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UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORMUS. COM. ZOOL
VOLUME 12 + 1959-1964
EDITORS
E. RayMonp Hat, Chairman
THEODORE H. EATON, JR.
Henry S. Fircu
ROBERT W. WILSON
MusEuM oF NATURAL HISTORY
UNIVERSITY OF KANSAS
LAWRENCE
1964
LIBRARY,
DEC 31 {965
HARVARD
UNIVERSITY
gS ~VA -L Bere
Museum oF NATURAL HISTORY
UNIVERSITY OF KANSAS
LAWRENCE
PRINTED BY
HARRY (BUD) TIMBERLAKE, STATE PRINTER
TOPEKA, KANSAS
1964
30-3964A
13.
14.
15.
CONTENTS OF VOLUME 12
Functional morphology of three bats: Eumops, Myotis, Macrotus. By
Terry A. Vaughan. Pp. 1-153, pls. 1-4, 17 figs. July 8, 1959.
The ancestry of modern Amphibia: a review of the evidence. By Theo-
dore H. Eaton, Jr. Pp. 155-180, 10 figs. July 10, 1959.
The baculum of microtine rodents. By Sydney Anderson. Pp. 181-216,
49 figs. February 19, 1960.
A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By
Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figs.
May 2, 1960.
Natural history of the Bell Vireo, Vireo bellii Audubon. By Jon C. Barlow.
Pp. 241-296, 6 figs. March 7, 1962.
Two new pelycosaurs from the Lower Permian of Oklahoma. By Richard
C. Fox. Pp. 297-307, 6 figs. May 21, 1962.
Vertebrates from the Barrier Island of Tamaulipas, México. By Robert K.
Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp.
309-345, pls. 5-8. June 18, 1962.
Teeth of edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 figs.
October 1, 1962.
Variation in the muscles and nerves of the leg in two genera of grouse
(Tympanuchus and Pedioecetes). By E. Bruce Holmes. Pp. 363-474,
20 figs. October 25, 1963.
. A new genus of Pennsylvanian fish (Crossopterygii, Coelacanthiformes )
from Kansas. By Joan Echols. Pp. 475-501, 7 figs. October 25, 1963.
. Observations on the Mississippi Kite in southwestern Kansas. By Henry
S. Fitch. Pp. 503-519. October 25, 1963.
. Jaw musculature of the Mourning and White-winged doves. By Robert L.
Merz. Pp. 521-551, 22 figs. October 25, 1963.
Thoracic and coracoid arteries in two families of birds, Columbidae and
Enerueae. By Marion Anne Jenkinson. Pp. 553-573, 7 figs. March 2,
The breeding birds of Kansas. By Richard F. Johnston. Pp. 575-655,
10 figs. May 18, 1964.
The adductor muscles of the jaw in some primitive reptiles. By Richard
C. Fox. Pp. 657-680, 11 figs. May 18, 1964.
Index, Pp. 681-694.
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AUG- 61959
7 AD
UMIVERSITY
UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY
Volume 12, No. 1, pp. 1-153, 4 pls., 17 figs.
July 8, 1959
245 4
ix)
lia
Functional Morphology of Three Bats:
Eumops, Myotis, Macrotus
BY
TERRY A. VAUGHAN
UNIVERSITY OF KANSAS
LAWRENCE
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 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 figures in text. April 9, 1948.
Vol. 3. *l. 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 figures in text. June 29, 1951.
8. Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473-
530, 49 figures in text, 18 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,
1 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. 1-803, 73 figures in
text, 37 tables, August 25, 1952.
2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry
WY hk and Lewis L. Sandidge. Pp. 305-338, 5 figures in text. August
24, 1953.
The silky pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker.
Pp. 389-347, 1 figure in text. February 15, 1954.
North American jumping mice (Genus Zapus). By Philip H. Krutzsch, Pp.
349-472, 47 figures in text, 4 tables. April 21, 1954.
Mammals from Southeastern Alaska. By Rollin H. Baker and James S.
Findley. Pp. 478-477. April 21, 1954.
Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479-
487. April 21, 1954.
Subspeciation in the montane meadow mouse, Microtus montanus, in Wyo-
ming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text.
July 28, 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. 588-586. 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 Coahuila,
pee. By Robert J. Russell and Rollin H. Baker. Pp. 591-608. March
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.
ES eA A?
