal pocket ("cyst") removed from an adult T. ornata and
cut open to show two larval hot flies (Sarcophaga cistudinis) ( X 2, May

15, 1956).

Figs. 1-3. Stages in covirt.ship of T. o. orrxata: male pursuing female and
biting her shell; male lunging at female in attempt to mount; and, male
just after mounting female (X'')- Fig. 4. 7\ o. onuita smelling food
(X 1 )• I'ig- 5. T. o. ornata in attitude of alertness after deteeting intruder
(X^s). Fig. 0. Traeks of T. o. ornata in muddy ravine (X5«) (June 5,

1956).

PLATE 29

Fig. 1. A small, nearly-healed, injury on the carapace of an adult T. o. ornata
( X 2 ) . Note regenerated epidermis at bottom of depression and two pieces
of dead bone ( "splinter scars" ) at upper right margin of depression. Fig. 2.
Injured area on the carapace of a juvenal T. o. ornata ( X 3) with dead bone
removed and laid to the right, exposing regenerating epidermis in its early
stages. Fig. 3. Anterior edge of carapace (held away with forceps) of
specimen of T. Carolina (KU 51461, Gulf Co., Florida) that had been badly
burned ( X % ) • Nearly all the scutes of the shell had fallen off and large
pieces of dead bone could be pulled away, exposing a gnarled mass of re-
generating bone and epidennis. Fig. 4. A fracture that has healed and
joined the tibia (upper bone) to the fibula in a specimen of T. o. ornata

(KU 1877, X 3J^).

PLATE 30

Top: A shell of T. o. (Jimilu ( X '2) as it was found at the Damin Farm
June 1, 1956. A serious injury (probably resulting from burns) had exposed
a large area of dead bone on the earapace. Center: Same shell with some
of scutes removed. Bottom: Same shell with dead bone removed to expose
regenerating epidermis and bone. Note that the injury involved several
of the neural bones; the turtle probably died as a result of this injury but
not before regeneration was approximately one-half completed.

Natural History of the Ornate Box Turtle 655

Terrapene ornata seems to concentrate its breeding season
(laying, incubation, and hatching of eggs) more nearly in the
middle of the warm season than does T. c. Carolina. This concen-
tration probably is an adaptation for breeding in open habitats
where, under environmental temperatures less equable than in
forest, eggs would develop more rapidly and hatch sooner but would
be less able to survive winter temperatures.

Males of T. o. ornata become sexually mature when younger and
smaller than females and rarely grow as large as females. Nichols
(1939a: 20) indicated the reverse to be true of T. c. Carolina; Nichols
further indicated that growth continued some six to eight years after
puberty. Most individuals of T. o. ornata attain maximum size
within two to three years after puberty.

Although it is difficult to be certain about the adaptive value of
color and pattern, it seems that in box turtles, as in many other kinds
of animals, patterns and colors most nearly blending with those of
the habitat have some selective value in providing concealment
from enemies. The pattern of linear radiations in T. o. ornata close-
ly resembles the patterns formed by hght passing through grasses
and associated vegetation and camouflages the turtle. In a similar
manner, partial or complete loss of radial markings in T. o. luteola
seems to provide concealment in habitats where vegetation is sparse
and where blending with the substrate is of survival value. The
patterns of blotches and broken radiations in most of the subspecies
of T. Carolina likewise provide camouflage by tending to match
patterns formed by the light passing through a leafy canopy.

Although ornate box turtles are omnivorous, they probably de-
pend on insects as a dietary staple. In years when preferred kinds
of insects were unusually abundant, the turtles grew more than in
other years. A large proportion of the insects eaten is obtained
by foraging in or near dung. Alteration of the dung community — at
least in a physical sense, but presumably also by influencing the
successional stages of the dung biota — is one of the few evident
effects of box turtles on the environment. Although certain kino-
stemids (Carr, 1952:93), emyids ( Deraniyagala, 1939:257; Love-
ridge and Williams, 1957:198), and testudinids (Loveridge and
Williams, op. cit. -.247) eat mammalian feces, T. ornata is seemingly
the only chelonian that habitually seeks its staple diet in dung.
The habit seems to be yet another specialization for terrestrial
existence. The carnivorous habits of T. ornata reverse the general
trend toward omnivorous and herbivorous habits in other turtles

10—773

656 UxrvERsiTY of Kansas Publs., Mus. Nat. Hist.

that have become partly (emyids) or wholly ( testudinids ) ter-
restrial.

It seems remarkable that none of the species of true tortoises
occurring in the grasslands of the world has developed insectivorous
habits or utilized the unique food niche (in regard to dung-forag-
ing) filled by ornate box turtles in the Great Plains; tortoises are,
as far as is known, strictly herbivorous. The ranges of Gopherus
and Terrapene are now almost mutually exclusive and the two
kinds do not compete with each other for food in the few places
where they occur together. It is known, however, that box turtles
(T. longinsulae, ornata-like, earliest known box turtle) and true
tortoises (genera Testudo and Gopherus, see Williams, 1950:25-26,
Fig. 2) occurred together in what is now the Great Plains in early
Pliocene times and probably for some time before and after this.
Assuming that food habits of fossil representatives of these genera
were somewhat like the habits of recent representatives, ornate box
turtles may have developed insectivorous habits at a time when
other food niches were filled by herbivorous tortoises. Box turtles
possibly survived subsequent changes in habitat that made it im-
possible for populations of large tortoises to exist in the Great
Plains.

SUMMARY

Box turtles of the genus Terrapene are emyid turtles that are
specialized for terrestrial existence. Two of the seven species now
recognized — T, ornata and T. Carolina — occur in the United States.
Terrapene Carolina inhabits forested areas in the east whereas T.
ornata is characteristic of open grassy areas in the west; the ranges
of the two species overlap in the broad belt of prairie-forest ecotone
in the central United States. Terrapene ornata is considered to
be the most specialized of Hving box turtles.

The natural history of T. o. ornata Agassiz was studied in the
period, 1953 to 1957. Intensive field studies were made in Douglas
County, northeastern Kansas, on a small area of prairie and on the
University of Kansas Natural History Reservation. Field observa-
tions were made also in a number of other places in eastern Kansas.
Laboratory studies supplemented field studies.

Habitats occupied are chiefly open areas; they vary in regard
to food supply, temperature, moisture, and kind of soil. The
grassy prairies of Nebraska, Kansas, Oklahoma, and northern Texas
seem to provide optimum habitat for ornate box turtles; in these
areas box turtles are active on a majority of days from April to

Natural History of the Ornate Box Turtle 657

October. The subspecies luteola is adapted to the more rigorous
and arid environment of the southwestern United States, where
activity may be possible for only a few weeks in the year. The
remainder of the year is spent in a state of quiescence. Factors
hmiting the distribution of T. ornata are: 1) the presence of a
substrate too hard to permit digging of nests and forms ( altitudinal
distribution in southwestern United States and distribution at
western edge of the range); 2) temperatures causing the ground
to freeze deep enough (approximately 30 inches) to kill turtles in
hibernacula (northern edge of range); and, 3) the lack of one or
more relatively wet periods in the course of the warm season, pre-
venting at least temporary emergence from quiescence (south-
western part of range). The activities of man probably have af-
fected population density in local areas but limit the geographic
range only in the north (Blanchard, 1923:19-20, 24) where in-
tensive cultivation probably has excluded the species.

Preferred habitat in northeastern Kansas is open rolling grassland
grazed by cattle; populations are most dense near natural breaks
in the grassy vegetation such as fences, scattered rocks on hillsides,
ravines, and stream-beds.

Mating occurs most commonly in spring and autumn; courtship
behavior includes pushing and biting on the part of the male.
In coitus the hind legs of the male are held tightly by the female;
the male falls backward after coitus, still clasped by the female.
A few sperm are stored in the oviducts; fertilization without re-
insemination can occur. The spermatogenic cycle begins in May
and reaches its peak in September, when large numbers of sperm
and spermatids are present in the testes; the cycle is completed in
October, when sperm pass into the epididymides. The testes are
smallest in spring and largest in September. Females are insem-
inated with sperm produced in the preceding year. The ovarian
cycle begins in midsummer, soon after ovulation, and continues
up to the time of the next ovulation. Follicular growth is rapid
in the period from spring emergence to ovulation. Large folhcles
remaining after ovulation represent, in many instances, eggs that
will be laid later in the same season. Follicular atresia is never
great enough to account for the destruction of all large follicles
remaining after ovulation. All mature females lay at least one
clutch of eggs per year. It is estimated that one-third of the fe-
males produces two clutches of eggs in a single season. Second
clutches contain fewer eggs than first clutches. An alternation

638 University of Kansas Publs., Mus. Nat. Hist.

of ovarian activity occurs, whereby one ovary is more active than
its partner in one season and less active in the next season. Al-
ternating activity of ovaries accounts in part for the reduced num-
ber of eggs in young females, breeding for the first time, and in
older, nearly senile females. Extrauterine migration of ova re-
sults usually in a more even distribution of eggs in the oviducts.
Corpora lutea constitute an accurate record of the number of eggs
produced by the ovary as well as the number of eggs laid.

Nesting occurs from May through July but is most common in
mid-June; some of the females nesting early in the season lay a
second clutch of eggs in July. Nests are dug in the earth by the
female using her hind legs. Preferred nesting sites are open, well-
drained places with a soft substrate. The nesting site is selected
after a period of wandering, in which tlie female tests the substrate
at a number of places; some females search for a nest site for
more than a week. Nest digging begins in the evening and is
usually completed after dark. Captive females dug a preliminary
cavity in which the body rested during the digging of the main
nest cavity. The entire clutch of eggs is laid in one nest. The
average number of eggs in 23 clutches was 4.7 (range, 2 to 8).
The average size of eggs tends to be inversely proportional to the
number of eggs in a clutch. Eggs increase in bulk by absorption
of water in the course of incubation. Immersion in water for short
periods does not harm eggs. The incubation period under favor-
able environmental conditions is approximately 65 days; cool, damp
conditions prolong the incubation period and probably constitute
an important factor of prenatal mortahty in certain years. Eggs
that do not hatch before winter probably do not survive. Emer-
gence of hatchlings from the nest may, however, be delayed until
spring if the soil is dry in autumn. Hatchlings can probably es-
cape freezing by burrowing into the walls of the nest. Infertility
and prenatal mortahty account for at least 40 per cent of the eggs
laid, according to laboratory findings. Progeny of a single adult
female (considering factors of mortality, multiple layings, and
average age of puberty) would number approximately 300 after 20
years. Reproductive processes probably continue throughout life,
although possibly at a somewhat reduced rate in later life.

Young box tiutles are active soon after hatching but become
quiescent if allowed to burrow in soil or if they are covered with
damp cotton. Some captive hatchlings take live food in the first
days of life but others do not eat until the following sjDring; initia-

Natural History of the Ornate Box Turtle 659

tion of growth is coincident with initiation of regular feeding. The
yolk sac retracts mainly during hatching; it sometimes ruptures
after hatching. The caruncle remains on the beak for a variable
length of time, but never is present in the spring following hatch-
ing.

Major growtli-rings on the epidermal laminae are formed regu-
larly, one after each season of growth, in the first 10 to 14 years of
life. Minor growth-rings occur between major rings and are
shallower. Growth of epidermal laminae results from the formation,
in spring, of a new layer of epidermis beneath the existing scute.
The peripheral projection of the new layer is distinct in texture and
color from the older part of the scute and is separated from it by a
major growth-ring. Minor growth-rings form when growth slows or
temporarily stops during periods of quiescence; no new layer of
epidermis is formed. Growth-rings constitute an accurate record of
growth that can be studied at any time in the life of the turtle; they
are accurate indicators of age only as long as regular annual growth
persists.

Growth in the season of hatching depends on early hatching and
early emergence from tlie nest. Turtles that remain in the nest until
spring probably do not grow. Slightly less than half of the free-
hving individuals studied grew in the season of hatching. Pre-
cociousness in early life often results in the attainment of sexual
maturity at an earHer than average age.

Growth is rapid at first ( increments in plastral length average 68,
29, and 18 per cent, respectively, in the first three years) and then
slows gradually until puberty. Attainment of sexual maturity is
more closely correlated with size than with age. Males mature
when smaller (76 per cent were mature when plastron 100 to 109
mm. long) and younger (average age, eight to nine years) than
females (66 per cent were mature when plastron 110 to 119 mm.
long, average age at maturity, ten to eleven years) but females
grow larger than males. A few individuals of each sex reach
puberty tliree to four years sooner than average.

The average number of growing days per season is approximately
160. Amount of growth in any season depends on chmatic factors
that influence food supply and foraging conditions. Growth rate is
directly correlated with precipitation, being highest when large
populations of grasshoppers and long periods of favorable weather
occur in tlie same year. Zones of epidermis formed in years when
growth was especially slow or especially fast constituted landmarks

660 UxrvERsiTY of Kansas Publs., Mus. Nat. Hist.

that were helpful in interpreting growth-histories. Growth stops
two to thiee years after puberty. The total growing period is esti-
mated to be not more than 15 to 20 years. Longevity is estimated to
be approximately 50 years.

A number of changes in structure and appearance occur in the
period from hatching to puberty. Fontanelles of the bony shell
close at or before puberty. Movable parts of the plastron are not
functional until the fourth year. Markings on the carapace change
from a series of dots to distinct, straight-sided radiations, and a
similar pattern develops on the plastron. Markings on the heads of
females resemble those of juveniles but males have greenish heads.
Males further difFer from females in having a red iris, more brightly
colored antebrachial scales, and a turned in first toe.

Analysis of some 500 body temperatures (Centigrade) obtained
under natural conditions revealed the following: the optimum
temperature for activity is near 30 degrees; box turtles emerge from
cover usually when body temperature is 24 degrees or higher, and
almost never when the body temperature is below 15 degrees; body
temperature is raised to optimum by basking in open areas although
activity begins at suboptimum temperatures if basking is impossible;
cover of dens, burrows, or forms is sought when the body tempera-
ture rises above 30 degrees; and, maximum and minimum body
temperatures that would be lethal to box turtles (for prolonged
periods) are approximately 40 and zero degrees, respectively.
Laboratory experiments showed speed of response to environmental
temperature to be inversely proportional to bulk; hatchlings could
be chilled or warmed more tlian twice as fast as adults and were
active within a narrower range of temperature. Ornate box turtles
in general are subject to a narrower range of thermal activity than
are aquatic turtles that occur in the same areas.

Box turtles are dormant approximately five and one-half months
of the year — from late October to mid-April. Warm weather in
November and late March sometimes stimulates temporary activity
but dormancy is uninterrupted from mid-November to early March.
Forms, dens, and burrows are used as hibernacula. Depth of
hibernacula is dependent on severity of temperatures and amount of
vegetational cover; hibernacula in open grassland were seven to 18
inches deep whereas those in wooded areas were six inches or
shallower. Box turtles are ordinarily solitary when hibernating.
Injuries and deaths due to freezing probably occur in the coldest
part of the winter. The lowest body temperature of a turtle that

Natural History of the Ornate Box Turtle 661

survived a winter was 2.7 degrees; an individual, the temperature
of which was nearly zero for several days, subsequently died.
Turtles burrow upward at the end of hibernation and remain just
below the surface for a week or two before emerging. The primary
stimulus for emergence seems to be a period of warm moist weather.

Populations of T. ornata observed under natural conditions were
chiefly carnivorous, although captives ate a variety of animal and
vegetable matter. Insects, consisting chiefly of beetles, caterpillars,
and one species of grasshopper, comprised approximately 89 per
cent (by volume) of tiie food present in stomachs. Beetles (chiefly
scarabaeids and carabids) are obtained in or near dung and seem
to constitute the most important staple element of the diet. Piles of
dung, disturbed by turtles in the course of their foragings, were
characteristic "sign" of T. ornata in the areas studied.

Insects form the bulk of the diet for most of the year, although
certain other foods, when especially abundant for short periods
(mulberries for example), are eaten in large quantity or eaten to
the exclusion of all other foods. Ornate box turtles occasionally eat
the eggs and young of ground-nesting birds and slightly damage
vegetables, but in no instance do these feeding habits significantly
affect the economy of man. Box turtles probably benefit man by
destroying large numbers of crop-damaging insects (locustids and
noctuid caterpillars).

Box turtles were more numerous than most kinds of reptiles at
the Damm Farm and were the most conspicuous of any kind of rep-
tile. One hundred and ninety-four turtles were marked; one-fourth
of these were recaptured at least twice. Population density in cer-
tain areas of favorable habitat ranged from 2.6 to 6.3 turtles per
acre. The total number of individuals on the study area was esti-
mated to be 286. The marked population consisted of 53 per cent
adult or subadult females, 31 per cent adult males, and 16 per cent
juveniles of undetermined sex. Only six individuals had plastra
shorter than 60 millimeters. Small box turtles are not so rare as
these samples indicate; they are infrequently obtained because
their smallness and ability to blend with the substrate make them
difficult to see. More females than males were found in all months
of the season of activity, excepting April and August when more
males were found; the preponderance of females was greatest in
the nesting season ( June and July ) .

Ornate box turtles walk with the shell held off the substrate.
They are able to chmb steep embankments or low barriers with

662 University of Kansas Publs., Mus. Nat. Hist.

some facility. Swimming ability is sufficient to permit survival in
water and traversal of water-barriers but ornate box turtles almost
never swim voluntarily.

Daily activity consists of periods of basking, foraging, and rest,
the durations of which are influenced by temperature and humidity.
There is no activity after dark except that of nesting females. After
several days of activity there is a period of rest; rest periods seemed
not to be correlated with climatic conditions. The average distance
traveled per day in summer is 200 to 300 feet. Movements of
gravid females are more extensive (average, 363 feet per day) than
those of other members of the population; one individual traveled
approximately one-fourth of a mile in a single day. Turtles re-
moved from their normal home ranges traveled farther per day
than any other group. Movements in autumn are less extensive
(average, 152 feet per day) than at other times in the season of
activity.

Individual box turtles tended to remain in small areas for long
periods; these areas were interpreted as home ranges. The esti-
mated average radius of 44 home ranges was 278 feet (average
area, 5.6 acres ) . The average area of eight home ranges that were
actually measured was five acres. General suitability of habitat and
certain physical features of terrain (rock fences, ravines, barren
fields) that acted as barriers were thought to be the most impor-
tant factors governing size of home range. Of two turtles removed
more than one-fourth of a mile from their home ranges, one homed
and one did not. Home ranges of turtles of all ages and sexes
overlap broadly. There was no indication that territoriality or social
hierarchy existed in the population studied.

Box turtles are subject to injury from natural causes that include
fire, cold, molestation by predators, and trampling by cattle. Auto-
mobiles and farm machinery now constitute major causes of mor-
tality and serious injury. Capacity to recover after serious injury
is great but there is increased chance for secondary injury, infec-
tion, and predation in the period of recovery. Pits on the shell from
unknown causes ("shell pitting") are less common in ornate box
turtles than in other kinds of turtles.