(Continued on inside of back cover)
AUG - 6 1959
J
UNIVERSITY
. UNIVERSITY OF KANSAS PUBLICATIO
\ MusEUM OF NATURAL HISTORY
Volume 12, No. 1, pp. 1-153, 4 pls., 17 figs.
July 8, 1959
Functional Morphology of Three Bats:
Eumops, Myotis, Macrotus
BY
TERRY A. VAUGHAN
UNIVERSITY OF KANSAS
LAWRENCE
1959
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL History
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson
Volume 12, No. 1, pp. 1-153, 4 pls., 17 figs. in text, 3 tables
Published July 8, 1959
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED IN
THE STATE PRINTING PLANT
TOPEKA, KANSAS
1959
Functional Morphology of Three Bats:
Eumops, Myotis, Macrotus
BY
TERRY A. VAUGHAN
CONTENTS
FENSTRODUGEION Us. 5,5 cast co ayteiine fr. Oe ates autare teaet ee tarsL Aran, <0sicptoee
Materials and. Methods 26-0 bs sow tarps Se ae chews Sle ccs eae
Remarksyom, Bats SEUGIed 926 oe sien he eek ee
TEOCOMOTOR ADEHAVIOR® 327 v0e. hao hin aa ea ose See ie he eso
BUENOS s PELOUS: . aa Worn a itn es eet ance a een rete
Roosting Habits and Terrestrial Locomotion ............
oraring, Habits and, PMolt 02 Sanat Sine ta ese
IVY GUIS VC EER ax Sica See, ten re es ae mea en ie ee
Roosting Habits and Terrestrial Locomotion ............
Boragine. Habitscand PMP «20g ae ie fees We oe
INIACTOLUS: CaltOrmiCUSe oe fa cnt es oes aig oc ae
Roosting Habits and Terrestrial Locomotion ............
Porapine slapits andy WMOME oes 49 tae ata eee ee ne
AERODYNAMIC» CONSIDERATIONS. «2505 ols ape oid oa Od Boo
OSTEO OG ant eee Ln etna ay asa ee ON eer rhe ee
ETILFOCUCLOLYANCIMATKS! 4c) cis is On tee an Peeve tei rato
Werte praia GOlMmin taka Hee sees aa Rome ak tea renin sen ee
SCO TITAN Cache oe eee ek Ate a Ber Hg Ue ee nc owe ce eas Meeerave Steed es Zoo a
ITED S 5 epee Sips ee PN ees cfd: A Dene em La A areas tn eS
RectoraliGirdleand:Ihimb--.= 225 hereon ee cee ee ae
Seo Mee or tary co ae Sell fo Sat perenne Mea e arte cot
ave ens ahs Seca ok harm Fork Mina cep ee ene te oy aeaaa ree ty Deh te
INA ATAU le ee oe eet Ro ty al OR
Pelvic Cirdievand, limber. 6.2 85 se a oe
nnomimnates DONE as eo oS aS te Sele
GUUS EES 2 on Oe a sR ee
WabiaganaeLibimlaw meee aon oho pets ae A ero
PES eee ss ee I eRe een Das yeas
4 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. HIst.
PAGE
NIVOROGY AM eet «Cae oie tioe Ratha oo CugOs eee Rae: 67
Introductory Remarks (eet rs 3 Se ea SUM ems eer 67
MuscleseUnique:to Bats’ 2<.).\..08% 40. 5. et ecce eet a 67
Muscles of the Pectoral Girdle and Limb ................ 70
MrapezsuswGroupy ©. eo cies wes | iad ose a PORE 70
€osto-spino-seapular ‘Group 22) 6.255... sas eet aa
Ratissimus-subscapular ‘Group’ ...2..5 7.0.6.9 ee 73
Deltorde Groupe sean soot has ot She wees ee eee 75
Supraseapular ‘Group.'.25. a6. 8 ai. Soe eee CHE
MmecepsiGroupy! & i555 5.04 coin? cos ss 3 Ye oe ep eee 78
Fixtensor Group or MOrearm,. 45.2. «52 5: ) one a ee ee 80
IPectOralise Groups)... elds yk oe or > Se ee 85
Blexor: Group.of Arm) |. 64.00.24 «snd Ge pen en 88
MlexonsGroupsof Porearm: (ibs. c.cce2 epee 91
HxtensorGroupsor Manus? jc.) (nace os ee ne ee 94
nlexorn Grouprol Wanus)a:0)2 65.5 ¥2 ot A aoe eee 94
Muscles of Pelvic Girdle and Limb ...:.... .20..2:22.:-%- 97
MI ACUS MG TOU ID 4 iy ree ee les sche frac Wat MEM beg ne 98
Cluteale Groupsets hens riasite soe eh oat epee 100
Ouadriceps, Memos; Groupp. 35.2.2. h ae ca eee 102
Mibialebixtensor Group: essence see ene ee ee te eee 103
Reroneale Groupie nese lets ees Veen ey wa srnchee. see 104
(NG ctormG rou pee ey ea ech sie ee toe eee: 105
ischiotrochantericaiGroup 4.6...) gacsdss ea to ee ee 108
Hamstring Groups ace tc eae gee a ee Se aac ees 108
Flexor Groups Ob Weg! tne cots. nea versas geal ale aoe ea 110
lexan GroupsorsPes seit uo we uth dor eee e mee seiama- 112
GWONCEUSIONS ea nica ete ah dente Nees a ean ance entre camer 114
Ga pia OnSetOrue liGhbys. - ale acae Bt ser a cag en ets aA ee 114
iheeMechantes of Bat Michty mines ok «nie sou) ete 119
Comparisons’of. the Bats Studied <4... 0.0.4.7 ks ee 125
Evolutionary C OnsideraliOMs! (see sets tcete ee eee eat: 127
SS EIGN EAT Wa Bet Pere Rae sacle eh cute vied ero hea, te tratlch a caine atee ets AS) a 131
IER ATURE 2 Co CEED ate ie ee ee teen la Ree or Ua, 135
INTRODUCTION
Although bats have fascinated man for centuries, many of the
osteological and muscular specializations that enable these animals
to fly have not been studied. Certain aspects of the life histories
of bats have been carefully investigated, and the biology of some
bats is well understood, but little is known of the foraging habits
or of the details of flight or terrestrial locomotion in many bats.
The primary objectives of the study here reported on were: (1) to
investigate locomotion in bats by means of field studies on the
foraging and roosting habits of three North American bats, the
western mastiff bat (Eumops perotis, Molossidae), the cave myotis
(Myotis velifer, Vespertilionidae), and the leaf-nosed bat (Macro-
tus californicus, Phyllostomidae); (2) to describe and compare
the appendicular myology and osteology of these bats; (3) to
ascertain the functional significance of their basic osteological and
myological adaptations for flight; (4) to find how the differences
between the habits of the three bats under study are reflected in
their appendicular morphology; (5) to compare some of the major
morphological adaptations for flight in bats and birds.
The bats listed above were chosen for study because they differ
from one another widely with respect to habits and morphology
and seem to illustrate much of the diversity occurring between
bats of the suborder Microchiroptera. It was thought that ana-
tomical differences between the three kinds of bats mentioned
might be ascertained more readily and might be more indicative
of contrasting trends within the suborder than would less striking
differences between closely related animals.
Except for scattered observations at earlier dates, field work
on this study began in June 1953, and was also carried out in parts
of the summers of 1954 and 1957. A total of roughly 60 days in
the field was devoted to this project. The laboratory section of
the study was started in December of 1952 and was terminated in
December 1957.
In the nineteenth century a number of papers were published
on the anatomy of bats, but the number of species covered by
these reports was small. Among the authors who investigated
bat anatomy in this period were Cuvier (1800-1805), Kolenati
(1857), and Humphrey (1869). Macalister (1872) studied the
myology of 19 species of bats representing the families Pteropidae,
Rhinolophidae, Megadermidae, Phyllostomidae and Vespertilion-
(5)
6 UNIvERSITY OF Kansas Pusts., Mus. Nat. Hist.
idae, and corrected numerous errors made by earlier workers.
Little attention has been given to the myology of bats since Mac-
alister’s paper was published and it remains a major reference on
the subject. Eisentraut (1936), on the basis of moving picture
filmstrips of flying bats, carefully described the wing movements
made by bats in various types of flight. The life history of Eumops
perotis has been considered by Howell (1920a) and Krutzsch
(1955). The habits of Myotis velifer have been reported on by
Stager (1939) and Twente (1955). Only scattered notes have been
published on the behavior of Macrotus californicus and its life
history is poorly known.
I am pleased to acknowledge the help of Professor E. Raymond Hall,
under whose guidance this study was conducted. For important help in
various ways I extend my sincere thanks to Mr. Sydney Anderson, Dr. George
W. Byers, Dr. A. Byron Leonard, Dr. Charles Pitrat, and Dr. Robert W.
Wilson, all of the University of Kansas, and to Dr. Philip H. Krutzsch of
the University of Pitttsburgh, Mr. J. R. Alcorn of Fallon, Nevada, Mrs. G, M.