Ectoparasites infesting T. ornata are larvae of chigger mites
(genus Tromhicula) and larvae of bot flies (Sarcophaga cvitu-
dinis). Ectoparasites usually have little adverse effect on the tur-
tles, although heavy infestations cause occasional injury or death.

Few natural enemies other than man are known; however most

Natural History of the Ornate Box Turtle 663

wild carnivores as well as opossums, large birds, and domestic
dogs and cats are suspect as predators. The incidence of predation
on eggs and small juveniles is far greater than on older juveniles
and adults. Adults of T. ornata occasionally attack smaller indi-
viduals.

Ornate box turtles are able to detect the presence of intruders,
by sight, from a distance of several hundred feet in open country;
apparently, intruders are not detected until seen. Defensive be-
havior is passive; the shell is closed tightly in response to painful
stimuli and, in some instances, at the sight of an intruder. Juveniles
usually void odoriferous fluid from the musk glands when handled
but adults do so only in response to pain or injury. The function of
the musk glands is unknown; possibly the odor of musk is a means
of sexual identification or stimulation. Although the musk is prob-
ably distasteful to predators, repellent odor alone seems to be of
doubtful value as a defense mechanism.

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1955. Further observations on the desert tortoise, Gophenis agassizi, of
California. Copeia, 1955(2) :113-118, 1 pi., 2 figs., 2 tables.
MnxER, M. R.

1948. The seasonal histological changes occurring in the ovary, corpus
luteum, and testis of the viviparous lizard, Xantusia vigilis. Univ.
California Publ. Zool., 47:197-224.
MrrsuKtTRi, K.

1895. How many times does the snapping turtle lay eggs in one season?
Zool. Mag. Tokyo, 7 (85): 143-147.
Neill, W. T.

1948a. Hibernation habits of amphibians and retpiles in Richmond Coimty,

Georgia. Herpetologica, 4(3) :107-114.
1948b. Odor of young box turtles. Copeia, 1948(2) :130.
Nichols, J. T.

1939a. Data on size, growth and age in the box turtle, Terrapene Carolina.
Copeia, 1939( 1) :14-20, 2 figs., 2 tables.

668 University of Kansas Publs., Mus. Nat. Hist.

1939b. Range and homing of individual box turtles. Copeia, 1939(3):
125-127.
NoRBis, K. S., and Zweifel, R. G.

1950. Observations on the habits of the ornate box turtle, Terrapene
omata (Agassiz). Nat. Hist. Misc., no. 58, 4 pp.
Odum, E, p., and Kuenzler, E. J.

1955. Measurement of territory and home range size in birds. Auk,
72(2): 128-137, 3 figs., 2 tables.
Oliver, J. A.

1955. The natural history of North American amphibians and reptiles.
Van Nostrand, xi -f 359 pp., 74 figs., 14 tables.

Ortenburger, a. I., and Freeman, B.

1930. Notes on some reptiles and amphibians from western Oklahoma.
Publ. Univ. Oklahoma Biol. Surv., 2(4):175-188, 2 maps.
Packard, R. L.

1956. The tree squirrels of Kansas. Univ. Kansas Mus. Nat. Hist,
Misc. Publ., no. 11, 67 pp., 2 pis., 10 figs.

Penn, G. H., Jr., and Pottharst, K. E.

1940. The reproduction and dormancy of Terrapene major in New Or-
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Peters, J. A.

1948. The box turtle as a host for dipterous parasites. Amer. Midi.
Nat., 40(2):472-474.

Rainey, D. G.

1953. Death of an ornate box turtle parasitized by dipterous larvae.
Herpetologica, 9(2):109.

RiSLEY, P. L.

1933. Observations on the natural history of the common musk turtle,
Stemotherus odoratus (Latreille). Pap. Michigan Acad. Sci., Arts
and Letters, 17:685-711, figs. 78-79, 1 table.

1938. Seasonal changes in the testis of the musk turde, Stemotherus
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RODECK, H. G.

1949. Notes on box turtles in Colorado. Copeia, 1949( l):32-34.

ROKOSKY, E. J.

1948. A bot-fly parasitic in box turdes. Nat. Hist. Misc., no. 32, 2 pp.

SCHNECK, J.

1886. Longevity of turtles. Amer. Nat., 20 ( 9 ) : 897.
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1937. Some materials to the problem of the reptilian post-embryonic
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1939. The body temperature of reptiles in natural surroundings.
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Smith, H. M., and James, L. F.

1958. The taxonomic significance of cloaca! bursae in turtles. Trans.
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Natural History of the Ornate Box Turtle 669

Smith, H. M., and Ramsey, L. W.

1952. A new turtle from Texas. Wasmann Joum. Biol., 10(l):45-54,
1 pi., 1 table.
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1954. An analysis of 100 years of grasshopper populations in Kansas
(1854-1954). Trans. Kansas Acad. Sci., 57(4):397-433, 1 pi.,
5 figs.
SxicacEL, L. F.

1950. Populations and home range relationships of the box turtle, Terra-
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1936. The occurrence of flints and extinct animals in pluvial deposits
near Clovis, New Mexico. Part III, — Geology and vertebrate
paleontology of the late Quaternary near Clovis, New Mexico.
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text.
Streceeb, J. K., Jr.

1908. The reptiles and batrachians of McLeiman County, Texas. Proc.
Biol. Soc. Washington, 21:69-84.
SmiFACE, H. A.

1908. First report on the economic features of turtles of Permsylvania.
Zool. Bull., Pennsylvania Dept. Agr., 6(4-5): 105-195, pis. 4-12,
16 figs., tables.
Taylor. W. E.

1895. The box tortoises of North America. Proc. U. S. Nat. Mus., 17
(1019):573-588, 7figs.
Wever, E. C, and Vebnon, J. A.

1956. Auditory responses in the common box turtle. Proc. Nat. Acad.
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WnxiAMS, E.

1950. Testudo cubensis and the evolution of western hemisphere tortoises.
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1948. Studies of the desert tortoise Gopherus agassizii. Ecol. Monogr.,
18:145-200, 25 figs., 4 tables.

Transmitted August 27, 1959.

PRINTED IN

THE STATE PRINTING PLANT

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I960

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INDEX TO VOLUME 11

New systematic names are in boldface type

IMOS. CO?^P. ZOBl]

LIBRARY

NOV 181960
UNIVERSITY

Abutilon theophrasti, 82

Acacia, 8

acacia, 508

acadicus, Aegolius, 207

Accipiter

cooperi, 53

cooperii, 178, 457

gentilis, 177. 456

stiiatus, 178

suttoni, 456-57

velox, 457
Acer, 409

Negundo, 536

negundo, 84
acorn woodpecker, 471
Acris gryllus, 91
Actitis macularia, 195, 462
acuta, Anas, 174, 455
adaptations in ornate box turtle, 650
aedon. Troglodytes, 251
Aegolius acadicus, 207
Aeronautes

saxatalis, 468

aestivalis, Hybopsis, 385-86, 419, 432-

33
aflBnis,

Aythya, 176, 455

Gambusia, 368, 371, 376-77, 385-
86, 388-90, 395, 397
agassizi, Gopherus, 557, 563, 573, 600,

611, 627, 630, 642-43
agassizii, Gopherus, 338, 340-42
agave, 448
Agelaius

megapotamus, 498

phoeniceus, 283
Agkistrodon, 318, contortrix, 124, 646
aglaiae, Pachyrhamphus, 473
Agonoderis obliqulus, 31
Aimophila

boucardi, 451, 509

cassini, 509

tenuirostris, 451, 509
Aix sponsa, 175
alba, Tyto, 200
albeola, Bucephala, 456
albescens, Certhia familiaris, 483
albicolljs, Zonotrichia, 314
albifrons,

Anser, 173

Catherpes mexicanus, 485

albiventris, Platypsaris aglaiae, 473

albonotatus, Buteo, 457

alcyon,

Megaceryle, 212
Megaceryle alcyon, 469
alder flycatcher, 77, 229
alexandri, Archilochus, 469
alfalfa, 85, 123, 138, 232, 241, 278,

295
alfreddugesi, Eutrombicula, 58
alfreddugesi, Trombicula, 54
alfreddugesi, Trombicula, 643-44
Algae, 361
algae, green, 353

alosoides, Hiodon, 383, 415, 432-33
alpestris, Eremophila, 232
alticola, Melospiza lincolnii, 512
Amazilia chalconota, 469
amblops, Hybopsis, 393
Ambrosia

artemisiifolia, 85

elatior, 536

triBda, 82
Ambystoma tigrinum, 88
Ameriurus melas, 377, 427
Ameiva, 17, 33, chrysolaema, 32
American

avocet, 462

bittern, 77, 172, 454

coot, 193, 461

eel, 376, 428

elm, 82, 241, 248, 259, 264, 271-73,
278, 289, 536

goldfinch, 505

hawk, rough-legged, 317

redstart, 77, 279, 497

rough-legged hawk, 317

toad, 78, 89, 165, 318

widgeon, 455

woodchuck, 77, 194
americana,

Certhia familiaris, 483

Fulica, 193, 461

Justicia, 353, 360-61

Mareca, 455

Parula, 494

Prunus, 82, 259

Recurvirostra, 462

Spiza, 295

Tilia, 85

Ulmus, 259

— Univ. Kansas Publ., Mus. Nat. Hist., Vol. 11, 1958-1960.

(671)

672 Unrersity of Kansas Publs., Mus. Nat. Hist.

americanum, Xanthoxj lum, 535
americanus,
Coccyzus, 198
Cocc>zus americanus, 464
Ulmus, 82, 409, 535
Ammodramus
bairdii, 508
perpallidus, 508
savannarum, 303
amoenissima, Polioptila caerulea, 449.

451, 488-89
amoenus, Carphophis, 117
Amphispiza

bilineata, 448, 509
grisea, 449, 509-10
opuntia, 451, 509-10
Amyda

mutica, 519, 523-24
muticus, 519
spinifera, 524
Anas

acuta, 174, 455
carolinensis, 175, 455
discors, 175, 455
platyrhynchos, 174, 454
septentiionalium, 455
strepera, 454
Andropogon, 409

gcrardi, 84-85, 353, 536
scoparius, 85, 353, 536
angelica, Leptotila verreauxi, 464
Anguilla bostoniensis, 376, 428
angustifolia, Typha, 158
anhinga, 453
Anhinga anhinga, 453
anhinga, Anhinga, 453
ani, groove-billed, 448, 465
annularis, Pomaxis, 380, 385-86, 388-

89, 391, 429, 432-33
anomalum, Campostoma, 370,375,379
385-86, 388-89, 395-96, 432-33 '
Anser albifrons, 173
anthinus, Passerculus sandwichensis,

508
Anthus

pacificus, 490
rubcscens, 490
Antrostomus macromystax, 467
annus, Ilelianthus, 82
Aphelocoma
couchii, 479
cyanotis, 479
potasina, 480
aphrasta,

Erophilia alpestris, 477
Otocoris alpestris, 477
Aplodinntus grunniens, 383, 385-86

388-89, 430, 432-33
ApocyniuTi cannabinum, 85
aquaticus, Scalopus, 30, 133, 156
aquilonaris, Geophis, 329

arachnoides. Pyxis, 588

Araneida, 621

arborea, Spizella, 307

arcansanum, Etheostoma zonale, 384,

393
Arciiilochus
alexandri, 469
colubris, 212, 469
arctiid caterpillar, 620
Arctiidae, 621
Ardea

herodias, 170, 453
treganzai, 453
wardi, 453
arenacea, Spizella pusilla, 511
argutus, Rubus, 84
aries, Ovis, 75
Aristida oligantha, 84
arizonae,

Caprimulgus vociferus, 467
Peucedramus taeniatus, 494
Spizella passerina, 511
Arkansas River shiner, 368
Anna, 31

Armadillidium vulgare, 621
amiadillo, 52, 58
Arremonops rufivirgata, 506
Artemesia ludoviciana, 536
arteinesiifolia. Ambrosia, 85
artemisae, Molothrus ater, 500
Asclepias, 83
ash, 293, 353, 409

northern prickly, 535
prickly, 256
ash-throated flycatcher, 474
asiatica, Zenaida asiatica, 463
Asio otus, 206
asio, Otus, 200
aspen, 449, 486
Aster, 82, 536
aster, 82, 84, 536
ater,

Molothrus, 287
Molothrus ater, 500
Atlapetes dilitus, 505
atlapetes, rufous-capped, 449, 452,

505
atratus, Coragyps, 456
atricapilla, Virco, 491
atricapillus. Parus, 241
atricristatus, Parus atricristatus, 448-

49, 481
atrogularis, Spizella atrogularis, 512
atromaculatus,

Semotiius, 414, 419, 432-33
Semotilus atromaculatus, 419
attenuatus, Ophisaurus, 53, 104
auduboni.

Dendroica auduboni, 495
Hylocichla guttata, 487-88
Audubon's warbler, 495

Index to Volume 11

673

aura,

Cathartes, 176

Cathartes aura, 456
auratus,

Carassius, 419, 432-33

Colaptes, 213
aureolum, Moxostoma, 418, 432-33
auricollis, Icteria virens, 496
aurifroDS, Centurus aurifrons, 470
Auriparus

flaviceps, 481

ornatus, 481
aurocapillus, Seiurus, 275
autumnalis, Dendrocygna, 454
avocet, American, 462
awnless

brome, 74

brome grass, 83, 190, 306
Aythya

affinis, 176, 455

coUaris, 176
Aztec thrush, 452, 487

badger, 646-47

bairdi, Dryobates scalaris, 472

bairdii, Ammodramus, 508

Baird's sparrow, 508

bald eagle, 77, 187, 317

baldwini, Vemonia, 83

Baltimore oriole, 77, 285

banded darter, 384

band-tailed pigeon, 449, 451, 463

bank swallow. 77, 233

Baris, 31

bark beetle, 221

barley, 85, 295

bam,

owl, 77, 200, 317, 465

swallow, 81, 85, 234, 322. 478
barred owl, 77, 203, 319
Bartramia longicauda, 195
bass, 354, 584

black, 395

largemouth, 380, 429, 437

spotted, 380

white, 384, 428, 437
basswood, 465, 469
bat, red, 79, 134
beaver, 317

becard, rose-throated, 473
beefcattle, 535
beetle, 661

bark, 221

carabid, 620

chrysomelid, 230, 266, 293

groimd, 31

ladybird, 31

scarabaeid, 241, 620

tiger, 31
Bell vireo, 81, 270-71, 321
bellii,

Chrysemys picta, 557

Vireo, 270

Bell's vireo, 448-49, 492

belted kingfisher, 77, 212, 317, 469

bendirei, Faico columbarius, 458

berlandieri,

Gopherus, 338-41, 628
Thryothorus ludovicianus, 484
Bevidck's wren, 484
bicolor,

Geophis, 832

Iridoprocne, 478

Parus, 245
Big Blue River Basin, 404
big bluestem, 84-85, 141, 153, 163,

282, 295
bigeye shiner, 365
bigmouth

buffalo, 416

buffalo fish, 364
biguttata, Hybopsis, 393
bifineata, Amphispiza bilineata, 448,

509
bindweed, 106, 304
Birds from Coahviila, Mexico, 445
bird's nest fungus, 622
bittern, American, 77, 172, 454
black

bass, 395

buffalo, 363, 416

bullhead, 77, 87, 377, 390, 427

cherry, 536

crappie, 381, 429

oak, 82, 204, 271, 293

phoebe, 451, 474

rat snake, 54, 79, 121, 238, 261,
321-22

snake, 51

vulture, 450, 456

walnut, 271, 354, 535

willow, 536
black and white warbler, 77, 272, 493
black-banded topminnow, 376
black-bellied tree duck, 454
blackberrv, 71, 84-85, 134, 151, 178,

230, 243, 254, 256, 268, 278, 301,

320
black-billed cuckoo, 77, 199, 465
blackbird, 392

Brewer's, 499

red-winged, 77, 498

rustv, 286

yellow-headed, 77, 282, 498
Blackburnian warbler, 77, 275
black-capped

chickadee, 81, 221, 241, 250, 265,
319

vireo, 491
black-chinned

hummingbird, 469

sparrow, 512
black-crested titmouse, 448-49, 451,

481

674

University of Kansas Publs., Mus. Nat. Hist.

black-crowned night heron, 77, 172,

454
black-eared bushtit, 451, 482
black-fin minnow, 420
blackhead minnow, 426
black-headed grosbeak, 502
blackjack oak, 19
blackpoll warbler, 77, 275
black-tailed gnatcatcher, 489
black-throated

gray warbler, 495

green warbler, 77, 275, 496

sparrow, 448-49, 451, 509-10
blandingi, Emydoidea, 532, 554, 653
Blarina, 133, brevicauda, 30, 130
blennius, Notropis, 365, 385-86, 394-

95, 423
blennoides, Etheostoma, 393
blue

goose, 77, 173

grosbeak, 77, 294, 321, 451, 503

heron, great, 76-77, 170, 453

jay, 70, 81, 235, 318-19

sucker, 416

vervain, 536
bluebird, 231, 242, 267, 319

eastern, 81, 260, 488

mountain, 488

western, 488
bluegill, 382. 429, 437, 584
bluegrass, 83, 85, 123, 140, 155, 195,

296
blue-grav gnatcatcher, 81, 263, 448-

49, 451, 488-89
bluestem, 74, 196, 305, 390, 409, 536

big, 84-85, 141, 153, 163, 282, 295

grass, 281

little, 85, 141, 295

prairie, 109, 118, 161
blue-throated hummingbird, 469
blue-winged teal, 77, 175, 455, 618
bluntface shiner, 367
bluntnose minnow, 373, 426
boat-tailed grackle, 499
bob-white, 80, 190, 319, 321, 448,

459, 618
Boleosoma nigrum, 430
bombifrons, Scaphiopus, 88
Bombycilla cedrorum, 265, 490
bonesct, false, 82

bony shell of ornate box turtle, 586
boops, Notropis, 365, 368, 370, 372,

385-86, 388-89, 393-94, 396-97
borealis,

Lasiunis, 134

Nuttallornis. 232, 476
boreus, Myiarchus crinitus, 473
Bos taurus, 75

bostoniensis, Anguilla, 376, 428
hot fly. 643, 645-46, 662
Botaurus lentiginosus, 172, 454
boucardi, Aimophila ruficeps, 451, 509

Bouteloua curtipendula, 536
box turtle, 51, 54, 58, 538
eastern, 77, 99
ornate. 322, 531
western, 78, 99
boxelder, 84, 536

brachidactylus, Geothlypis trichas, 496
brachyrhynchos, Corvus, 648
Branta

canadensis, 172
leucopareia, 454
breeding, six-lined racerunner, 32
brevicauda, Blarina, 30, 130
breviceps, Lepomis raegalotis, 382,