Richards of Vidal, California, and to my wife, Hazel A. Vaughan. All photo-
graphs and drawings are by the author.
MATERIALS AND METHODS
Most of my field work was done in southern California, in Los Angeles,
San Bernardino, Riverside and San Diego counties. Some additional ob-
servations were made on Eumops in Merced and Tuolumne counties, Cali-
fornia, The field studies were carried out mainly in localities away from
human habitation, and were supplemented by observations of bats in the
laboratory. Photographs were taken in the field and in the laboratory with
an Eastman Bantam Special and a Heiland Strobonar electronic flash. Bats
were obtained by shooting, by the use of nets, and by stretching wires about
one inch above the surfaces of ponds. Specimens of several kinds of bats
other than those serving as the central subject of this report were taken in
various parts of Kansas, and in Washington county, Utah, and were used as
comparative material.
The animals saved for anatomical investigations were preserved in the
field in a solution of one part formalin to eight parts water. An opening was
made into the visceral cavity of each specimen to facilitate rapid preservation,
and some specimens of Eumops were injected. After several weeks, generally
when the specimens were brought to the laboratory, they were transferred to
a solution of 70 percent alcohol. All of the dissections were made with the
aid of a low power binocular microscope equipped with a 6.6X, a 13X, and
a 80X objective. A number of bats of each of the three species under study
were dissected. No volumetric determinations on the muscles were made.
For any one muscle, individual variation seemed to be slight in size, in place
of origin and in place of insertion.
Skeletons used are in the University of Kansas Museum of Natural History.
The drawings of bones are from the following specimens: Eumops perotis,
73128, 73214; Myotis velifer, 52465; Macrotus californicus, 52458. The fol-
FUNCTIONAL MORPHOLOGY OF THREE BATS 7
lowing preserved specimens were used and are mostly in the above named
museum; specimens marked TV are in the collection of the writer.
Chilonycteris personata Wagner—19 mi. E San Andrés Tuxtla, 1000 ft., Vera-
cruz, 1 (KU 24636).
Pteronotus davyi fuluus (Thomas)—19 mi. E San Andrés Tuxtla, 1000 ft.,
Veracruz, 1 (KU 24646).
Macrotus mexicanus bulleri H. Allen—12 mi. N and 3 mi. W Los Mochis,
Sinaloa, 1 (KU 60609).
Macrotus californicus Baird—35 mi. N Blythe, Riverside Co., California, 18
(KU 76538-76555 ).
Glossophaga soricina leachii (Gray)—2 mi. ESE Tepanatepec, Oaxaca, 1
(KU 70492).
Choeronycteris mexicana Tschudi—3 mi. E Raboso, Puebla, 1 (KU 67382).
Leptonycteris nivalis nivalis (Saussure )—% mi. W Aduana, 1600 ft., Sonora,
1, (KU 25005).
Artibeus hirsutus Anderson—% mi. W Aduana, 1600 ft., Sonora, 1 (KU 25072).
Myotis velifer brevis Vaughan—35 mi. N Blythe, Riverside Co., California, 18
(KU 76556-76573 ).
Eptesicus fuscus fuscus (Palisot de Beauvois )—1 mi. SE Leavenworth, Leaven-
worth Co., Kansas, 1 (KU 76574).
Lasiurus cinereus cinereus (Palisot de Beauvois)—12% mi. N and 5% mi. W
St. Francis, Cheyenne Co., Kansas, 1 (KU 52432).
Tadarida brasiliensis (Saussure)—8 mi. NE Ocotlan, 5100 ft., Jalisco, 5 (KU
32007, 32009, 32011, 32012, 32014).
Chaerophon luzonus Hollister—Luzon, Philippine Islands, 1 (KU 10524).
Tadarida molossa (Pallas)—Zion National Park, Washington Co., Utah, 3
TV 731-733); 2 mi. E La Palma, Michoacan, 10 (KU 38291, 38296, 38298,
38301-38305, 38307).
Eumops perotis californicus (Merriam)—Los Angeles Co.: 3 mi. S and 1
mi. W Newhall, 1 (KU 76575); % mi. NW Chatsworth, 1 (KU 76576); 1
mi. W Chatsworth, 1 (KU 76577). Riverside Co.: 4 mi. SW Lakeview, 3
(KU 76578-76580). San Diego Co.: 1% mi. N Barrett, 5 (KU 76581-
76582; TV 709, 710, 712).
Molossus nigricans Miller—1 mi. N Sebana Grande, Managua, Nicaragua, 1
(KU 71033).
Molossus bondae J. A. Allen—Turrialba, Costa Rica, 1 (KU 57151).
Molossus obscurus E. Geoffroy Saint-Hilaire—I]ha Madre de Deus, Bahia,
Brazil, 3 (KU 41153-41155).
Molossus coibensis J. A. Allen—3 mi. SW Managua, Nicaragua, 1 (KU 71007).
REMARKS ON THE BAtTs STUDIED
The western mastiff bat (Eumops perotis) is a member of the
family Molossidae. This family occurs mainly in the tropical, sub-
tropical and desert regions of both hemispheres. The range of
the species in the southwestern United States extends as far north
as central California; E. perotis has also been recorded from Texas,
the state of Sonora, Mexico, Cuba, and parts of northern South
America. In the southwestern United States this bat inhabits arid
desert regions, grassland areas, sections dominated by chaparral,
and occurs locally in the yellow pine belt of the Sierra Nevada
Mountains of California. (The presence of E. perotis in the latter
8 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
area is hitherto unreported. I have heard the distinctive cries of
mastiff bats on many occasions in Yosemite and Hetch Hetchy
Valleys, Yosemite National Park, and there are several specimens
in the Yosemite National Park Museum.) This is a large bat of
roughly 155 mm. to 185 mm. total length. The broad ears face
ventrolaterad and are connected across the top of the snout. The
flight membranes are tough and leathery; the wings are long and
narrow, and approximately the distal half of the tail extends be-
yond the uropatagium. The color of the pelage is dark gray or
brownish gray.
The cave myotis (Myotis velifer) belongs to the family Vesper-
tilionidae; this family is nearly world-wide in distribution except
in arctic regions. This bat occurs from south-central Kansas through
the southwestern United States, and south to Guatemala. It is
known also from the southern tip of Baja California. The cave
myotis inhabits tropical, subtropical and temperate regions. It is
a small “mouse eared” bat, having a total length of from approxi-
mately 92 mm. to 105 mm. The pelage is usually pale brown;
the wings are broad and the uropatagium extends to the end of
the tail.
The leaf-nosed bat (Macrotus californicus) belongs to the New-
World family Phyllostomidae. The range of this family includes the
warmer parts of the southwestern United States, Mexico, Central
and South America, and the Bahama Islands. This species in-
habits the arid deserts of the southwestern United States as far
north as southern Nevada, south to Baja California and Sonora,
Mexico. The leaf-nosed bat is of medium size, with a total length
of roughly 93 mm. to 103 mm., and is distinctive in appearance.
The ears are large and are connected across the forehead. There
is a small nose-leaf, and the body is pale grayish brown dorsally
with whitish underparts. The flight membranes are thin and deli-
cate; the wings are broad and the tail is slightly shorter than the
long hind limbs and extends several millimeters beyond the uro-
patagium.
LOCOMOTOR BEHAVIOR
Some understanding of both terrestrial and aerial locomotion is
necessary in order to interpret correctly the morphology of the
bats under study. Because terrestrial locomotion is important only
in connection with roosting in most insectivorous bats, and aerial
locomotion is used primarily in foraging, a consideration of the
foraging and roosting habits of each of the bats is important in a
study of their functional morphology.
FUNCTIONAL MORPHOLOGY OF THREE BATS 9
Eumops perotis
Roosting Habits and Terrestrial Locomotion
Most of my observations on free living individuals of this species
were made in the chaparral belt of coastal southern California. I
examined 22 roosting sites that were being used or had recently
been used by Eumops. Of these sites only eight were used by
colonies of bats; most of the other roosting sites were used by
single bats, whereas a few roosts harbored groups of two or three
bats.
Away from human habitations, this bat generally seeks diurnal
refuge in crevices in rocks that form vertical or nearly vertical
cliffs, or that are situated on steep slopes. All of the occupied
crevices that I examined were more than a foot deep and had
entrances that were at least two inches wide and six inches long
and were at the bottoms or sides of the crevices. A larger crevice
is preferred, however, and some of the slitlike openings were five
to 10 feet long. Several crevices that were occupied by colonies of
Eumops were at least 10 feet deep; the total depths of these larger
crevices were not ascertained. Large, exfoliating slabs of rock
seemed most often to form crevices suitable for these bats. Because
granitic rocks and consolidated sandstones are likely to weather by
exfoliation and form deep, vertical crevices suitable for retreats
for Eumops, this bat is most common in broken terrain where ex-
tensive exposures of these rocks occur.