393
breweri, Spizella breweri, 511
Brewer's

blackbird, 499
sparrow, 511
brewsteri^ Empidonax traillii, 475
broad-tailed hummingbird, 469
broad-winged hawk, 53, 77, 184-186,

457
brome, 161

grass, 49, 123. 128, 138-39, 141,
143, 168, 224, 278, 281, 287,
296, 304

grass, awnless, 74, 83, 190, 306
Bromus

inermis, 83

japonicus, 159, 535-36
bronzed grackle, 77, 286
brook silversides, 378
brooksi, Passerculus sandwichensis,

508
brown

creeper, 81, 221, 245, 249, 265,
449, 483

snake, 78, 114

thrasher, 81, 256

towhee, 507
brown-headed cowbird, 500
brown-throated wren, 451, 483
brunncscens, Passerculus sandwichen-
sis, 508
bubalus, Ictiobus. 363-64, 374, 385-

86, 388, 416, 432-33
Bubo

mayensis, 466

palicscens, 466

virgianus, 201
Bucephala albcola, 456
buchanani, Notropis, 366, 385-86, 388-

89, 397. 423, 432-33
buckbrush, 535-36
buffalo, 437, 452

bigmouth, 364, 416

black. 363, 416

fish, 354

smallmouth, 363-64, 416, 435
buffalo-bur. 536
buft-bellicd hummingbird, 469

Index to Volume 11

675

buff-breasted flycatcher, 449-450, 476

bufilehead, 456

Bufo

terrestris, 89
woodhousei, 90
bullfrog, 78, 96, 164, 171, 318, 537
bullhead, 437

black, 77, 87, 377, 390, 427

yellow, 377, 427
bullsnake, 79, 122, 648
bulockii. Icterus bulockii, 499
Bullock's oriole, 499
bunting,

indigo, 81, 85, 294, 321, 503

lark, 507

painted, 77, 295, 504

varied, 503
bursarius, Geomys, 88, 138
burrowing owl, 467
burrows, six-lined racerunner, 28
bush, creosote, 540
bushtit, black-eared, 451, 482
Buteo

albonotatus, 457

borealis, 457

fuertesi. 457

jamaicensis, 180, 648

lagopus, 187

lineatus, 183

nitidus, 458

platypterus, 58, 186, 457

regalis, 458

swainsoni, 457
Butoroides virescens, 171, 454

caballus, Equus, 75
cacti, 448

columnar, 8
cactopliilus, Dendrocopos scalaris,

450, 472
cactus wren, 449, 451, 484
caerulea,

Guiraca, 294

Guiraca caerulea, 503

Polioptila, 263

Polioptila caerulea caerulea, 448,
488
caerulescens, Chen, 173
cahooni, Troglodytes brunneicollis, 451
Calamospiza melanocorys, 507
Calendra parvulus, 621
calendula,

Regulus, 265

Regulus calendula, 489
calif ornianus, Geococcyx, 465
calif ornicum, Glaucidium gnoma, 466
calligaster, Lampropeltis, 51, 123
CaUipepla

castanogastris, 459-60

palUda, 450, 459-60
squamata, 451, 459-60
Calothorax lucifer, 468

calvatus, Trionyx muticus, 519-20
Campostoma

anomalum, 370, 375, 379, 385-86,

388-89, 395-96, 432-33
plumbeum, 426, 431
camurus, Notropis, 367, 370, 373, 379,

385-86, 388-89, 393, 396-97
Campylorhynchus
couesi, 451, 484
guttatus, 449, 485
Canada

goose, 77, 172
warbler, 77, 279
canadense,

Stizostedion, 430, 432-33
Teucrium, 83, 212
canadensis,
Branta, 172
Cercis, 82
Erigeron, 82
Grus, 192
Sitta, 249
Wilsonia, 279
cancellatus, Geophis, 333-34
canicaudus, Richmondena cardinalis,

502
Canis

familiar is, 75
latrans, 162, 646
cannabinum, Apocynum, 85
canon wren, 485
Cantharidae, 621
Capella gallinago, 195
Caprimulgidae, 452
Caprimulgus
arizonae, 467
carolinensis, 208
vociferus, 209
caprodes, Percina, 385-86, 388, 391
carabid beetle, 620
Carabidae, 621
caracara, 458
Caracara cheriway, 458
Carassius auratus, 419, 432-33
carbonaria, Percina caprodes, 378
cardinal, 81, 85, 223, 290, 303, 307,

318, 502
cardinalis, Richmondena, 290
Carex, 536

carinatum, Moxostoma, 364, 393
carolensis, Sitta, 248
Carolina,

Porzana, 193, 461

Terrapene, 99, 449, 532-35, 539,
543, 545, 548, 551, 553-54, 557-
59, 561-63, 569, 572, 585, 595-
96, 611-12, 616, 622-23, 626-
28, 630, 634, 636-38, 640, 643,
646-48, 650-54, 656
Terrapene Carolina, 654-55
Carohna wren, 81, 252, 319, 448, 484

676

University of Kansas Publs., Mus. Nat. Hist.

carolinae,

Cottus, 393

Vireo huttoni, 448, 451, 492
carolinensis.

Anas, 175, 455

Capriinulgus, 208

Dumetclla, 256

Sciunis, 136

Zenaidura macroura, 463
carolinus,

Centurus, 214

Euphagus, 286
carp, 365, 388, 395-96, 418, 435, 437
Carphophis amoenus, 117
carpio,

Carpiodes, 386, 388, 431-33

Carpiodes carpio, 363, 416

Cyprinus, 365, 385-86, 388, 418,
431-33
Carpiodes

carpio, 363, 386, 388, 416, 431-33

forbesi, 416-17, 432-33. 438

velifer, 363, 394, 417
Carpodacus

cassinii, 504

centralis, 505

frontalis, 505

nigrescens, 505

potosinus, 504

purpureas, 298
carpsucker, 437

highfin. 363, 417

plains, 416

river, 363, 416, 418, 435
Casmerodius egretta, 454
caspicus, Podiceps, 453
Cassia, 8
Cassidix

mexicanus, 500

nionsoni, 500

prosopidicola, 499
cassini, Aimophila, 509
cassinii,

Carpodacus, 504

Vireo solitarius, 493
Cassin's

finch, 504

kingbird. 473

sparrow, 448, 509
castanogastris, Callipepla squamata,

448, 459-60
cat,

domestic, 648, 663

house. 317
catbird, 77, 256
catclaw, 481. 485, 491, 504
catcnatus, Sistnirus, 51
catenifer, Pitiiophis, 122, 648
caterpillar. 617, 618, 620

arctiid, 620

noctuid, 620, 622

catesbeiana, Rana, 96, 537, 618
catfish, 354

channel, 377, 427, 435, 437

flathead. 377, 428, 435, 437

madtom, 395
Cathartes

aura. 176, 456

teter. 456
Catherpes albifrons, 485
Catosomus commersonnii, 414, 418,

432-33
cattait 84, 93, 158, 165, 316
caudacutus, Passerherbulus, 304
cave swallow, 448, 478
ceanothus, 506
cedar,

red, 206, 249

waxwing, 77, 265, 490
cedrorum, Bombycilla, 265, 490
celata, Vemiivora. 273, 493
celsum, Toxostoma curvirostre, 451,

486
Celtis occidentalis, 536
centralis, Carpodacus mexicanus, 505
Centurus

aurifrons, 470

carolinus, 214

incanescens, 470-71
cepedianum, Dorosoma, 363, 385-86,

388, 415, 432-33
Cerambycidae, 621
Cercis canadensis, 82
Certhia

albescens, 483

americana, 483

familiaris, 249

montana, 483
Chaenobryttus giilosus, 384
Chaetura pelagica, 211
chalconota, Amazilia yucatanensis,

469
Chalcis, 31

chalybea, Progne chalybea, 479
chalybeus, Geophis, 333
channel

catfish, 377, 427, 435, 437

darter, 379
Chara, 353, 361
Charadrius vociferus. 194, 461
chat,

long-tailed. 77. 278, 321

yellow-breasted, 279, 496
Chautauqua, Cowley and Elk Comi-
ties, present flora, 353
Chelonidae, 552

Chelydra seri^cntina, 98, 559, 611
Chen

caerulescens, 173

hyperborea, 173
Chenopodium, 82
chcriway. Caracara, 458

Index to Volume 11

677

cherry,

black, 536

wild, 270
chess, Japanese, 159, 536
chestnut, 125, 271

oak, 203, 248, 293
chickadee, 189, 242, 245, 249, 318,
320 322

black-capped, 81, 221, 241, 250,
265, 319

Mexican, 449, 481
chicken, 179, domestic, 162
chigger, 53, 54, 58, mite, 643-44,

662
chihuahuae, Colaptes cafer, 470
chimney swift, 77, 211
chinquapin oak, 206-07, 263
chipping sparrow, 72, 77, 85, 308, 511
Chloroceryle

hachisukai, 469

tentrionalis, 470
chlorosomum, Etheostoma, 393
Chlorura chlorura, 506
chlorura, Chlorura, 506
Chondestes

grammacus, 304

strigatus, 508
Chordeiles

howelh, 468

minor, 210

texensis, 468
chorus frog, 78, 93-94, 165, 318
Chrysemys, 569, 571, 628-29

beUii, 557

picta, 101, 552, 554, 559, 585, 611,
627, 643
Chrosomus erythrogaster, 419. 431-

433
chrysolaema, Ameiva, 32
chrysomelid beetle, 230, 266, 293
Chrysomelidae, 621
chrysoparia, Dendroica, 496
chrysops, Roccus, 384, 428, 432-33
chub,

creek, 419

silver, 419

speckled, 365, 419
chuck-will's widow, 77, 208, 322
cicada. 618
cineraceus,

Otus asio. 465

Regulus calendula, 489-90
cinerascens, Myiarchus cinerascens,

474
cinereoargenteus, Urocyon, 164
cinnamon teal, 455
Circus cyaneus, 53, 188, 458
ciris, Passerina, 295
Cirsium, 82

Cistophorus platensis, 255
cistudinus, Sarcophaga, 643, 645-46,

662

clay-colored sparrow, 77, 308, 511
clemenciae, Lampornis clemenciae,

469
Clemmys, 589, insculpta, 653
cliff swallow. 77, 235, 478
climate,

of Chautauqua, Cowley, and Elk
counties, 351

of Big Blue Basin, 416
sweet clover, 191
clypeata. Spatula, 175, 455
Cnemidophorus, 17, 22, 33, 56

gularis, 18

inomatus, 43

octolineatus, 17

perplexus, 20, 38, 43

sacki, 17, 20, 30

sexlineatus, 17-18, 21, 32, 106, 554

tessellatus, 20, 31, 38

tigris, 17, 20-21, 28, 31, 38
coachwhip, 51

Coahuila, Mexico, birds from, 445
coahuila, Terrapene, 532, 654
Coccyzus

americanus, 198, 464

erythropthalmus, 199, 465

occidentalis, 464
cockroach, 31

coeruleum, Etheostoma, 379
coffee tree, 271. Kentucky, 284
Colaptes

auratus, 213

chihuahuae, 470

collaris, 470

nanus, 470
Coleoptera, 621-22
Colima warbler, 494
Colinus

texanus, 448, 459

virginianus, 190
collared Hzard, 25, 51, 67, 76, 78,

102, 187, 318, 321. 601, 616-17
collaris,

Aythya, 176

Colaptes cafer, 470

Crotaphytus, 25, 51, 601, 617
color and markings of ornate box

turtle, 593
colossus, Eumolops, 621
Coluber, 318

constrictor, 51, 58, 118

flagellum, 51, 58
colubris, Archilochus, 212, 469
Columba

fasciata, 463

livia, 75
Colirmbigallina passerina, 212
columnar cacti, 8
commersonnii, Catostomus, 414, 418,

432-433
common

egret, 454

678

Univ'ersity of Kansas Publs., Mus. Nat. Hist.

common — Concluded

garter snake, 79, 114, 322

nighthawk, 468

raven, 480

shiner, 421

water snake, 77, 113
compositus Troglodytes brunneicoUis,

451, 483
confinus, Poecetes gramineus, 51
constrictor, Coluber, 51, 58, 118
Contopus

pertinax, 476

veliei, 476

wens, 230, 476
contortrbc, Agkistrodon, 124, 646
Cooper hawk, 77, 178, 191
Cooper's hawk, 53, 319, 457
cooperi,

Accipiter, 53

MyTiarchus tyrannulus, 474

Piranga rubra, 449, 451, 501
cooperii, Accipiter, 178, 457
coot, 77, American, 193, 461
copelandi, Percina, 379, 385-86, 388-

89 393-94
copperhead, 79, 124, 129, 321-22, 646
Coragyps atratus, 456
coralberry, 146-47, 153, 155, 161, 278
corn, 159, 164, 191, 407, 535, snake, 52
Comus

Drummondi, 536

drummondii, 82, 287
comutus, Notropis, 432-33
coronata, Dendroica, 274, 495
Corvus

brachyrhynchos, 238, 646, 648

cr>ptoleucus, 480, 646

sinuatus, 480
cotton rat, 160, 163, 206-07, 321, his-
pid, 80, 149, 154
cottontail, 54, 69, 71, 79, 163, 178,

181, 202, 243, 321-22, eastern, 134,

622
Cottonwood, 19, 214, 285, 353, 409,

498, 502, 535-36
cottonwood-elm association, 199
Cottus carolinae, 393
couchii, Aphelocoma ultramarina, 479
Coues' flycatcher, 476
couesi, Campvlorhynchus brunneicap-

illum, 451,484
cow, 392
cowbird, 81, 269, 287

brown-headed, 500
coyote, 52-53, 58, 67, 77, 159, 162,

169, 322, 646-47, 649, melon, 617
crab apple, 82, 270, 278, 290, 294, 301
crabgrass, 85
cragini, Ethoostoma, 384
Cragin's darter, 384
crane, sandhill, 77, 192, 461

crappie, 437

black, 381, 429

white, 380, 429
Crataegus mollis, 82
crayfish, 622
creek chub, 419
creel census of fishes of Big Blue Basin,

435
creeper, 249

brown, 81, 221, 245, 249, 265, 449,
483

Virginia, 273
Crematogaster, 31, 96
creosote, 462, bush, 540
crested flycatcher, 80, 223

great, 473

Wied's, 450, 474
cricket frog, 78, 91, 94, 164, 171, 196,

318
Crissal thrasher, 451, 487
crissalis, Vermivora, 494
crinitus, Myriarchus, 223, 473
cristata, Cyanocitta, 235
Crotalus horridus, 52, 128
Crotophaga sulcirostris, 465
Crotaphvtus collaris, 25, 51, 102, 601,

617
crow, 81, 202, 238, 319, 646, 648
Crustacea, 621

cryptoleucus, Corvus, 480, 646
crysoleucas,

Notemigonus, 372, 385

Notropis, 386
Cryptotis parva, 132
cryptus, Tnr>'omanes bewickii, 484

CUCKOO,

black-billed, 77, 199, 465

yellow-billed, 80, 198-99, 448, 464
cucuUatus, Icterus cucullatus, 499
Cucurbita foetidissima, 617
Cuora, 532
Curculionidae, 621
curlew, long-billed, 462
currucoidcs, Sialia, 488
curtipendula, Bouteloua, 536
curve-billed thrasher, 448, 451, 486
cutgrass, rice, 141
cutworm, 31

cvanea, Passerina, 294, 503
cyanellus, Lepomis, 87, 381, 385-86,

388-92, 394-97, 421-22, 429, 431-

33
cyancus. Circus, 53, 188, 458, 648
cyanocephalus, Euphagus, 499
Cyanocitta

cristata, 235

macrolopha, 479
c van Otis, Aphelocoma coerulescens,

' 479
Cyathus striatiis, 622
Cyclagras, 8

Index to Volume 11

679

Cyclemys, 532

Cycleptus elongatus, 393, 416

cycles of activity and temperature re-
lationship, six-lined racerunner, 20

cypress, 465

cyprinella, Ictiobus, 385-86, 388, 416,
432-33

Cyprinus carpio, 365, 385-86, 388, 418,
431-33

Cyrtonyx

mearnsi, 461
montezumae, 461

dace, southern redbelly, 419
dactyloides, Tripsacum, 353
Dama virginiana, 168
dandelion, 212
darter,

banded, 384

channel, 379

Cragin's, 384

johnny, 430

orangethroat, 379, 430

redfin, 384

slenderhead, 378
deep-bodied sucker, 388-89, 395-96
deer, 163, white-tailed, 77, 168, 322
deer mouse, 79, 143, 321

prairie, 145

woodland, 145
defense in the ornate box turtle, 648
DeKay snake, 616
dekayi, Storeria, 114, 616
delawarensis, Larus, 463
deliciosus, Notropis, 376, 386, 395,
404, 423, 431-32, 438

Notropis deliciosus, 423, 431
deltoides, Populus, 409, 535
Dendrocopos

cactophilus, 450, 472

giraudi, 449, 472

icastus, 450, 471

intermedius, 448, 472

pubescens, 220

symplectus, 448, 472

villosus, 218
Dendrocygna autumnalis, 454
Dendroica

auduboni, 495

chrysoparia, 496

coronata, 274, 495

fusca, 27*5

memorabihs, 495

morcomi, 495

nigrescens, 495

nigrifrons, 495

occidentalis, 496

petechia, 274

striata, 275

townsendi, 495

virens, 275, 496
Dermochelydae, 552

desert tortoise, 600, 611
dextra, Piranga flava, 501
Diabotrica 12-punctata, 621
Diadophis punctatus, 116
diamond-backed terrapin, 552
diaphora, Erophilia alpestris, 477
dickcissel, 68, 81, 85, 295, 321
Didelphis marsupialis, 129, 648
diet of ornate box turtle, 617
differential grasshopper, 622
difFerentiahs, Melanoplus, 620-21
difBcilis, Empidonax, 476
diffusa, Jussiaea, 353, 361
Digitaria sanguinalis, 85
dilutus, Atlapetes pileatus, 505
dioica, Gymnocladus, 284
Dionda nubila, 393
Diplopoda, 621
Diptera, 621
discolor,

Leptodeira, 3, 4

Tantalophis, 4-7
discors.