The roosting sites examined all had several characteristics in
common. All of the sites had moderately large openings that could
be entered from below. Measured through the anterior part of
the thorax, the thickest part of its body, Eumops is roughly one inch
thick. This bat occupies crevices the openings of which are at
least twice as wide as its body, probably because entrances of this
size may be entered rapidly and easily. The entrances are usually
horizontally oriented and face downward. According, these bats
can leave their roosts by simply dropping from the crevice, and they
can alight by swooping up into it from below. The entrances to
several of the roosts were more than twelve feet long. All of the
roosts were crevices that became narrow enough at some point to
enable the bats to wedge themselves into the space and have their
dorsal and ventral surfaces against the rock surfaces. On several
occasions bats were frightened and forced themselves into narrow
spaces so tightly that when shot and killed the bats remained in
place.
10 University OF Kansas Pusts., Mus. Nat. Hist.
Characteristically Eumops roosts in crevices that are high above
the ground and have unobstructed approaches. Howell, who stud-
ied colonies of Eumops in buildings in southern California, stated
(1920a:112) that these bats always choose a roost below which
there is a drop of at least 20 feet, so that the bats can, by dropping
from the roost, gain flying speed. I agree with Howell that Eumops
chooses high roosting places. All the roosts found by me were
in rocks in broken, hilly or mountainous country, where steeply slop-
ing terrain provided additional space in which the bat could gain
speed. These roosts were situated from six to 40 feet above the
slope, and a nearly straight drop of at least six feet was always
available beneath the crevice. The occupied crevices that were
fairly close to the ground were always in rocks exposed on extremely
steep slopes.
The character of the vegetation limits the choice of roosting sites.
Most of the roosts were in semi-arid country that supported low
chaparral, the most conspicuous constituents of which were Cali-
fornia buckwheat (Eriogonum fasciculatum), greasewood (Ade-
nostoma fasciculatum), black sage (Salvia mellifera), white sage
(Salvia apiana), and coastal sagebrush (Artemisia californica). In
these areas the plants were low and did not seriously obstruct the
approaches to cliffs and outcrops of rock. In the hills one mile
west of Chatsworth, Los Angeles County, however, the chaparral
was tall and dense locally, and in this area Eumops was not so com-
mon as IJ had expected, probably because tall vegetation encroached
on many of the likely-looking crevices and did not allow a clear
approach from below.
The crevice shown in figure 2, plate 1, harbored roughly six
Eumops at the times of most of my visits, and is typical of many
roosts found in localities having extensive exposures of granitic
rocks. The crevice was in a large boulder of granodiorite situated
roughly 100 yards from the base of a chaparral-covered slope three
miles northeast of Perris, Riverside County.
On several occasions a roosting place was kept under observa-
tion during the early morning hours when Eumops was returning
from foraging. Without exception the bats entered the roost by
swooping up into the entrance from below, grasping the surface
of the rock with both the thumbs and hind feet, and crawling up
into the crevice. This maneuver is generally executed so rapidly
that the bat simply seems to disappear after reaching the mouth of
the crevice; this makes the action difficult to analyze. This bat
FUNCTIONAL MORPHOLOGY OF THREE BATS ll
seems never to cling just inside the entrance of the crevice after
alighting, but crawls as rapidly as possible into the inner parts of
the retreat. This behavior may be important in keeping the en-
trance clear when bats are entering the roost in rapid succession.
Also, it probably makes the bats less vulnerable to predation when
they are entering their roosts.
In its roost Eumops crawls forward at varying rates of speed. In
merely shifting its position slightly, the animal usually crawls
slowly and appears awkward and faltering in its movements, but
at other times crawls fairly rapidly, even when moving but short
distances. When entering the roost after a night’s foraging, or when
frightened and forced to move long distances, this bat loses its ap-
pearance of awkwardness and can move with remarkable speed.
The bat can crawl almost as rapidly within the confines of a nar-
row crevice as across a level cement floor, and in a crevice is able
to maintain forward speed until its dorsum and venter are in contact
with the rock surfaces. At this point the bat turns around, and
in an upside-down position, wedges itself into the crevice by crawl-
ing backward. The hind feet reach behind the animal, grope about,
and when a solid foothold is obtained puil the bat backward; the
bat pushes with its wings, keeping the thumb and pad at the ventral
base of the first metacarpophalangeal joint against the substrate.
I have never observed this bat resting in any other position than
upside-down, with its body wedged either tightly into a crevice or
with the dorsum and venter at least touching the rock. When
roosting in buildings Ewmops seems to behave differently, for
Howell (1920a:114) found individuals hanging against the ridge
pole of an attic near Covina, Los Angeles County, California.
Eumops moves about in its roost at any hour of the day, but
according to my observations is most active in early morning and
late afternoon, at which times the bats are fairly vocal. On several
occasions a colony was observed in the morning and again in late
afternoon. In the morning the bats were invariably in the deeper
parts of the crevice, but in the afternoon usually some were at
places nearer the entrance. Individual bats roosting alone seem to
move relatively little in the daytime.
In some roosts, Eumops must craw] fairly long distances from the
part of the crevice in which it roosts to the launching place. In
large crevices these bats usually crawl far into the deeper recesses
to rest, and in several large crevices the bats habitually rested
roughly 15 feet from the mouth of the crevice. As the time for the
12 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. HIstT.
evening exodus approached, one or two bats at a time moved from
the resting place to near the launching place. Usually the bats did
not leave the roost in rapid succession, but launched themselves
singly, at irregular intervals. On August 20, 1953, in San Diego
County, a colony of Eumops was observed from 5:30 P.M. until
10:30 P.M. (these and all other times mentioned are Pacific Standard
times). The first Eumops launched itself at 7:55, and between this
time and 10:30 only six more bats left the crevice. At 10:30 one bat
was near the mouth of the crevice and four or five were still far
back in the crevice. Throughout the period of observation there
was much movement within the crevice. On June 21, 1954, at
another colony in San Diego County, it took 26 minutes (7:27 P.M.
to 7:53 P.M.) for the 19 adult members of the colony to take flight.
The mechanical arrangement of the hind limbs of Eumops does
not favor quadrupedal locomotion; nevertheless, because of the
strength of the musculature of the pectoral girdle, this bat can crawl
rapidly for fairly long distances. While the animal is crawling,
the body remains parallel to the ground and the ventral surface
of the bat is elevated slightly above the substrate. The tail curves
upward at an angle to the substrate of roughly 45 degrees and
probably serves as a sensory organ when the bat is moving either
frontward or backward in a crevice. The head is held low, in line
with the body. The limbs are used alternately, as is the case in
lizards and most cursorial animals. While the forelimb on one side
is at the forwardmost part of its stride the hind limb on the same
side is extended to the rear at the end of its propulsion stroke. The
limbs on the opposite side of the body are in the reciprocal phase
of the cycle.
During quadrupedal locomotion this bat keeps the digits fully
flexed and the wing-tip is folded against the medial surfaces of the
distal parts of the first and second metacarpals. The elbow and
humerus are held fairly close to the body. The calcar is laid back
near the posteromedial surface of the shank, and the uropatagium
is pulled toward the base of the tail. The proximal part of the
posterior edge of the plagiopatagium is pulled upward, out of the
way of the hind limb, by the partly flexed forelimb. The long axis
of the humerus is roughly parallel to the substrate throughout most
of the stride, while the forearm angles downward and the thumb
and thumb-pad contact the substrate during the rearward (propul-
sion) part of the stride. In the course of a stride the forelimb
makes a movement that resembles a paddling stroke. The fore-
FUNCTIONAL MORPHOLOGY OF THREE BATs 13
arm is partly flexed and is directed forward and laterad at the be-
ginning of the propulsion stroke of the stride; the humerus is par-
tially extended at this point in the stride. The propulsion part of
the stride is produced by the extension of the forearm and flexion
of the humerus, these actions tending to pull the forearm caudad
while keeping the carpus roughly the same distance from the body.
In the propulsion part of the stride the forearm moves through an
arc of some 40 degrees. At the start of the forward part of the
stride the forelimb is lifted from the substrate by abduction of the
entire limb, and is brought forward by flexion of the forearm and
extension of the humerus. This action is extremely rapid; although
the photograph of Eumops in the midst of the forward part of the
stride shown in figure 3, plate 4 was taken at one two-thousandth
of a second, the distal half of the forearm is blurred. At the end
of the forward part of the stride the forelimb is adducted and the
propulsion stroke begins. Because the humerus is directed laterad
while the animal crawls, the forelimbs are splayed out to the side.