Anas, 175

Anas discors, 455
distribution of known breeding birds

of Coahuila, 447
dog, domestic, 646, 663
dogbane, 85
dogwood, 82, 84-85, 113, 209, 256,

263, 268, 270, 287, 316, 536
doliata, Lampropeltis, 124
domestic

cat, 648, 663

chicken, 162

dog, 646, 663

pigeon, 75
domesticus,

Felis, 75

Passer, 280

Passer domesticus, 497
Dorosoma cepedianum, 363, 385-86,

388, 415, 432-33
dorsale, Toxostoma dorsale, 451, 487
Douglas fir, 487, 496
dove,

ground, 464

Inca, 464

mourning, 80, 196, 231, 319-20,
463

white-fronted, 464

white-winged, 463
downy woodpecker, 69, 80, 215, 219-

20, 245, 250, 265, 319
Drepanoides, 8

drum, freshwater, 383, 430, 435
Drummondi, Comus, 536
dnunmondii, Comus, 82, 287
Dryobates bairdi, 472
duck,

black-bellied tree, 454

lesser scaup, 176

680

University of Kansas Publs., Mus. Nat. Hist.

duck, — Concluded

ring-necked, 77, 176

wood, 77, 175
duckv^'eed, 353
Duellman, William E., Systematic

status of the colubrid snake, Lep-

todeira discolor Giinther, 3
dugesii, Geophis, 333-34
Dumetella carolinensis, 256
dumosum, Toxostoma dorsale, 451,

487
duquesnii, Moxostoma, 393
duquesnei, Moxostoma niacrolepido-

tum, 418
dwarf oak, 268, 271
dysleptus, Parus atricristatus, 451, 481

eagle, bald, 77, 187, 317

eared grebe, 453

earless lizard, 52

earth snake, smooth, 77, 115

G3.stGrn

bluebird, 81, 260, 488

box turtle, 77, 99

cottontail, 134, 622

goldfinch, 81, 299

kingbird, 77, 222

meadowlark, 81, 281, 498

mole, 133

pewee, wood, 80, 230, 476

phoebe, 474

ring-necked snake, 116

woodrat, 67, 80, 151
economic importance of ornate box tur-
tle, 534
ectoparasites of ornate box turtle, 643
eel, American, 376, 428
egret, common, 454
egretta, Casmerodius albus, 454
eidos, Parus sclateri, 480
Elaphe, 318

guttata, 52

obsoleta, 52, 54, 121
elatior. Ambrosia artemisiifolia, 536
elderberry, 256
elegans,

Geochelone, 552

Pseudemys scripta, 571

Thamnophis, 548
elf owl, 450, 467
elm, 83, 123, 125, 134, 136, 161. 197,

199, 201, 209, 214, 218-19, 224-25,

235, 238. 242-43, 250, 263, 265,

267-68. 270, 277. 279, 289, 293-

94, 296, 299, 308, 314, 353, 409,

469, 535, American, 82, 241, 248,

259, 264, 271-73, 278, 289, 536
elm-hackberry fringe-forest, 353
elongatus, Cycleptus, 393, 416
embryonic development of ornate box

turtle, 560

Empidonax

brcwsteri, 475

difficilis, 476

flaviventris, 229

griseus, 475

hammondii, 475

liellmayri, 476

immemoratus, 476

minimum, 230

minimus, 475

occidentalis, 476

pygmaeus, 476

salvini, 476

traillii, 229

trepidus, 475

wrightii, 475
Emydoidca, 588, blandingi, 532, 554,

653
Emyidae, 532, 588
Emys, 532, 588, orbicularis, 642-43
English sparrow, 81, 85, 280
enthvmia, Eremophila alpestris, 477
Epeira, 621
Equus caballus, 75
Eremophila

alpestris, 232

enthymia, 477
eremophilus, Thryomanes bewickii,

484
ericrypta, Melospiza georgiana, 513
Erigeron canadensis, 82
Erolia

melanotos, 462

minutilla, 462
Erophilia

aphrasta, 477

diaphora, 477
erythrocephalus, Melanerpcs, 217
ervthrogaster,
'Chrosomus, 419, 431-33

Hirundo rustica, 478
( r\'throphthalmus,
' Coccvzus, 199, 465

Pipilo, 300
erythrorhynchos, Pelecanthus, 170, 453
erythrurum, Moxostoma. 364, 385-86,

388, 391, 393, 396, 418
Essox lucius, 437
Etheostoma

arcansanum, 384, 393

blcnnoides, 393

chlorosomum, 393

cooruleum, 379

cragini, 384

gracile, 393

lineolatum, 393

nigrum, .393, 430, 432-33

piHchellum, 379, 430

punctulatum, 393

saxatile, 393

Index to Volume 11

681

Etheostoma — Concluded

spectabile, 370, 375, 385-86, 388-
89, 391, 395-96, 432-33

whipplii, 384, 393
Eugenes fulgens, 469
Euglena rubra, 353
Eumeces, 319

fasciatus, 53, 109, 113, 584, 623

multivirgatus, 52

obsoletus, 53, 110, 113, 623

septentrionalis, 113
Eumolops colossus, 621
Eupatcrium altissimum, 82
eupatoroides, Kuhnia, 82
Euphagus

carolinus, 286

cyanocephalus, 499
Euphorbia, 82
Eupoda montana, 462
eurhyncha, Guiraca caerulea, 451, 503
Eutrombicula alfreddugesi, 58
excubitorides, Lanius ludovicianus,

490-91
exilis, Noturus, 393
Extrarius

sesquialis, 419

tetranemus, 419

Falco

bendirei, 458
mexicanus, 189, 458
sparverius, 53, 189, 459
falcon, prairie, 77, 189, 458
false boneset, 82
familiaris,
Canis, 75
Certhia, 245
fasciata, Columba fasciata, 463
fasciatus, Eumeces, 53, 109, 113, 584,

623
fathead minnow, 77, 86, 374, 426
Felis domesticus, 75
ferox, Trionyx, 520, 523
ferruginous hawk, 458
fertility and prenatal mortality of or-
nate box turtle, 564
field sparrow, 68-69, 82, 85, 231, 308.

319, 511
finch,

Cassin's, 504
house, 504
purple, 77, 298
fir, Douglas, 487, 496
fir-pine-aspen association, 469
fish,

bigmouth buffalo, 364
bufFalo, 354
smallmouth buffalo, 364
Fishes of the Big Blue River Basin,

Kansas, 403
Fishes of Chautauqua, Cowley, and
Elk counties, Kansas, 347

fissicornis, Prionus, 621
Fitch, Henry S.,

Home ranges, territories, and sea-
sonal movements of vertebrates
of the Natural History Reserva-
tion, 65

Natural history of the six-lined
racerimner ( Cnemidophorus sex-
lineatus), 13
five-lined skink, 67, 78, 109, 111, 584
flagellum.

Coluber, 51, 58

Masticophis, 51
flammeolus, Otus fiammeolus, 466
flammeus, Pryocephalus rubinus, 451.

477
flammulated owl, 466
flathead catfish, 377, 428, 435, 437
flat-headed snake, slender, 124
flavescens, Perca, 430
flaviceps, Auriparus flaviceps, 481
flavifrons, Vireo, 271, 492
flavipes, Totanus, 195, 462
flaviventris, Empidonax, 229
flavomarginatus, Gopherus, 337
flavus, Noturus, 383, 428, 432-33
flicker, 177, 215

red-shafted, 470

yellow-shafted, 80, 213
floridana, Neotoma, 151
floridanus, Sylvilagvis, 54, 134, 622

fly,

bot, 643, 645-46, 662

robber, 617
flycatcher,

alder, 77, 229

ash-throated, 474

buflF-breasted, 449-50, 476

Coues', 476

crested, 80, 223

gray, 475

great crested, 473

Hammond's, 475

least, 77, 230, 475

olive-sided, 77, 232, 476

pine, 449, 475

scissor-tailed, 77, 222, 473, 475

Traill, 475

vermilion, 448-49, 451, 477

western, 476

Wied's crested, 450, 474

Wright's, 475

yellow-bellied, 77, 229
foetidissima, Cucurbita, 617
foliosus, Potamogeton, 353, 361, 379
food habits, six-lined racerunner, 31
forbesi, Carpiodes, 416-17, 432-33, 438
forficata, Muscivora, 222, 473
Formica, 31
Fonnicidae, 621
formicivorus, Melanerpes formicivorus,

471

682

University of Kansas Publs., Mus. Nat. Hist.

formosus, Oporornis, 276
fossils of Terrapene, 534
fox, 159

gray, 77, 164

red, 77, 163, 322

grape, 270

sparrow, 77, 315

squirrel, 69, 79, 137, 246, 321-22
foxtail, 85, 141, 153, 160, 296, 315-16
fragrant sumac, 301
Franklin gull, 77, 196, 317
franklinii, Spermophilus, 136
Franklin's ground squirrel, 77, 136
Fraxinus, 409
freckled madtom, 383
frenata, Mu stela, 166
freshwater drum, 383, 430, 435
frog,

chorus, 78, 93-94, 165, 318

cricket, 78, 91, 94, 164, 171, 196,
318

leopard, 78, 97, 164, 171, 318, 537
frontalis,

Carpodacus mexicanus, 505

Notropis comutus, 421, 431
fuertesi, Buteo jamaicensis, 457
fulgens, Eugenes fulgens, 469
Fulica americana, 193, 461
fulva,

Siaha sialis, 488

Vulpes, 163
Fundulus,

kansae, 376, 385-86, 394-95, 428,
432-33

notatus, 376, 385-86, 388-92, 396-
97

zebrinus, 376
fungus, bird's nest, 622
fusca, Dendroica, 275
fuscescens, Hylocichla, 260

gadwell, 454
gaigei,

Piplio erythrophthalmus, 506

Pipilo maculatus, 507
Galapagos tortoise, 342
galbula. Icterus, 285
gaUinago, Capella, 195
gambeli, Lanius ludovicianus, 491
gambelii, Zonotrichia leucophrvs, 512
Gambusia affinus, 368, 371, 376-77,

385-86, 388-90, 395, 397
gar, 395, 437

longnose, 362, 415

shortnose, 363, 415

spotted, 383
garfish, 354
garter snake, 321, 548, common, 79,

114, 322
Caslroplnyne olivacea, 95
Gastropoda, 621
gentilis, Accipiter, 177, 456

Geochelone, 342

elegans, 552
Geococcyx californianus, 465
geology of Big Blue Basin, 405
Geomys bursarius, 88, 138
Geophis

aquilonaris, 329

bicolor, 332

cancellatus, 333-34

chalybeus, 33

dugesii, 333-34

latifrontalis, 332

semidoliatus, 333
georgiana, Melospiza, 316
Geothlypis

brachidactylus, 496

nelsoni, 496

cccidentalis, 496

trichas, 277
gerardi, Andropogon, 84-85, 353, 536
germander, 83, 212
getulus, Lampropeltis, 52, 75
ghost shiner, 366, 423
giant ragweed, 82, 93, 99, 155, 159,

166, 220, 241, 244, 296
gilvus,

Vireo, 272

Vireo gilvus, 493
girardi, Notropis, 368, 385-86. 394-95
giraudi, Dendrocopus scalaris, 449, 472
gizzard shad, 363, 415
glabra, Rhus, 84, 146, 536
glass hzard, 70, 321, slender, 78
glass snake, slender, 104
Glaucidium

cahfornicum, 466

gnoma, 466
gnatcatcher, 321

black-tailed. 489

blue-grav, 81, 263, 448-49, 451,
488-89
gnoma, Glaucidium gnoma, 466
golden

redhorse, 364, 379

shiner, 372
golden-checked warbler, 496
golden-crowned kinglet, 77, 221, 245,

250, 264-65, 489
golden-fronted woodpecker, 448-49,

470-71
goldenrod, 82, 84, 295
goldeye, 415
goldfinch, 319-20

American, 505

eastern, 81, 299

lesser, 505
goldfish, 419
goose,

blue, 77, 173

Canada, 77, 172

snow. 77, 173

white-fronted, 77, 173

Index to Volume 11

683

gooseberry, 135, 209, 277

goosefoot, 82

gopher,

pocket, 88, 123
plains pocket, 138
Gopherus, 652, 656

agassizi, 557, 563, 573, 600, 611,

627, 630, 642-43
agassizii, 338, 340-42
berlandieri, 338-41, 628
flavomarginatus, 337
polyphemus, 338, 340-42
goshawk, 77, 177, 317, 449, 456
gracile, Etheostoma, 393
gracilis,

Melospiza lincolnii, 513
Peromyscus maniculatus, 145
Procambaris, 621
Tantilla, 124
grackle,

boat-tailed, 499
bronzed, 77, 286
grama, side-oats, 141, 295, 536
gramineus,

Pooecetes, 304
Potamogeton, 353
grammacus, Chondestes, 304
grape, 214, 257, 619, fox, 270
grass,

awnless brome, 83, 190, 306
bluestem, 281

brome, 49, 123, 128, 138-39, 141,
143, 168, 224, 278, 281, 287,
296, 304
Indian, 85, 141, 295, 536
Johnson, 353
muhly, 155, 161
poverty, 143
slough, 353, 536
switch, 85, 141, 295
three-awn, 84, 106, 140, 304
grasshopper, 31, 103, 214, 581-82, 601,
617-18, 659, 661
differential, 622
sparrow, 77, 303, 508
gray,

flycatcher, 475
fox, 77, 164
hawk, 458

squirrel, 69, 79, 136, 321-22
treefrog, 78, 93
warbler, black-tluoated, 495
gray-backed thrush, 77
gray-breasted martin, 448, 479
gray-cheeked thrush, 259
great

blue heron, 76-77, 170, 453
crested flycatcher, 473
horned owl, 466
Great Plains

narrow-moutlied frog, 79

Great Plains — Concluded
narrow-moutlied toad, 95
skink, 67, 78, 110, 187, 321
greater yellowlegs, 462
grebe, 317
eared, 453
pied-billed, 77, 170
green

algae, 353
heron, 77, 171, 454
kingfisher, 448, 450, 469
sunfish, 77, 381, 390, 392, 429
warbler, black -throated, 77, 275,
496
green-tailed towhee, 506
green-winged teal, 77, 175-76, 455
grey rat snake, 52
grisea, Amphispiza bilineata, 449, 509-

10
griseus,

Empidonax, 475
Vireo, 268
groove-billed ani, 448, 465
grosbeak,

black-headed, 502
blue, 77, 294, 321, 451, 503
rose-breasted, 77, 293
ground,
beetle, 31
dove, 464
skink, 78, 109
ground squirrel, Frankhn's, 77, 136
growth

of adult ornate box turtle, 578

of epidermal laminae of ornate box

turtle, 568
of juveniles of ornate box turtle,

575
of ornate box turtle, 565
of six-lined racerunner, 39
grunniens, Aplodinotus, 383, 385-86,

388-89, 430, 423-33
Grus canadensis, 192, 461
gryUus, Acris, 91
guadalupensis, Najas, 84, 360
Guiraca

caerulea, 294, 503
eurhyncha, 451, 503
interfusa, 503
gularis, Cnemidophorus sacki, 18
gull,

Franklin, 77, 196
Franklin's, 317
herring, 462
ring-billed, 463
gulosus, Chaenobryttus, 384
guttata,

Elaphe, 52
Hylocichla, 259, 487
guttatus, Campylorhynchus brunnei-

capillum, 449, 485
Gymnocladus dioica, 284

684

University of Kansas Publs., Mus. Nat. Hist.

habitat

and limiting factors of ornate box

turtle, 539
of ornate box turtle in Kansas, 542
of six-lined racerunner, 18
hachisukai, Chloroceryle americana,

469
hackberr>-, 83, 268, 353, 536
hairy woodpecker, 80, 214-15, 218,

319, 322, 448, 450, 471
llaldea valeriae, 115
halepense. Sorghum, 360
Haliacetus leucocephalus, 187
haliaetus, Pandion, 458
hammondi, Scaphiopus, 617
hammondii, Empidonax, 475
Hammond's flycatcher, 475
harlequin quail, 461
Harris' hawk, 448, 458
Harris sparrow, 82, 291, 307, 313-14
harrisi, Parabuteo unicinctus, 458
Hartlaub's warbler, 449, 494
harvest mouse, 52, 160, 206, 321
plains, 76, 79, 85, 140
western, 79, 140-41, 154
hastata. Verbena, 536
haw, 113, red, 82, 271
hawk, 322

broad-winged, 53, 77, 184-86, 457

Cooper, 77, 178. 191

Cooper's, 53, 319, 457

ferruginous, 458

gray, 458

Harris', 448, 458

marsh, 53, 77, 188. 317, 458, 648

pigeon, 459

red-shouldered, 77, 183

red-tailed, 77, 159, 180, 184, 187,

319-20, 457, 648
rough-legged, 77, 187
sharp-shinned, 77, 178, 456
sparrow, 53, 77, 85, 189, 459
Swainson's, 457
zone-tailed, 457
hawthome, 506
Hedymeles papago, 502
Helianthus annuus, 82
Helisoma. 621

hellmayri, Empidonax difficilis, 476
hepatic tanager, 501
hepatica, Piranga flava, 501
hermit

thrush. 77, 259, 487
warbler, 496
herodias, Ardea, 170, 453
heron, 317

black-crowned night, 77, 172, 454
great blue, 76, 77, 170, 453
green, 77. 171, 454
yellow-crowned night, 454
herring gull, 462
Heterodon platyrhinos, 52

heterophvllum, MyTiophvllum, 353,

360-61. 366, 370
hibernation of ornate box turtle, 611
hickory, 109, 123, 125, 137, 209, 218,
235, 293-94, 353
shagbark, 204, 248, 271
highfin carpsuckcr, 363, 417
Hiodon alosoides, 383, 415, 432-33
Hirundo

erythrogaster, 478
rustica, 235
hispid cotton rat, 80, 149, 154
hispidulus, Sitonia, 31
hispidus, Sigmodon, 149
hoactli, Nycticorax nycticorax, 454
hog-nosed snake, 52
Holbrookia maculata, 52
Home ranges, territories, and seasonal
movements of vertebrates of the
Natural History Reservation, 65
honey locust, 85, 93, 113, 147, 161,
198, 201, 203, 206-07, 214-15, 220,
239, 256, 260. 263, 270-71, 278,
289-90, 294
hooded

oriole, 499

yellowthroat, 449, 496
hoopesi, Stumella magna, 498
homed

lark, 80, 85, 232, 321-22, 477
owl, 77, 174, 193. 201, 240, 319,
466
horridus, Crotalus. 52, 128
horsefieldi, Testudo. 600
horseweed, 82
house

cat, 317
finch, 504

mouse, 80, 160, 321
sparrow, 317, 497
wren, 77, 251, 483
howelH, Chordeiles minor, 468
hudsonius, Zapus, 161
humifusa, Opuntia, 536
humilis, Lepomis. 369, 381, 385-86,

388-91, 395-97, 429, 431-33
hummingbird,

black-chinned, 469
blue-throated, 469
broad-tailed, 469
buff-bellied. 469
Lucifer, 468
Rivoh's. 469

ruby-throated, 77, 212, 469
liutton's vireo, 448-49, 451, 492
llybognathus

nuchalis, 423-26, 432-33
placita, 386, 395, 423-26
Ihbopsis

aestivalis, 385-86. 419, 432-33
amblops, 393
biguttata, 393

Index to Volume 11

685

Hybopsis — Concluded

hyostomus, 419

storeriana, 419, 432-33

tetranemus, 365, 394-95, 419

x-punctata, 393
hyemalis, Junco, 305
Hyla versicolor, 93
Hylocichla, 452

auduboni, 487-88

fuscescens, 260

guttata, 259, 487

minima, 259

mustelina, 258

ustulata, 259
Hymenoptera, 621
hyostomus, Hybopsis, 419
Hypentelium nigricans, 393
hyperborea, Chen, 173
Hypsiglena, 3

Hypsirhynchus, 8 . ■, . ac-t

hypugaea, Speotyto cuniculana, 467

icastus, Dendrocopos villosus, 450, 471

^''mS: 87, 377, 381, 385-86 388-
92, 394-97, 414, 427, 431-33,
438
nataUs. 377, 385-86, 388-89, 391,

393, 427, 432-33
punctatus, 377, 385-86, 388-89,
427. 431-33, 438
Icteria

auricollis, 496
virens, 278, 496
Icteridae, 453
Icterus

bulockii, 499
cucuUatus, 499
galbula, 285
parisonmi, 499
spurius, 284, 498
wagleri, 499

^^*bubalus, 363-64, 374, 385-86, 388,

416,432-33 ^ ^,„

cyprineUa, 364, 385-86, 388, 416,

432 33
niger, 363, 385-86, 388, 416, 432-

33

veUfer, 417
ihaca, Passerella, 315
immeroratus, Empidonax ditticilis, 4/b
Inca dove, 464
inca, Scardafella, 464
incanescens, Centurus aurifrons, 470-

71
Indian grass, 85, 141, 295, 536
indigo bunting, 81, 85, 294, 321, 503
inermis, Bromus, 83
injuries of ornate box turtle, 638
inomatus, Cnemidophorus, 43

insculpta, Clemmys, 653
interfusa, Guiraca caerulea, 503
intermedius,

Dendrocopos villosus, 448, 472

Meleagris gallopavo, 461
ioensis, Pyrus, 82
Ipomoea, 8

Iridoprocne bicolor, 478
ironweed, 83

Ps'altriparus melanotis, 451, 482
Psaltriparus minimus, 482
ivy, poison, 214, 268, 274

jack rabbit, 162
jamaicensis, Buteo, 180, 648
Japanese chess, 159, 536
iaponicus,

Bromus, 159, 535-36

Trionyx, 552

'""^blue, 70, 81, 235, 318-19
Mexican, 479
scrub, 449, 479
Steller's, 449, 479
Joe-Pye weed, 82
johnny darter, 430
Johnson grass, 353
Juglans nigra, 535

jumping mouse, 80, 321, meadow, Ibl
junco, 82, 245, 291, 307, 313, 322
Mexican, 510
slate-colored, 305
Junco

hyemalis, 305
paUiatus, 510-11
juniper, 540

Juniperus virgianus, 249
Jussiaea diffusa, 353, 361
Justicia americana, 353, 360-61

kansae, Fundulus, 376, 385-86, 394-

95, 428, 432-33
Kentucky

coffee-tree, 284

warbler, 68, 81, 276, 320
kiUdeer, 77, 194, 317, 461
killifish, 376

plains, 428
king snake, 52, 75, prairie, 51, 77, 123
kingbird, 85

Cassin's, 473

eastern, 77, 222
kingfisher, _ ^ ^^

Belted, 77, 212, 317, 469

green, 448, 450, 469

loWen-crowned, 77, 221, 245, 250,
264-65, 489

ruby-crowned, 77, 265, 489
Kinixys, 588 „^^ ^^„ ^-

klauberi, Terrapene, 532, 650, 652-53
Kuhnia eupatoroides, 82

686

Unix-ersity of Kansas Publs., Mus. Nat. Hist.

Labidesthes sicculus, 378, 385-86,

388-91, 393, 396-97
Lactuca scariola, 82
ladder-backed woodpecker, 448-50,

472
ladybird beetle, 31
lagopus, Buteo, 187
Lampomis clemenciae, 469
Lampropeltis

calligaster, 51, 123

doliata, 124

getulus, 52, 75

rhombomaculata, 124
Lampyridae, 621
land

terrapin, 534

tortoise, 534
Lanius

excubitorides, 490-91

gambeU, 491

ludovicianus, 266

mexicanus, 491

migrans, 490
largemouth bass, 380, 429, 437
lark

bunting, 507

horned, 80, 85, 232, 321-22, 477

sparrow, 77, 85, 304, 508
Larrea tridentata, 462
Larus

delawarensis, 463

pipixcan, 196

smithsonianus, 462
Lasiurus borealis, 134
laterale, Lygosoma, 109
latifolia,

Sagittaria, 84, 353, 361

Typha, 158, 353
latifrontalis, Geophis, 332
latrans, Canis, 162, 646
least

flycatcher, 77, 230, 475

sandpiper, 462
Leconte sparrow, 77, 304
Leersia oryzoides, 141
Legler, John M.,

Natural history of the ornate box
turtle, Terrapene omata ornata
Agassiz, 529

A new snake of the genus Geophis
from Chihuahua, Mexico, 329

A new tortoise, genus Gopherus,
from north-central Mexico, 337
Leimadophis, 8

lentiginosus, Botaurus, 172, 454
leopard frog, 78, 97, 104, 171, 318,

537
lepida,

Phainopepla nitens, 490

Tachycineta thalassina, 451
Lepidoptcra, 621-22

Lepisosteus

osseus, 385-86, 388-89, 395, 414-
15, 432-33

oxyurus, 362, 415

platostomus, 363, 393, 415, 432-33

productus, 383
Lepomis

breviceps, 382, 393

cyanellus, 87, 381, 385-86, 388-92,
394-97, 421-22, 429, 431-33

humilis. 369, 381, 385-86, 388-92,
395-97, 429, 431-33

macrochirus, 382, 385-86, 388-89,
414, 429, 432-33

megalotis, 385-86, 388-91, 396

microlophus, 384
Leptodeira, 3

discolor, 3, 4
Leptotila angelica, 464
lespedeza, 83-84, 106, 140, 143, 191,

304
Lespedeza striata, 83
lesser

nighthawk. 468

scaup, 77, 176, 455

yellow-legs, 77, 195, 462
lettuce,

prickly, 159

wild, 82
leucocephalus, Haliaeetus, 187
leucopareia, Branta canadensis, 454
leucophrys,

Zonotrichia, 313

Zonotrichia leucophrys, 512
leucopterus, Mimus polyglottos, 485
leucopus, Peromyscus, 146
limicola, Rallus, 193
Lincoln sparrow, 77, 315
lincolnii,

Melospiza, 315

Melospiza lincolnii, 512
Lincoln's sparrow, 512
linden, 85
lineatus, Buteo, 183
lineolatum, Etheostoma flabellare, 393
lipovskyana, Trombicula, 644
Lisscmvs, 588, punctata, 552
little

bluestem, 85, 141, 295

short-tailed shrew, 79, 132
livia, Columba, 75
lizard,

collared, 25, 67, 51, 76, 78, 102,
187, 318, 321, 601, 616-17

earless, 52

glass, 70, 78, 321

scaly, 52

slender glass, 78
lloydi, Psaltriparus melanotis, 451, 482
locust, 139, 199, 218, 296, 301

honey, 85, 93, 113, 147, 161, 198,
201. 203, 206-07, 214-15. 220,

Index to Volume 11

687

locust honey — Concluded

239, 2^6, 260, 263, 270-71, 278,

289-90, 294
Locustidae, 621

bgg"erhead sh'rike, 77, 85, 266, 490-91
logperch, 378
long-billed
curlew, 462
marsh wren, 485
thrasher, 448, 486
longear sunfish, 382
long-eared owl, 77, 206
longevity of the ornate box turtle, 5»o
longicauda, Bartramia, 195
longinsulae, Terrapene, 534, 656
longnose gar, 362, 415
long-tailed

chat, 77, 278, 321
weasel, 77, 166
loon, 317

lotor, Procyon, 52, 164, 646
Louisiana water-thrush, 77, 276
lucifer, Calothorax, 468
Lucifer hummingbird, 468
lucius, Essox, 437
ludoviciana,

Artemesia, 536
Piranga, 501
ludovicianus,
Lanius, 266
Pheucticus, 293
Thrvothorus, 252
lutea, Nelumbo, 353, 360-61, 366
luteola, Terrapene ornata, 372, 449,
533^34, 540-43, 597, 617-18, 629-
30, 654-55, 657
lutescens, Setaria, 536
lutrensis, Notropis, 369-71, 385-86,
388-92, 395-97, 414, 421, 431-33,

438
Lygosoma laterals, 109

Madura pomifera, 536 oc c«

macrochirus, Lepomis, 382, 38o-8o,

388-89, 414, 429, 432-33
macrolepidotum, Moxostoma, 418
macrolopha, Cyanocitta stelleri, 479
macromystax, Antrostomus vociferus,

467
macropterus, Spinus pinus, 505
macroura, Zenaidura, 196
macularia, Actitis, 195, 462
maculata, Holbrookia, 52
maculatus, Pheucticus melanocephalus,

503
madrone, 469
madtom,

catfish, 395

freckled, 383
Magicicada septendecim, 618
magister, Sceloporus, 22

magna Sturnella, 281

Malaclemys terrapm, 552, 5bi-b^

mallard, 77, 174, 454

maniculatus, Peromyscus, 143

maple, 409, 465, 469

Mareca americana, 455

marginella, Zenaidura macroura, 4bo5

marilandica, Quercus, 353

marine turtle, 552

Marmota monax, 136

"'"hawk, 53, 77, 188, 317, 458, 648
wren: long-billed, 485; short-billed,
77, 255
marsupialis, Didelphis, 129, 648

grav-breasted, 448, 479

purple, 77, 235, 479
massasauga, 51
Masticophis flagellum, 51
mayensis. Bubo virginianus, 46b

mayfly, 31 . ,^„

mccallii, CHus asio, 4bb
meadow jumping mouse, 161
meadowlark, 85

eastern, 81, 281, 498

western, 77, 282, 498
mealworm, 619
meamsi,

Cyrtonyx montezumae, 4bi

Zenaida asiatica, 463
medius, Vireo bellii, 448-49, 492
Megaceryle alcyon, 212, 469
megalotis,

Lepomis, 385-86, 388-91, 396

Reithrodontomys, 52, 141
megapotamus, Agelaius phoeniceus,

498
Melanei-pes,

erythrocephalus, 217
formicivorus, 471
melanocephalus, Pheucticus melano-
cephalus, 502
melanocorys, Calamospiza, 507
melanoleucus,
Pituophis, 52

Totanus, 462 , ^r.^ m

Melanoplus, 31, difiFerentialis, 620-21
melanops, Minytrema, ob5, vibb-ao,

388, 390-91, 394, 396
melanotos,
Eroha, 462
Sitta pygmaea, 482
melanura, PoUoptila melanura, 4«y
melas,

Ameiurus, 377, 427
Ictalurus, 87, 377, 381, 385-86,
388-92, 394-97, 414, 427, 431-
33, 438
Meleagris intermedins, 461
melodia, Melospiza, 316
melon, coyote, 617

688

Unr'ersity of Kansas Publs., Mus. Nat. Hist.

Melospiza

allicola, 512

crico-pta. 513

georgiana, 316

gracilis, 513

lincolnii, 315, 512

melodia, 316
memorabilis, Dendroica auduboni, 495
Mephitis mephitis, 52, 167, 646
mephitis. Mephitis, 52, 167, 646
mesquite, 485, 495. 502, 540
Metcalf, Artie L., Fishes of Chautau-
qua, Cowley and Elk Counties,

Kansas, 347
Mexican

chickadee, 449, 481

jay, 479

iunco, 510
mexicana,

Sialia mexicana, 488

Sitta carolinensis, 451, 482

Terrapene, 532, 654

Vermivora superciliosa, 494
mexicanus,

Cassidix mexicanus, 500

Falco, 189, 458

Lanius ludovicianus, 491

Myriarchus mexicanus, 474

Pyrocephalus rubinus, 448-49, 477
Micrathene whitneyi, 467
microlophus, Lepomis, 384
Micropterus,

punctulatus, 380, 385-86, 388-89,
393, 396

salmoides. 380, 385-86, 388, 391,
429, 432-33
Microtus, 30

ochrogaster, 30, 70, 152

pinetorum, 70, 155

scalapsoides, 158
micrus, Vireo griseus, 491
migrans, Lanius ludovicianus, 490
migratorius, Turdus, 257
miharius, Sistrurus, 52
milk snake, 77, 124
milkweed, 83

milo, 85, 232, 307, 313, 535
mimic sliiner, 372
Mimus

leucopterus, 485

polyglottus, 255
Minckley, W. L., Fishes of the Big

Blue River Basin, Kansas, 403
minima,

Hylocichla, 259

Petrochelidon pyrrhonota, 478
minimus, Empidonax, 230, 475
mink, 317
minnow. 395

black-fin, 420

blackhead, 426

bluntnose, 373, 426

minnow, — Concluded

fathead. 77, 86, 374, 426

mountain, 373

parrot. 374

plains suckcmiouth, 420

suckermouth, 372
minor,

Chordeiles, 210

Philohela, 194
minutilla, Erolia, 462
Minytrema melanops, 365, 385-86,

388. 390-91, 394. 396
mirabilis,

Phenacobius, 372, 385, 420, 431-32

Pimephales, 386, 388
missuriensis, Notropis deliciosus, 368,

395, 423, 431
mite, chigger, 643-44, 662
miurus, Noturus, 393
Mniotilta varia, 272, 493
mockingbird, 77, 255, 485
mole, 79, 156, eastern, 133
mollis, Crataegus. 82
Molothrus

artemisae, 500

ater, 287, 500

obscurus, 500
molt, six-lined racerunner, 46
monax, Marmota, 136
monsoni, Cassidix mexicanus, 500
montana,

Certhia famiharis, 483

Eupoda, 462
montanensis, Trombicula, 644
montanus,

Oreoscoptes, 487

Piplio maculatus, 506

Reithrodontomys, 140
montezumae, Cyrtonyx montezumae,

461
morcomi, Dendroica petechia, 495
Morus rubra, 536, 619
mosquito, 31
mosquitofish, 376
motacilla, Seiurus, 276
mountain

bluebird, 488

minnow, 373

plover, 462
mourning dove, 80, 196, 231, 319-20,

463
mouse,

deer, 79, 143, 321

harvest, 52, 76, 79, 85, 140-41, 154,
160, 206, 321

house, 80, 160, 321

jumping, 80, 321

meadow jumping, 161

oldfield, 29

plains harvest, 76, 79, 85, 140

prairie deer, 145

western harvest, 79, 140-41, 154

Index to Volume 11

689

mouse, — Concluded

white-footed, 80, 143, 147, 160,
207-08, 321

woodland deer, 145
movements

of ornate box turtle, 626

of six-lined racerunner, 26
Moxostoma

aureolum, 418, 432-33

carinatum, 364, 393

duquesnei, 418

duquesnii, 393

erythrurum, 364, 385-86, 388, 391,
393, 396, 418

macrolepidotum, 418

pisolabrum, 364, 393
Mucorales, 640
mud-dauber, 250
mugwort, 536

Muhlenbergia Schreberi, 535
muhly grass, 155, 161
mulberry, 601-02, 618-19, 661, red,

536
mullein, 536

multistriatus, Scolytus, 221
multivirgatus, Eumeces, 52
murinus, Thryomanes bewickii, 484
Muscivora forficata, 222, 473
Mus musculus, 160
musculus, Mus, 160
musk tiu-tle, 548
muskrat, 77, 158, 317
Mustela frenata, 166
mustelina, Hylocichla, 258
mutica, Amyda, 519, 523-24
muticus,

Amyda, 519

Trionyx, 519-20, 522-25
Myadestes, 452, townsendi, 488
Myiarchus

boreus, 473

cinerascens, 474

cooperi, 474

crinitus, 223, 473

mexicanus, 474
Myiochanes richardsonii, 476
Myriophyllum heterophyllum, 353,

360-61, 366, 370
myrtle warbler, 77, 245, 274

naiad, 84

Najas guadalupensis, 84, 360

nanus, Colaptes cafer, 470

narrow-mouthed toad, 67, 93, 318,
Great Plains, 78, 95

Nashville warbler, 77, 274

natalis, Ictalurus, 377, 385-86, 388-89,
391, 393, 427, 432-33

Natrix sipedon, 113

Natm^al history of the ornate box tur-
tle, Terrapene omata omata Agas-
siz, 529

Natural history of the six-lined race-
runner ( Cnemidophorus sexlinea-

tus), 13
neglecta,

Stumella, 282

Stumella neglecta, 498
Negumbo, Acer, 536
negundo, Acer, 84
nelsoni,

Geothlypis nelsoni, 496

Sitta carolinensis, 451

Terrapene, 532, 650
Nelumbo lutea. 353, 360-61, 366
Neotoma floridana, 151
nesting of the ornate box turtle, 554
nevadensis, Passerculus sandwichensis,

508
New snake of the genus Geophis from

Chihuahua, Mexico, 329
New softshell turtle from the south-
eastern United States, 519
New tortoise. Genus Gopherus, from

north-central Mexico, 337
niger,

Ictiobus, 363, 385-86, 388, 416,
432-33

Sciurus, 137
night heron,

black-crowned, 77, 172, 454

yellow-crowned, 454
nighthawk, 77, 179, 210, 468

common, 468

lesser, 468
nightshade, 82-83
nigra,

Juglans, 535

Salix, 536
nigrescens,

Carpodacus mexicanus, 505

Dendroica, 495
nigricans.

Hypentelium, 393

Sayomis nigricans, 451, 474
nigriflora, Parula pitiayumi, 494
nigrifrons, Dendroica auduboni, 495
nigrita, Pseudacris, 94
nigromaculatus, Pomaxis, 381, 385-86,

429, 432-33
nigrum,

Boleosoma nigrum, 430

Etheostoma, 432-33

Etheostoma nigrum, 393, 430
nitens, Phainopepla nitens, 490
nitidus, Buteo, 458
noctuid caterpillar, 620, 622
Noctuidae, 621
noctumus, Noturus, 383, 393
nodosus, Potamogeton, 353, 370
Nomotettix, 31
northern

pike, 437, 438

prickley ash, 535

690

University of Kansas Publs., Mus. Nat. Hist.

northern — Concluded

redhorse, 364. 418

water-thrush, 77, 276
norvegicus, Rattus, 159
Norway rat, 80, 159
notatus,

Fundulus, 376, 385-86, 388-92,
396-97

Pimephales, 372-74, 379, 385-86,
388-89, 393, 396, 426, 432-33

Zygonectes, 376
Notemigonus crysoleucas, 372, 385
Notochelys, 532
Notropis, 390, 395-96

blennius, 365, 385-86, 394-95, 423

boops, 365, 368, 370, 372, 385-86,
388-89, 393-94, 396-97

buchanani, 366, 385-86, 388-89,
397, 423, 432-33

camerus, 367, 370, 373, 379, 385-
86, 388-89, 393, 396-97

comutus. 432-33

crysoleucas, 386

dehciosus, 376, 386, 395, 404, 423,
431-32, 438

frontalis, 421, 431

girardi, 368, 385-86, 394-95

lutrensis, 369, 370-71, 385-86, 388-
92, 395-97, 414, 421, 431-33,
438

missuriensis, 368, 395, 423, 431

percobromus, 369-70, 385-86, 388-
90, 395, 397, 420, 432-33

pilsbryi, 393

rubellus, 366, 368, 370, 372, 385-
86, 388-89, 396-97, 420, 432-33

rubrifrons, 420

spilopterus, 393

topeka, 371, 410, 423, 432-33

umbratilis, 371, 379, 382, 385-86,
388-92, 396-97. 420, 432-433

volucellus, 367, 372, 385-86, 388,
393-94, 396-97
Noturus

exilis, 393

flavus, 383, 428, 432-33

miurus, 393

nocturnus. 383, 393
noveboracensis,

Seiurus, 276

Vireo griscus, 491
nubila, Dionda, 393
nuchalis,

Hybognathus, 423, 432-33

Hybognathus nuchalis, 424-26

Sphyrapicus varius, 471
Numenius parvus, 462
nutallii, Phalaenoptilus nutaUii, 467
nutans, Sorghastrum, 85, 353, 536
nuthatch,

pigmy, 482

nuthatch, — Concluded

red-breasted, 77, 249

white-breasted, 77, 248, 451, 482
Nuttallomis boreahs, 232, 476
Nyctanassa violacea, 454
Nyclicorax

hoactli, 454

nycticorax, 172
nycticorax, Nycticorax, 172

oak, 109, 123, 125, 137, 201, 209, 214,
218, 235-36, 271, 294, 353-54,
409. 469, 489, 492, 498

black. 82, 204, 271, 293

blackjack, 19

chestnut, 203, 248, 293

chinquapin, 206-07, 263

dwarf, 268, 271

post, 19, 85

red, 85, 203, 217, 271
oak-liickory, 136, 178, 196, 203, 210,
289

woodland, 85, 263

woods, 167
oak-hickory-elm, 245
oats, 85, 295
oberholseri, Toxostoma cvurvirostre,

448, 486
obliqulus, Agonoderis, 31
obhtus, Passerculus sandwichensis,

508
obscurus, Molothrus ater, 500
obsoleta, Elaphe, 52, 54, 121
obsoletus,

Eumeces, 53, 110, 113, 623

Salpinctes obsoletus, 485
occidentalis,

Celtis, 536

Coccyzus americanus, 464

Dondroica, 496

Empidonax difRcilis, 476

Geothlypis trichas, 496

Platanus, 409
ochrogaster, Microtus, 30, 70, 152
octohneatus, Cnemidophorus sacki, 17
odoratus, Sternotherus, 548, 559, 561,

624, 649
Oepidopinae, 621
oldfield mouse, 29
oligantha, Aristida, 84
olivacea,

Gastrophrvne, 95

Piranga, 288
olivaceus, Viroo, 271, 493
olivaris, Pylodictis, 377, 385-86, 388-

89, 428, 432-33, 438
olive

sparrow, 448, 506

warbler, 494
olive-backed

thrush, 77, 259

warbler, 494

Index to Volume 11

691

olive-sided flycatcher, 77, 232, 476
Ondatra zibethicus, 158
Ophisaurus attenuatus, 53, 104
Oporomis formosus, 276
opossum, 67, 71, 79, 129, 167, 169,

243, 321-22, 538, 648, 663
Opuntia humifusa, 536
opuntia, Amphispiza bilineata, 451,

509-10
orange, osage, 93, 113, 139, 142, 151-

52, 178, 198-99, 206, 208, 239-41,

246, 256, 263, 271, 278, 280, 290,

294, 301, 313-14, 536
orange-crowned warbler, 77, 273, 493
orangespotted sunfish, 381, 429
orangethroat darter, 379, 430
orbicularis, Emys, 642-43
orbiculatus, Symphoricarpos, 535
orchard oriole, 77, 284, 498
Oreoscoptes montanus, 487
orestera, Vermivora celata, 494
oriantha, Zonotrichia leucophrys, 512
or ien talis, Piplio erythrophthalmus,

506

origin of breeding birds of Coahuila,

452
oriole,

Baltimore, 77, 285

Bullock's, 499

hooded, 499

orchard, 77, 284, 498

Scott's, 499

Wagler's, 499
ornata,

Terrapene, 51, 54, 99, 527-669

Terrapene ornata, 527-669
ornate box turtle, 322, 527-669

adaptations, 650

bony shell, 586

color and markings, 593

defense, 648

diet, 617

economic importance, 534

ectoparasites, 643

embryonic development, 560

fertility and prenatal mortality, 564

growth and development, 565

growth in adult, 578

growth of epidermal laminae, 568

growth of iuveniles, 575

habitat, 542

habitat and limiting factors, 539

hibernation, 611

injuries, 638

longevity, 585

movements, 626

nesting, 554

populations, 623

predators, 646

reproduction, 543

sexual dimorphism, 596

ornate box turtle — Concluded

temperature relationships, 598

wear on shell, 595

weight, 586

years unfavorable for growth, 582
omatus, Auriparus, flaviceps, 481
Orthoptera, 621-22
oryzoides. Leersia, 141
osage orange, 93, 113. 139, 142, 151-

52, 178, 198-99, 206-08, 239-41,

246, 256, 263, 271, 278, 280, 290,

294, 301, 313-14, 536
osprey, 458
osseus, Lepisosteus, 385-86, 388-89,

395, 414-15, 432-33
Otocoris aphrasta, 477
otter, 317
Otus

asio, 200

cineraceus, 465

flammeolus, 466

mccallii, 466

semplei, 466

suttoni, 465-66
otus, Asio, 206
oven-bird, 77, 275
Ovis arias, 75
owl, 322

bam, 77, 200, 317, 465

barred, 77, 203, 319

burrowing, 467

elf, 450, 467

flammulated, 466

great horned, 466

horned, 77, 174, 193, 201, 240,
319, 466

long-eared, 77, 206

pygmy, 466

saw-whet, 77, 207

screech, 77, 200, 322, 465-66
oxyurus, Lepisosteus osseus, 362, 415

Pachyrhamphus aglaiae, 473
pacificus, Anthus spinoletta, 490
paddlefish, 383
painted

bunting, 77, 295, 504

redstart, 497

turtle, 77, 101, 317, 557, 585, 643
pallescens. Bubo virginianus, 466
palliatus, Junco phaenotus, 510-11
pallida,

Callipepla squamata, 450, 459-60

Petrochelidon fulva, 478

Spizella, 308, 511
pallidus, Spinus tristis, 505
palm, 492

Pandion hahaetus, 458
Panicum virgatum, 85, 353, 536
papago, Hedymeles melanocephalus,

502
Parabuteo harrisi, 458

692

University of Kansas Publs.. Mus. Nat. Hist.

parisorum, Icterus, 499
parkmanii. Troglodytes aedon, 483
parrot,

minnow, 374

thick-billed, 449, 464
Parula

americana, 494

nigrilora, 494
parula warbler, 494
Parus

atricapillus, 241

atricristatus, 448-49, 481

bicolor, 245

dvsleptus, 451, 481

eidos, 480

sclateri, 481
parva, Cryptotis, 132
parvulus, Calendra, 621
parvus, Numenius americanus, 462
Passer domesticus, 280, 497
Passerculus

anthinus, 508

brooksi, 508

brunnescens, 508

nevadensis, 508

oblitus. 508
Passerella iliaca, 315
Passerherbulus caudacutus, 304
Passerina

ciris, 295

cyanea. 503

pallidior, 504

versicolor, 503
passerina,

Columbigallina, 464

Spizella, 308
pecan, 498

pectinata, Spartina, 353, 536
pectoral sandpiper, 462
pelagica, Chaetura, 211
Pelecanus erytlirorhynchus, 170, 453
pelican, white, 77, 170, 453
Pelusios, 588
Perca flavescens, 430
perch,

ring, 430

yellow, 430
Percina

caprodes, 385-86, 388, 391

carbonaria, 378

copelandi, 379, 385-86, 388-89,
393-94

phoxocephala, 378-79, 385-86, 388-
89, 393
percobromus, Notropis, 369-70, 385-

86, 388-89, 395. 397, 420. 432-33
peregrina, Vermivora, 273
Pcromyscus, 30

i gracilis, 145
eucopus, 146
maniculatus, 143
polionotus, 29

perpallidus, Amniodramus savanna-
rum, 508
perplexus, Cnemidophorus, 20, 38, 43
perspicuus, Pimephales vigilax, 374
pertinax, Contopus pertinax, 476
petechia, Dendroica, 274
Petrochelidon

minima, 478

pallida, 478

pyrrhonota, 235
Peucedramus arizonae, 494
pewee, 321

wood, 72

wood, eastern, 80, 230, 476

wood, western, 476
Phainopepla

lepida, 490

nitens, 490

nutallii, 467
phainopepla, 490
Phalangida, 621
phalarope, Wilson's, 462
Pharyngodon wameri, 53, 58
Phasianidae, 452
Phenacobius mirabilis, 372, 385, 420,

431-32
Phengodidae, 621
Pheucticus

ludovicianus, 293

maculatiis, 503

melanocephalus, 502
Philohela minor, 194
phoebe, Savornis, 224, 474
phoebe, 80,' 189, 223-24, 231, 255, 320

black, 451, 474

eastern, 474

Say's, 474
phoeniceus, Agelaius, 283
Photinus p\Talis, 621
Photuris, 621
phoxocephala, Percina, 378-79, 385-

86, 388-89, 393
Picidae, 452
picta,

Chrysemvs, 101, 552, 554, 559, 585,
611, 627,643

Setophaga picta, 497
pied-billed grebe, 77, 170
pig, 162
pigeon,

band-tailed, 449, 451, 463

domestic, 75

hawk, 459
pigmy

nuthatch, 482

rattlesnake, 52
pike,

northern, 437-38

yellow, 430
pileolata, Wilsonia pusilla, 497
pilsbryi, Notropis zonatus, 393

Index to Volume 11

693

Pimephales

mirabilis, 386, 388

notatus, 372-74, 379, 385-86, 388-
89, 391, 393, 396, 426, 432-33

perspicuus, 374

promelas, 86, 372, 374, 385-86,
388, 414, 426, 432-33

tenellus, 372-74, 385-86, 388-89,
393 397

vigilax,' 372-73, 385-86, 388-89
pine, 449, 494

flycatcher, 449, 475

siskin, 77, 299, 449, 505

snake, 52

vole, 80, 131, 154-55

white, 469, 496
pinetorum, Microtus, 70, 155
pinicola, Ridgwayia, 487
pifion, 484
pintail, 77, 174, 455
pinus,

Spinus, 299

Spiniis pinus, 505
pipiens, Rana, 97, 537, 618
Pipilo

erythrophthalmus, 300

gaigei, 507

potosinus, 507

texanus, 507
pipit, water, 490
pipixcan, Larus, 196
Piplio

gaigei, 506

montanus, 506

orientalis, 506
Piranga

cooperi, 449, 451, 501

dextra, 501

hepatica, 501

ludoviciana, 501

olivacea, 288

rubra, 289, 448, 501
Pituophis

catenifer, 122. 640

melanoleucus, 52
pisolabrum, Moxostoma aureolum,

364. 393
placita,

Hybognathus, 386, 395, 423

Hybognathus nuchalis, 424-26
plains

carpsucker, 416

gopher, 77, 138

harvest mouse, 76, 79, 85, 140

killifish, 428

minnow, 420

pocket gopher, 77, 138

shiner, 369, 420

spadefoot toad, 77, 88

suckermouth minnow, 420

toad, 77, 88
Platanus occidentalis, 409

platensis, Cistophorus, 255
platorynchus, Scaphirrhynchus, 414,

432-33
platostomus, Lepisosteus, 363, 393,

415, 432-33
platycercus, Selasphorus platycercus,

469
platypterus,

Buteo. 58, 186

Buteo platypterus, 457
Platypsaris albiventris, 473
platyrhinos, Heterodon, 52
platyrhnchos. Anas, 174, 454
plesius, Telmatodytes palustris, 485
plover,

mountain, 462

upland, 77, 185
plum, 268, wild, 82, 259, 270-271,

279, 301
plumbeum, Campostoma anomalum,

426, 431
plumbeus, Vireo solitarius, 493
Poa pratensis, 83, 535
pocket gopher, 88, 123, plains, 77,

138
Podiceps caspicus, 453
podiceps, Podilymbus, 170
Podilymbus podiceps, 170
Poecetes confinis, 508
Poecilichthys pulchellus, 430
poison ivy, 214, 268, 274
pokeweed, 257
polionotus, Peromyscus, 29
Polioptila

amoenissima, 449, 451, 488-89

caerulea, 263, 448, 488

melanura, 489
Polioptilinae, 452
Polistes, 148
polyglottus, Mimus, 255
Polygonum, 82
Polygyra, 31, 621
Polyodon spathula, 383
polyphemus, Gopherus, 338, 340-42
Pomaxis

annularis, 380, 385-86, 388-89, 391,
429. 432-33

nigromaculatus, 381, 381, 385-86,
429, 432-33
pomifera, Madura, 536
pondweed, 84
Ponera, 31

Pooecetes gramineus, 304
poor-will, 467

populations of ornate box turtle, 623
Populus deltoides, 409, 535
Porzana Carolina, 193, 461
post oak, 19
Potamogeton, 84

foliosus, 353, 361, 379

gramineus, 353

nodosus, 353. 370

694

University of Kansas Publs., Mus. Nat. Hist.

potasina, Aphelocoma sieberii, 480
potosinus,

Carpodacus mexicanus, 504

Pipilo fuscus, 507
poverty grass, 143
prairie

bluestem, 109, 118. 161

deer mouse, 145

dog, 467

falcon, 77, 189, 458

king snake, 51, 77, 123

skink, 77. 113

vole, 30, 67, 80, 94, 105, 116, 152,
160, 206, 321
pratensis, Poa, 83, 535
pratincola, Tyto alba, 465
predation and parasitism on six-lined

racerunner, 48
predators of ornate box turtle, 646
prickly

ash, 256, 535, northern, 535

lettuce, 159

pear, 536
Prionus fissicomis, 621
Procambaris gracilis, 621
Procyon lotor, 52, 164, 646
productus, Lepisosteus, 383
Progne

chalybea, 479

subis, 235, 479
promelas, Pimephales, 86, 372, 374,

385-86, 388, 414, 426, 432-33
propinquus, Turdus migratorius, 487
proportion of the tail and its regenera-
tion, six-lined racerunner, 43
prosopidicola, Cassidix mexicanus, 499
Prosopis, 8
Prunus

americana, 82, 259

serotina, 270, 536
psaltria, Spinus psaltria, 505
Psaltriparus

iulus, 451, 482

lloydi, 451, 482
Pseudablabes, 8
Pseudacris nigrita, 94
Psciidemys, 569, 571. 628-29

elegans, 571

scripta, 552, 554, 558, 561-63, 572,
574, 584, 611, 627
Pseudolcptodeira, 3-4
Psittacidac, 452
pnboscons, Dcndrocopos, 220
pulchclium, Etheostoma spectabile,

379, 430
puklifllus, Poecilichthys spectabilis,

430
punctata, Lissemys, 552
punctatus,

Diadophis, 116

Ictalnrus, 377, 385-386, 388-89,
427, 431-33, 438

punctulatum, Etheostoma, 393
punctulatus, Micropterus, 380, 385-86,

388-89, 393, 396
purple

fi^nch, 77, 298

martin, 77, 235, 479
purpureus, Carpodacus, 298
pusilla,

Spizella, 308

Spizella pusilla, 512

Wilsonia, 279
putorius, Spilogale, 52, 167
pygmaeus, Empidonax fulvifrons, 476
pygmy owl, 466
Pylodictis olivaris, 377, 385-86, 388-

89, 428, 432-33, 438
Pyralidae, 621
pvTalis, Photinus, 621
Pyrocephalus

flammeus, 451, 477

mexicanus, 448-49, 477
pyrrhonota, Petrochelidon, 235
Pyrrhuloxia

sinuata, 502

texana, 502
pyrrhuloxia, 502
Pyrus ioensis, 82
Pyxis arachnoides, 588

quail, 538

harlequin, 461

scaled, 448, 450-51, 459
Qucrcus, 409

marilandica, 353

rubra, 84

stellata, 353

velutina, 82, 353
querula, Zonotrichia, 313
Quiscalus quiscula, 286
quiscula, Quiscalus, 286

rabbit, 169, 177, 538, jack, 162
raccoon. 52, 80, 163-64, 169, 177,

646-47
racer, yellow-beUied, 79, 118, 321-22
racerunner, six-lined, 11-62, 69, 76, 78,

106, 321
ragweed, 85. 191, 242, 536, giant, 82,

93, 99, 155, 159. 166. 220, 241,

244, 296
rail,

Sora, 193

Virginia, 77, 193
Rallus limicola, 193
Rana

catesbeiana, 96, 537, 618

pipiens, 97, 537, 618
rat,

black, snake. 54. 79, 121, 238, 261,
321-22

cotton, 160, 163, 206-07, 321

grey, snake, 52

Index to Volume 11

695

rat, — Concluded

hispid cotton, 80, 149, 154

Norway, 80, 159

snake, 52, 54, 79, 121, 238, 261,
321-22
rattlesnake,

pigmy, 52

timber, 52, 77, 128
Rattus norvegicus, 159
raven,

common, 480

white-necked, 480, 646
Recurvirostra americana, 462
red

bat, 79, 134

cedar, 206, 249

fox, 77, 163, 322

haw, 82, 271

mulberry, 536

oak, 85, 203, 217, 271

shiner, 369, 390, 421
red-bellied woodpecker, 69, 80, 214,

218-219, 268, 319
red-breasted nuthatch, 77, 249
redbud, 82, 212, 265
red-eyed

towhee, 81, 300

vireo, 81, 271, 448, 493
redear sunfish, 384
redfin

darter, 384

shiner, 371, 382, 390, 420
red-headed woodpecker, 80, 216-17
redhorse,

golden, 364, 379

northern, 364, 418
red-shafted flicker, 470
red-shouldered hawk, 77, 183
redstart,

American, 77, 279, 497

painted, 497
red-tailed hawk, 77, 159, 180, 184,

187, 319-20, 457, 648
redwing, 85, 283, 321
redwinged blackbird, 77, 498
regalis, Buteo, 458
Regulus

calendula, 265, 489

cineraceus, 489-90

satrapa, 264, 489
Reithrodontomys, 30

megalotis, 52, 141

montanus, 140
relationships, six-lined racerunner, 17
reproduction in ornate box turtle, 543
Retinella, 621
rhombomaculata, Lampropeltis calli-

gaster, 124
Rhus

glabra, 84, 146, 536

trilobata, 127
Rhynchopsitta tenisi, 464

rice cutgrass, 141, 316

richardsonii, Miochanes richardsonii,

476
Richmondena

canicaudus, 502

cardinahs, 290
Ridgwayia pinicola, 487
ring perch, 430
ring-billed gull, 463
ring-necked

duck, 77, 176

eastern, snake, 116

snake, 79, 116
Riparia riparia, 233
ripaiia, Riparia, 233
river

carpsucker, 363, 416, 418, 435

shiner, 365

turtle, 317
RivoU's hummingbird, 469
roadrunner, 465
robber fly, 617

robin, 77, 214, 257, 286, 449, 487
Roccus clirysops, 384, 428, 432-33
rock wren, 485

rose-breasted grosbeak, 77, 293
rose-throated becard, 473
rostratum, Solanum, 536
rosyface shiner, 370, 420
rough-legged

American, hawk, 317

hawk, 77, 187
rough-vdnged swallow, 77, 233
rubellus, Notropis, 366, 368, 370, 372,

385-86, 388-89, 396-97, 420, 432-

33
rubescens, Anthus spinoletta, 490
rubra,

Euglena, 353

Moms, 536, 619

Piranga, 289

Piranga rubra, 448, 501

Quercus, 84

Ulmus, 535
rubrifrons, Notropis, 420
Rubus argutus, 84
ruby-crowned kinglet, 77, 265, 489
ruby-throated hummingbird, 77, 212,

469
ruficapilla, Vermivora, 274
ruficollis, Stelgidopteryx, 233
rufivirgata, Arremonops rufivirgata,

506
rufous-capped atlapetes, 449, 452, 505
rufous-crowned sparrow, 451, 509
rufous-sided towhee, 506
rufum, Toxostoma, 256
rustica, Hirundo, 235
rusty blackbird, 77, 286
ruticilla,

Setophaga, 279

Setophaga ruticilla, 497

696

Unr'ersity of Kansas Publs., Mus. Nat. Hist.

sacki, Cnemidophorus, 17, 20, 38

sage thrasher, 487

Sagittaria latifolia, 84, 353, 361

salamander, tiger, 77, 88, 317

Salix, 409, nigra, 536

salnioides,

Micropterus, 385-86, 388, 391, 432-
33

Micropterus salmoides, 380, 429
Salpinctes obsoletus, 485
salvini, Empidonax difficilis, 476
sandhill crane, 77. 192, 461
sand shiner, 368, 423
sandpiper,

least, 462

pectoral, 462

solitary, 77, 195

spotted, 77, 195, 462
sanguinalis, Digitaria, 85
sapsLicker, yellow-bellied, 77, 218, 471
Sarcophaga cistudinis, 643, 645-46,

662
Sarcophagidae, 621
satrapa,

Regulus, 264

Regulus satrapa, 489
sauger, 430
savannah sparrow, 508
savannanim, Ammodramus, 303
saw-whet owl, 77, 207
saxataUs, Aeronautes saxatalis, 463
saxatile, Etheostoma, 393
saya, Sayomis saya, 474
Sayomis

nigricans, 451, 474

phoebe, 224, 474

saya, 474

semiatra, 451, 474
Say's phoebe, 474

scalapsoides, Microtus pinetorum, 158
scaled quail, 448, 450-51, 459
Scalopus aquaticus, 30, 133, 156
scaly lizard, 52
Scaphiopus

bonibi Irons, 88

hammondi, 617
Scaphirhynchus platorhynchus, 410,

414. 432-33
Scarabaeidae, 621
scarabaeid beetle, 241, 620
Scardafella inca, 464
scariola, Lactuca, 82
scarlet tanager, 77, 288
Scarophaga, 646
scaup,

lesser. 77, 455

lesser, duck, 176
Sccloporus, 53

magistor, 22

undulatus, 52

wood!, 22
scissor-tailed flycatcher, 77, 222, 473

Sciurus

carolinensis, 136
niger, 137
Schreberi, Muhlenbergia, 535
sclateri,

Aeronautes saxatahs, 468
Parus sclateri, 481
Scolytus multistriatus, 221
scoparius, Andropogon, 85, 353, 536
Scott's oriole, 499
screech owl, 77, 200, 322, 465-66
scripta, Pseudemvs, 552, 554, 558,

561-63, 572, 574, 584, 611, 627
scrofa. Sus, 75
scrub

jay, 449, 479

oak, 353
sedge, 134, 536
Seiurus

aurocapillus, 275

motacilla, 276

noveboracensis, 276
Selasphorus platycercus, 469
semiatra, Sayomis nigricans, 451, 474
semidoliatus, Geophis, 333
Semotilus atromaculatus, 414, 419,

432-33
semplei, Otus asio, 466
sennetti, Toxostoma longirostris, 486
septendecim, Magicicada, 618
septentrionalis, Eumeces, 113
septentrionalium, Anas cyanoptera,

455
serotina, Prunus, 270, 536
serpentina, Chelydra, 98, 559. 611
sesquialis, Extrarius, 419
Setaria, 85, 316

lutescens, 536

viridis, 160, 315
Setophaga

picla, 497

ruticilla, 279, 497
sexlineatus, Cnemidophorus, 17-18, 21,
32, 106, 554

breeding, 32

biu-rows, 28

cycles of activity and temperature
relationship, 20

food habits, 31

growth, 39

habitat, 18

molt, 46

movements, 26

predation and parasitism, 48

proportion of the tail and its regen-
eration, 43

relationships, 17
sexual dimorphism of ornate box tur-
tle, 596
shad, 437, gizzard. 363, 415
shagbark hickory, 204, 248, 271
sharp-shinned hawk, 77, 178, 456

Index to Volume 11

697

sheep, 162
shiner,

Arkansas River, 368
bigeye, 365
bluntface, 367
common, 421
ghost, 366, 423
golden, 372
mimic, 372
plains, 369, 420
red, 369, 390, 421
redfin, 371, 382, 390, 420
river, 365
rosyface, 370, 420
sand, 368, 423
Topeka, 371, 423
shortnose gar, 363, 415
short-tailed shrew, 79, 130, 321

little, 79, 132
shoveller, 77, 175-76, 455
shovelnose sturgeon, 414, 435
shrew

little short-tailed, 79, 132
short-tailed, 79, 130, 321
shrike, loggerhead, 77, 85, 266, 490-91
Sialia

currucoides, 488
fulva, 488
mexicana, 488
siahs, 260
siahs, Sialia, 260
sicculus, Labidesthes, 378, 385-86,

388-91, 393, 396-97
side-necked terrapin, 588
side-oats grama, 141, 295, 536
Sigalphus, 31
Sigmodon hispidus, 149
silver chub, 419
silversides, brook, 378
silvery minnow, 423
sinuata, Pyrrhuloxia sinuata, 502
sinuatus, Corvus corax, 480
sipedon, Natrix, 113
sirtahs, Thamnophis, 114, 548
siskin, pine, 77, 299, 449, 505
Sistrurus

catenatus, 51
miliarius, 52
Sitonia hispidulus, 31
Sitta

canadensis, 249
carolensis, 248
melanotis, 482
mexicana, 451, 482
nelsoni, 451
six-lined racerunner, 11-62, 69, 76, 78,
106, 321
breeding, 32
burrows, 28
cycles of activity and temperature

relationship, 20
food habits, 31

sLx-lined racerunner — Concluded
growth, 39
habitat, 18
molt, 46
movements, 26
predation and parasitism, 48
proportion of the tail and its regen-
eration, 43
relationships, 17
skink, 52, 319, 623

five-hned, 67, 78, 109, 111, 584
Great Plains, 67, 78, 110, 187, 321
ground, 78, 109
prairie, 77, 113
skunk, 58

spotted, 80, 167, 322
striped, 52, 80, 167, 646-48
skunkbush, 127
slate-colored junco, 305
slender

fl.at-headed snake, 124
glass lizard, 78
slenderhead darter, 378
slider, 571, 584
slough grass, 353, 536
smallmouth buflFalo, 363, 416, 435, fish,

364
smart^veed, 82, 93, 134, 316, 392
smithsonianus, Larus argentatus, 462
smooth

earth snake, 77, 115
sumac, 536
snail, 622
snake,

black, 51

black rat, 54, 79, 121, 238, 261,

321-22
brown, 78, 114

common garter, 79, 114, 322
DeKay, 616

eastern ring-necked, 116
garter, 321, 548
grey rat, 52
hog-nosed, 52
king, 52, 75
milk, 77, 124
pine, 52

prairie king, 51, 77, 123
ring-necked, 79
slender flat-headed, 124
slender glass, 104
smooth earth, 77, 115
water, 317
worm, 79, 117
snapping turtle, 77, 98, 317
snipe, Wilson, 77, 195
snow goose, 77, 173
social wasp, 148
Solanum, 82, rostratum, 536
Solidago, 82

solitaire. Towmsend's, 488
solitaria, Tringa, 195

698

University of Kansas Publs., Mus. Nat. Hist.

solitarius,

Vireo, 271

Vireo solitarius, 492
solitary

sandpiper, 77, 195

vireo, 77, 271, 492-93
song sparrow, 82, 291
sora, 77, 461
sora rail, 193

Sorghastrum nutans, 85, 353, 536
Sorghum halepense, 360
sorghum, 191, 407
southern redbelly dace, 419
sowbug, 622
spadefoot,

plains, 88

plains, toad, 77

toad, 317
sparrow,

Baird's, 508

black-chinned, 512

black-throated, 448-49, 451, 509-
10

Brewer's, 511

Cassin's, 448, 509

chipping, 72, 77, 85, 308, 511

clay-colored, 77, 308, 511

English, 81, 85, 280

field, 68-69, 82, 85, 231, 308, 319,
511

fox, 77, 315

grasshopper, 77, 303, 508

Harris, 82, 291, 307, 313-14

house, 317, 497

lark. 77, 85, 304, 508

Leconte, 77, 304

Lincoln, 77, 315, 512

olive, 448, 506

rufous-crowned, 451, 509

savannah, 508

song, 82, 291, 313, 316

swamp, 77, 316, 513

tree, 82, 291, 307, 322

vesper, 77, 304, 508

white-crowned, 77, 313, 512

white-throated, 77, 307, 314

Worthen's, 512
sparrow hawk, 53, 77, 85, 189
Spartina pectinata, 353, 536
sparverius, Falco, 53, 189, 459
spathula, Polyodon, 383
Spatula clypcata, 175, 455
speckled chub, 365, 419
spectabile, Etheostoma, 370, 375, 385-

86, 388-89, 391. 395-96, 432-33
Speotyto hypugaea, 467
Spermophilus Irankliuii, 136
Sphingidae, 621
Sphyrapicus

nuchalis, 471

varius, 218, 471
spiderwort, 618

Spilogale putorius, 52, 167
spilopterus, Notropis, 393
spinifer, Trionyx, 520, 523-24
spinifera, Amyda spinifera, 524
Spinus

macropterus, 505

pinus, 299, 505

psaltria, 505

tristis, 299
Spiza americana, 295
Spizella

arborea, 307

arenacea. 511

arizonae, 511

atrogularis, 512

breweri, 511

pallida, 308, 511

passerina, 308

pusilla, 308, 512

wortheni, 512
sponsa, Aix, 175
spotted

bass, 380

gar, 383

sandpiper, 77, 195, 462

skunk, 52, 80, 167, 322

sucker, 365
spruce, 449
spurge, 82-83
spurius. Icterus. 284, 498
squamata, Callipepla squamata, 451,

459-60
squirrel, 146, 169

fox, 69, 79, 137, 246, 321-22

gray, 69, 79, 136, 321-22

tree, 538
starling, 77, 216, 267, 282, 286, 317
starred tortoise, 552
Steganopus tricolor, 462
Stelgidopteryx ruficollis, 233
stellata, Quercus, 353
Steller's jay, 449, 479
stephensi, Vireo huttoni, 449, 492
Stemotherus odoratus, 548, 559, 561,

624, 649
Stizostedion

canadense, 430, 432-33

vitreum, 430
stonecat, 383, 428
stoneroller, 375, 379, 426
Storeria dekayi, 114, 616
storeriana, Hybopsis, 419, 432-33
streams of the Big Blue Basin, 407
strepera, Anas, 454
striata,

Dendroica, 275

Lespedeza, 83
striatus,

Accipiter, 178

Cyathus, 622
strigatus, Chondestes grammacus, 508
striped skunk, 52, 80, 167, 646-48

Index to Volume 11

699

Strix varia, 203

sturgeon, 354, shovelnose, 414, 435

Stumella

hoopesi, 498
magna, 281
neglecta, 282, 498
Sturnus vulgaris, 267
subis,

Progne, 235
Progne subis, 479
Succinia, 621
sucker,
blue, 416

deep-bodied, 388-89, 395-96
spotted, 365
white, 418
suckermouth minnow, 372, plains, 420
sulcirostris, Crotaphaga sulcirostris,

465
sumac, 84, 146
fragrant, 301
smooth, 536
summer tanager, 67, 81, 289, 320, 448-

49, 451, 501
sunfish, 421-22, 437

green, 77, 381, 390, 392, 429
longear, 382
orangespotted, 381, 429
redear, 884
sunflower, 82, 85, 99, 166, 241, 298-99
Sus scrofa, 75
suttoni,

Accipiter striatus, 456-57
Otus asio, 465-66
swainsoni, Buteo, 457
Swainson's hawk, 457
swallow,

bank, 77, 233
barn, 81, 85, 234, 322, 478
cave, 448, 478
cliff, 77, 235, 478
rough-winged, 77, 233
tree, 478

violet-green, 451, 477
swamp sparrow, 77, 316, 513
sweet clover, 191
swift,

chimney, 77, 211
white-throated, 468
switch grass, 85, 141, 295, 536
sycamore, 85, 165, 270-71, 409
Sylviidae, 452

Sylvilagus floridanus, 54, 134, 622
Symphoricarpos orbiculatus, 535
symplectus, Dendrocopos scalaris, 448,

472
Systematic status of the colubrid snake,
Leptodeira discolor Giinther, 3

Tachycineta

lepida, 451, 477
thalassina, 451, 478

Tachymenis, 8
tanager,

hepatic, 501

scarlet, 77, 288

summer, 67, 81, 289, 320, 448-49,

451, 501
western, 501
Tantilla graciUs, 124
Tantalophis, 3, 8-9

discolor, 4-7
tarbush, 540
taurus, Bos, 75
Taxidea taxus, 646
taxus, Taxidea, 646
teal

blue-winged, 77, 175, 455, 618
cinnamon, 455

green-winged, 77, 175-76, 455
tegu, 17
Teiidae, 17

Telmatodytes plesius, 485
temperature relationships of ornate

box turtle, 598
tenellus, Pimephales, 372-74, 385-86,

388-89, 393, 397
Tennessee warbler, 77, 273
tentrionaUs, Chloroceryle americana,

470
tenuirostris, Aimophila ruficeps, 451,

509
tem, 317

Terrapene, 531-32, 588-89
fossils, 534

Carolina, 99, 449, 532-35, 539, 543,
545, 548, 551, 553-54, 557-59,
561-63, 569, 572, 585, 595-96.
611-12, 616, 622-23, 626-28,
630, 634, 636-38, 640, 643, 646-
48, 650-56
coahuila, 532, 654
klauberi, 532, 650, 652-53
longinsulae, 534, 656
luteola, 372, 449, 533-34, 540-43,
597, 617-18, 629-30, 654-55,
657
mexicana, 532, 654
nelsoni, 532, 650
omata, 51, 54, 99, 527-669
triunguis, 628
yucatana, 532, 654
terrapin,

diamond-backed, 552
land, 534
side-necked, 588
terrapin, Malaclemys, 552, 561-62
terrestris, Bufo, 89
terrisi, Rhynchopsitta temsi, 464
tessellatus, Cnemidophorus, 20, 31, 38
Testudinidae, 532, 588, 638
Testudo, 656, horsefieldi, 600
teter, Cathartes aura, 456

roo

University of Kansas Publs., Mus. Nat. Hist.

tetranemus,

E.xtrarius, 419

Hybopsis aestivalis, 365, 394-95,
419
Teucrium canadense. 83, 212
texana, Pyrrhuloxia sinuata, 502
texanus,

Colinus virginianus, 448, 459

Pipilo fuscus, 507
texensis, Chordeiles acutipennis, 468
thalassina, Tachycineta thalassina, 451,

478
Thamnophis

elesans, 548

sirtalis, 114, 548
Thapsus, Verbascum, 536
theophrasti, Abutilon, 82
thick-billed parrot. 449, 464
thistle, 82, 159
thrasher,

brown, 81, 256

Crissal, 451, 487

curve-billed, 448, 451, 486

long-billed, 448, 486

sage, 487
three-awn grass, 84, 106, 140, 304
thrush,

Aztec, 452, 487

gray-backed, 77

gray-cheeked, 259

hermit, 77. 259, 487

northern, 77. 276

olive-backed, 77, 259

wood, 68, 71, 77, 258, 319
Thryomanes

cryptus, 484

eremophilus, 484

murinus, 484
Thryothorus

berlandieri, 484

ludovicianus, 252
tiger

beetle, 31

salamander, 77, 88, 317
tigrinum, Ambystoma, 88
tigris, Cnemidophorus, 17, 20-21, 28,

31, 38
Tilia americana, 85
timber rattlesnake, 52, 77, 128
titmouse, 249, 318, 320, 322

black-crested, 448-49, 451, 481

tufted, 81, 221, 244-45, 250, 265,
319
toad, 93, 321, 618

American, 78, 165, 318

common American, 89

Great Plains narrow-mouthed, 78,
95

narrow-mouthed, 67, 93, 318

plains spadefoot, 77

spadcfoot, 317

M'oodhouse's, 78, 85, 90, 317-18

Tomodon, 8

Topeka shiner, 371, 423

topeka, Notropis, 371, 410, 423, 432-

33
topminnow, black-banded, 376
tortoise,

desert, 600, 611

Galapagos, 342

land, 534

starred, 552
Totanus

flavipes, 195, 462

melanoleucus, 462
towhce, 307, 320, 322

brown, 507

green-tailed, 506

red-eyed, 81, 300

rufous-sided, 506
townsendi,

Dcndroica, 495

Myadestes towTisendi, 488

Townsend's

solitaire, 488

warbler, 495
Toxostoma

celsum, 451, 486

dorsale, 451, 487

dumosum, 451, 487

oberholseri, 448, 486

rufum, 256

sennetti, 486
Tradescantia, 618
Traill flycatcher, 475
traillii, Empidonax, 229
tree

duck, black-bellied, 454

sparrow, 82, 291, 307, 322

squirrel, 538

swallow, 478
treefrog, gray, 78, 93
treganzai, Ardea herodias, 453
trepidus, Empidonax aflfinis, 475
trichas, Geothlypis, 277
tridentata, Larrea, 462
trifida. Ambrosia, 82
trilobata, Rhus, 127
Tringa solitaria, 195

Trionyx

calvatus, 519-520

ferox, 520, 523

japonicus, 552

muticus, 519-20, 522-25

spinifer, 520, 523-24
Tripsacum dactyloides, 353
tristis, Spinus, 299
triunguis, Terrapene Carolina, 628
Trochilidae, 453
Troglodytes

aedon, 251

cahooni, 451, 483

compositus, 451, 483

Index to Volume 11

701

Troglodytes — Concluded

parkmanii, 483

troglodytes, 251
troglodytes. Troglodytes, 251
Trombicula, 662

alfreddugesi, 643-44

alfreddugesi, 54

lipovskyana, 644

montanensis, 644
trout, 354
Trypanurgos, 8
tufted titmouse, 81, 221, 244-45, 250,

265, 319
Tupinambis, 17
Tiurdidae, 452
Turdus

migratorius, 257

propinquus, 487
turkey, 450, 461

turkey vulture, 77, 176, 322, 456
turtle,

box, 51, 54, 58, 538

eastern, box, 77, 99

ornate, box, 322, 527-669

western, box, 78, 99

marine, 552

musk, 548

painted, 77, 101, 317, 557, 585, 643

river, 317

snapping, 77, 98, 317
Tuttle Creek Dam and Reservoir, 404
Typha, 277

angustifolia, 158

latifolia, 158, 353
Tyrannidae, 453
Tyrannus

tyrannus, 222

vociferans, 473
tyrannus, Tyrannus, 222
Tyto

alba, 200

pratincola, 465

Ubnus

americana, 259

americanus, 82, 409, 535

rubra, 535
umbratilis.

Notropis, 371, 379, 383, 385-86,
389-92, 386-97, 432-33

Notropis umbratilis, 420
undulatus, Sceloporus, 52
upland plover, 77, 185
Urban, Emil K., Birds from Coahuila,

Mexico, 445
Urocyon cinereoargenteus, 164
ustulata, Hylocichla, 259

valeriae, Haldea, 115
varia,

Mniotilta, 272, 493

Strix, 203

varied bunting, 503
varius,

Sphyrapicus, 218
Sphyrapicus varius, 471
veery, 77, 260

veliei, Contopus sordidulus, 476
velifer

Caipiodes, 363. 394, 417
Ictiobus, 417
velox, Accipiter striatus, 456-57
velutina, Quercus, 82, 353
velv.etleaf, 82
Verbascum Thapsus. 536
Verbena, 83, hastata, 536
Vernonia baldwini, 83
vermilion flycatcher, 448-49, 451, 477
verdin, 481
Vermivora

celata, 273, 493

crissalis, 494

mexicana, 494

orestera, 494

peregrina, 273

ruficapilla, 274

virginiae, 494
versicolor,

Hyla, 93

Passerina versicolor, 503
Vei-tebrata, 621
vervain, 83, blue, 536
vesper sparrow, 77, 304, 508
vigilax, Pimephales, 372-73, 385-86,

388-89
violacea, Nyctanassa violacea, 454
violet-green swallow, 451, 477
villosus, Dendrocopos, 218
virens,

Contopus, 230, 476

Dendroica, 275, 496

Icteria. 278, 496
Vireo

atricapilla, 491

bellii, 270

carolinae, 448, 451, 492

cassinii, 493

flavifrons, 271, 492

gilvus, 272, 493

griseus, 268

medius, 448-49, 492

micrus, 491

noveboracensis, 491

olivaceus, 271, 493

plumbeus, 493

soUtarius, 271, 492

Stephens!, 449, 492
vireo.

Bell, 81, 270-71, 321

Bell's, 448-49, 492

black-capped, 491

Mutton's, 448-49, 451, 492

red-eyed, 81, 271, 448, 493

solitary, 77, 271, 492-93

702

Uniyersity of Kansas Publs., Mus. Nat. Hist.

vireo, — Concluded

warbling, 77. 272, 493

white-eyed, 81, 268, 272, 491

yellow-throated. 77, 271, 448, 492
virescens, Butorides. 171, 454
virgatum, Panicum, 85, 353, 536
Virginia

creeper, 273

rail, 77, 193
virginiae, Vermivora, 494
virginiana,

Dama. 168

Juniperus, 249
virginianus.

Bubo, 201

Colinus, 190
Virginia's warbler, 494
virldis, Setaria, 160, 315
Vitis vulpina, 257
vitreum, Stizostedion, 430
vociferans, Tyrannus vociferans, 473
vociferus.

Caprimulgus, 209

Charadrius. 194

Charadrius vociferus, 461
vole, 71, 149, 165, 204, 207, 321

pine, 80, 131, 154-55

prairie, 30, 67, 80, 94, 105, 116,
152, 160, 206, 321
volucellus, Notropis, 367, 372. 385-86,

388, 393-94, 396-97
vulgare, Armadillidium, 621
vulgaris, Sturnus, 267
Vulpes fulva, 163
vulpina, Vitis, 257
vulture,

black, 450, 456

turkey, 77, 176, 322, 456

wagleri Icterus wagleri, 499

Wagler s oriole, 499

walleye, 430, 437-38

walnut, 137, 260, 264, 289, 353, 489,

black, 271, 354, 535
wapato, 84
warbler,

Audul)on's. 495

black and white, 77, 272, 493

Blackburnian, 77, 275

blackpoll, 77, 275

black-throated gray, 495

black-throated green, 77, 275, 496

Canada, 77, 279

Colima, 279

golden-cheeked, 496

Ilartlaub's, 449, 494

hermit, 496

Kentuckv, 68, 81, 276, 320

myrtle, 77, 245, 274

Nashville, 77, 274

olive, 494

olive-backed. 494

warbler, — Co ncluded

orange-crowned, 77, 273, 493

Parula, 494

Tennessee, 77, 273

Townsend's, 495

Virginia's, 494

Wilson, 77, 279

Wilson's, 497

yellow, 77, 274, 495
warbhng vireo, 77, 272, 493
wardi, Ardea herodias, 453
warmouth, 384
warneri, Pharyngodon, 53, 58
wasp, social, 148
water

snake, common, 77, 113

pipit, 490

thrush, Louisiana, 77, 276

snake, 317
waxwing, cedar, 77, 265, 490
wear on shell of ornate box turtle, 595
weasel, long-tailed, 77, 166
Webb, Robert G., Description of a new

softshell turtle from the southeast-
em United States, 519
weed, Joe-Pye, 82
weight of ornate box turtle, 586
western

bluebird, 488

box turtle, 78, 99

flycatcher, 476

harvest mouse, 79, 140-141, 154

meadowlark, 77, 282, 498

tanager, 501

wood pewee, 476
wheat, 85, 232, 295, 407, 535
whipplii, Etheostoma, 384, 393
whip-poor-'will, 80, 208-09, 322, 467
white

bass, 384, 428, 437

crappie, 380, 429

pehcan, 77, 170, 453

pine, 469, 496

sucker, 418
white-breasted nuthatch, 77, 248, 451,

482
white-crowned sparrow, 77, 313, 512
white-eved vireo, 81, 268, 272. 491
white-footed mouse, 80, 143, 147, 160,

207-208, 321
white-fronted

dove, 464

goose, 77, 173
white-necked raven, 480, 646
white-tailed deer, 77, 168, 322
white-throated

sparrow, 77, 307, 314

swift, 468
white-winged dove, 463
whitneyi, Micrathene, 467
widgeon, American, 455
Wicd's crested flycatcher, 450, 474

Index to Volume 11

703

wild

cherry, 270
lettuce, 82

plum, 82, 259, 270-71, 279, 301
willow, 19, 84, 134, 216, 229, 264-66,

293, 353, 409, 502, black, 536
Wilsonia

canadensis, 279
pileolata, 497
pusilla, 279
Wilson

snipe, 77, 195
warbler, 77, 279
Wilson's

phalarope, 462
warbler, 497
winter wren, 77, 251
wood duck, 77, 175
woodchuck, 77, 136, 317, American,

77
woodcock, American, 194
woodhousei, Bufo, 90
Woodhouse's toad, 78, 85, 90, 317-18
woodi, Sceloporus, 22
woodland,

oak-hickory, 85, 263
deer mouse, 145
woodpecker,
acorn, 471
downv, 69, 80, 215, 219-20, 245,

250, 265, 319
golden-fronted, 448-49, 470-71
hairy, 80, 214-15, 218, 319, 322,

448, 450, 471
ladder-backed, 448-50, 472
red-bellied, 69, 80, 214, 218-19,

268, 319
red-headed, 80, 216-17
wood

pewee, 72: eastern, 80, 230, 476;

western, 476
thrush, 68, 71, 77, 258, 319
woodrat, 71, 116, 135, 146, 148, 178,
318, 321
eastern, 67, 80, 151
woods, oak-hickory, 167
worm snake, 79, 117
wortheni, Spizella wortheni, 512
Worthen's sparrow, 512
wren,

Bewick's, 484
brown-throated, 451, 483
cactus, 449, 451, 484
caiion, 485

Carolina, 81, 252, 319, 448, 484
house, 77, 251, 483
long-billed marsh, 485
rock, 485

wren, — Concluded

short-billed marsh, 77, 255

winter, 77, 251
wrightii, Empidonax, 475
Wright's flycatcher, 475

Xanthocephalus xanthocephalus, 282,

498
xanthocephalus, Xanthocephalus, 282,

498
Xanthoxylum americanum, 535
Xenodontinae, 7
x-punctata, Hybopsis, 393

>ears unfavorable for growth of ornate

box turtle, 582
vellow

bullhead, 377, 427

perch, 430

pike, 430

warbler, 77, 274, 495
yellow-bellied

flycatcher, 77, 229

racer, 51, 79, 118, 321-322

sapsucker, 77, 218, 471
yellow-billed cuckoo, 80, 198-99, 448,

464
yellow-breasted chat, 279, 496
yellow-crowned night heron, 454
yellow-headed blackbird, 77, 282, 498
yellowlegs,

greater, 462

lesser, 77, 195, 462
vellow-shafted flicker, 80, 213
yellowtliroat, 77, 277, 496, hooded,

449, 496
yellow-throated vireo, 77, 271, 448,

492
yucatana, Terrapene, 532, 654
yucca, 448, 484, 508, 540

Zapus hudsonius, 161
zebrinus, Fundulus, 376
Zenaida

asiatica, 463

mearnsi, 463
Zenaidura

carolinensis, 463

macroura, 196

marginella, 463
zibethicus. Ondatra, 158
zone-tailed hawk, 457
Zonotrichia

albicollis, 314

gambelii, 512

leucophrys, 313, 512

oriantha, 512

querula, 313
Zygonectes notatus, 376

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA. KANSAS

I960

28-4430

(Continued from inside of front cover)

Vol. 8. 1. Life history and ecology of the five-lined skink, Eumeces fasciatus. By Henry
S. Fitch. Pp. 1-156, 26 figures in text. September 1, 1954.

2. Myology and serology of the Avian Family Fringillidae, a taxonomic itudy.
Bv William B. Stallcup. Pp. 157-211, 23 figures in text, 4 tables. November
15, 1954.

3. An pcologicnl study of the collared Uzard (Crotaphytiis coUaris). By Henry
S. Fifch. Pp. 213-274, 10 figures in text. February 10, 1956.

4. A field studv of the Kansas ant-eating frog, Gasfrophr>-ne olivacea. By Henry
S. Fitch. Pp. 275-306, 9 figures in text. February 10, 1956.

5. Check-list of the birds of Kansas. By Harrison B. Tordoff. Pp. 307-359, 1
figure in text. March 10, 1956.

fl. A population studv of the prairie vole (Microtus ochrogaster) in northeastern
Kansas. By Edwin P. Martin. Pp. 361-416, 19 figures in text. April 2,
1956.

7. TemptTatiire responses in free-livine amphibians and reptiles of northeastern
Kansas. By Henry S. Fitch. Pp. 417-476, 10 figures in text, 6 tables. June
1, 19.58.

8. Food of the crow. Cor\-us brach%Th\Tichos Brchm, in south-central Kansas. By
Dwighf Piatt. Pp. 477-498, 4 tables. June 8, 1956.

9. Ecological obsor\ntions on the woodrnf, Neotoma floridana. By Henry S.
Fitch and Dennis C. Rainey. Pp. 499-533, 3 figures in text. June 12, 1956.

10. Eastern woodrat, Nrotoma floridana: Life history and ecology. By Dennis
G. Bainey. Pp. 535-646, 12 plates, 13 figures in text. August 15, 1956.

Index. Pp. 647-675.
Vol. 9. 1. Speciation of the wandering shrew. By James S. Findle>'. Pp- 1-68, 18 fig-
ures in text. December 10, 1955.

2. Additional records and extensions of ranges of mammals from Utah. By
Stephen D. Durrnnt. SI. Raymond Lee. and Richard M. Hansen. Pp. 69-80.
December 10, 1955.

8. A new long-eared myotis (Mvotis evotis) from northeastern Mexico. By Rol-
lin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.

4. Subspeciation in the meadow mouse. Klicrotus pcnnsylvanicus, in Wyoming.
By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 19.56.

5. The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6
figures in text. May 19, 1956.

6. Additional remains of the multituberculafe geni« Eucosmodon. By Robert
W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.

7. Mammals of Coahuila. Mexico. By Rollin H. Baker. Pp. 125-335, 75 figurw
in text. June 15, 1956.

8. Comments on the taxonomic status of Apodemus peninsulae, with description
of a new subspecies from North China. Bv J. Knox Jones, Jr. Pp. 337-346,
1 figure in text, 1 table. August 15, 1956.

9. Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp.
347-351. Augiist 15, 1956.

10. A new bat (Genus Leptonvcferis) from Coahuila. By Howard J. Stains.
Pp. 353-356. January 21, 1957.

11. A new species of pocket gopher (Genus Pappogeomvs) from Jalisco, Mexico.
By Robert J. Russell. Pp. 357-361. January 21, 1957.

12. Geographic variation in the pocket gopher, Thomomys bottae, in Colorado.
By Phillip M. Youngman. Pp. 363-387, 7 figures in text. February 21,
1958.

13. New bog lemming (genus Svnaptomys) from Nebraska. By J. Knox Jones,
Jr. Pp. 385-388. May 12, 1958.

14. Pleistocene bats from San Josecito Cave, Nuevo Leon, Mexico. By J. Knox
Jones, Jr. Pp. 389-396. December 19. 1958.

15. New Subspecies of the rodent Baiomys from Central America. By Robert L.
Packard. Pp. 397-404. December 19, 1958.

16. Mamm.als of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405-
414. 1 figure in te.xt. .May 20, 1959.

17. Distribution, variation, and relationships of the montane vole, Microtus mon-
tanus. Bv Svdney Anderson. Pp. 415-511. 12 figures in text, 2 tables.
Aug\ist 1, 1959.

18. Conspecificitv of t^vo pocket mice, Perognathus goldmani and P. art\is. By
E. Raymond Hall and Mari]\-n Bailey Ogilvie. Pp. 513-518, 1 map.
January 14, 1960.

(Continued on outside of back cover)

(CoDtinued from inside of back cover)

19. Records of harvest mice, Reithrodontomys, from Central America, with
description of a new subspecies from Nicaragua. By Sydney Anderson and
J. Knox Jones, Jr. Pp. 519-529. January 14, 1960.

20. Small c.-univores from San Josecito Cave (Pleistocene), Nuevo Leon, Mexico.
By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, 1960.

21. Pleistocene pocket gophers from San Josecito Cave, Nuevo Leon, Mexico.
By Robert J. Russell. Pp. 539-548, 1 Bguie in text. January 14, 1960.

22. Review of the insectivores of Korea. Bv J. Knox Jones, Jr., and David
H. Johnson. Pp. 549-578. February 23, 1960.

More numbers will appear in volume 9.

VoL 10. 1. Studies of birds killed in nocturnal migration. By Harrison B. Tordoff and
Robert M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 1956.

2. Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.

3. The forest habitat of the University of Kansas Natural History Reservation.
By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figure*
in text, 4 tables. December 31, 1956.

4. Aspects of reproduction and development in the prairie vole (Microtia ochro-
gaster). By Henry S. Fitch. Pp. 129-161, 8 figures in text, 4 tables. Decem-
ber 19. 1957.

5. Birds found on the Arctic slope of northern Alaska. By James W. Bee.
Pp. 163-211, pis. 9-10, 1 figure in text. March 12, 1958.

6. The wood rats of Colorado: distribution and ecology. By Robert B. Finley,
Jr. Pp. 213-552, 34 plates, 8 figures in text, 35 tables. November 7. 1958.

7. Home ranges and movements of tlie eastern cottontail in Kansas. Bv Donald
W. Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, 1959.

8. Natural histor>' of the salamander, Aneides hardvi. By Richard F. Johnston
and Schad Gerhard. Pp. 573-585. October 8, 1959.

Mors numbers will appear in volume 10.

Vol. 11. 1, The svstematic status of the colubrid snake, Leptodeira discolor Giinther.
By William E. Duellman. Pp. 1-9, 4 figs. July 14. 1958.

2. Natural history of the six-lined racerunner. Cnemidophorus sexlincatus. By
Henry S. Fitch. Pp. 11-62, 9 figs., 9 tables. September 19, 1958.

3. Home ranees, territories, and season.il movements of vertebrates of the
Natural History Reservation. By Henry S. Fitch. Pp. 63-326, 6 plates, 24
figures in text, 3 tables. December 12, 1958.

4. A new snake of the genus Geophis from Chihuahua, Mexico. By John M.
Legler. Pp. 327-334, 2 figures in text. January 28, 1959.

5. A new tortoise, genus Gopherus, from north-central Mexico. By John M.
Legler. Pp. 335-343. April 24, 1959.

6. Fishes of Chautauqua, Cowley and Elk counties, Kansas. By Artie L.
Metcalf. Pp. 345-400, 2 plates, 2 figures in text, 10 tables, ^fay 6, 1959.

7. Fishes of the Big Blue River Basin, Kansas. By W. L. Minckley. Pp. 401-442.
2 plates, 4 figures in text, 5 tables. May 8, 1959.

8. Birds from Coahuila, Mexico. By Emil K. Urban. Pp. 443-516. August 1,
1959.

9. Description of a new softshell turtle from the southeastern United States.
By Robert G. Webb. Pp. 517-525, 2 pis., 1 figure in text, August 14, 1959.

10. Natural history of the ornate box turtle, Terrapene ornata ornata Agassiz. By
John M. Legler. Pp. 527-669, 16 pis., 29 figures in text. March 7, 1960.
Index will follow.
Vol. 12. 1. Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry
A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text, July 8, 1959.

2. The ancestry of modern Amphibia: a review of the evidence. By Theodore
H. Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, 1959.

3. The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49
figures in text. February 19, 1960.

More numbers will appear in volume 12.

Date Due