Accordingly, although the movement of the humeroradial joint is
anteroposterior, the plane of action of the limb is directed laterad,
and is not vertical to the substrate as in cursorial mammals. The
long forelimb, therefore, can go through the motions of the stride
without markedly increasing the height of the space necessary to
allow passage of the animal’s body. Because the limb is directed
more or less laterad the adductors must act to enable it to bear
part of the weight of the body and the abductors must help in the
forward part of the stride. Thus, not only are the extensors and
flexors of the forelimb brought into play in quadrupedal locomotion,
but the adductors and abductors are of considerable importance.
The hind limb of Eumops is used in reptilian fashion in quadru-
pedal locomotion. The long axis of the femur is held more or less
parallel to the substrate or angles slightly dorsad from the pelvis,
and the shank is vertical to the substrate or, in the propulsion part
of the stride, is directed slightly caudad. At the start of the pro-
pulsion part of the stride the femur is directed craniolaterad, the
shank is extended beyond right angles to the femur, and the foot
is roughly in alignment with the long axis of the body. In produc-
ing the propulsion part of the stride the femur is pulled caudad;
the shank is flexed through the middle part of the stride, and then
extended at the end of the propulsion part of the stride, when the
femur is directed caudolaterad. When the feet are in contact
with the substrate the toes are spread widely apart. The direction
14 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
of movement at the hip joint in quadrupedal locomotion, then, is
anterior-posterior; the plane of movement extends laterad. The
direction of movement at the knee joint is dorsal-ventral; the major
plane of movement of the shank in the stride is anterior-posterior.
The propulsion is provided mainly by anterior-posterior movements
of the femur and by the extension and flexion of the shank. The
adductors and abductors of the hind limb are also important, for
the former supports the weight of the posterior part of the body
and the latter helps in the forward part of the stride.
The limb action used by Eumops in crawling seems awkward,
but if it is remembered that this bat is primarily a crevice-dweller,
its type of locomotion is seen to be remarkably effective. The
action of the limbs is directed largely laterad, so that the animal
requires little more “headroom” when crawling than when at rest,
and can crawl at full speed in a fairly narrow crevice without hav-
ing the movement of the limbs restricted. Howell (1920a:116)
comments that mastiff bats are agile on the ground, and describes
the action of the forelimbs in crawling as being “over-hand.” A1-
though the bat does sometimes give this impression, high-speed
photographs show that movement of the forelimbs is directed
mainly laterad, even when the animal is not in a confining space
(Pl. 4, Figs. 2, 3).
Foraging Habits and Flight
The spectacular manner in which Eumops launches into flight
has been described briefly by Howell (1920a:117) and Krutzsch
(1955:410). All of my observations of these bats taking flight were
made on individuals emerging from crevices in rock. When a
group of mastiff bats are roosting together they become increasingly
active as darkness approaches. When poised at the mouth of the
crevice, shortly before launching into flight, they chatter loudly
and emit loud “smacking” noises; as a bat launches itself it gives
a series of piercing, high-pitched cries. The bat drops from the
mouth of its crevice and dives rapidly downward until sufficient
air speed is gained to allow the bat to begin level flight. The wings
usually give several powerful strokes as soon as the animal drops
clear of the mouth of the crevice, and the bat dives at an angle of
roughly 45 degrees for some 10 to 20 feet before it pulls upward
in a wide arc and assumes level flight. At a colony three miles east
of El Cajon, San Diego County, it was possible to estimate fairly
accurately the distance the bats dropped before leveling off. The
crevice from which the bats emerged was in a large granite boulder
FUNCTIONAL MORPHOLOGY OF THREE BATS 15
on the west side of a ridge and was approximately 22 feet above a
steep chaparral-covered slope that was dotted with large boulders.
The bats habitually dove downward and away from the cliff to
about opposite the level of the base of the colony-rock before be-
ginning level flight; thus most of these bats dropped between 20
and 25 feet. At this roost occasional individuals were observed to
leave the roost in a different fashion. Instead of leveling off after
dropping some 20 feet, these bats cleared the boulder directly be-
neath the colony-rock by a few powerful wing-beats and then re-
sumed their steep dives; with erratic strokes of the wings and rapid
twists and turns they slanted down the slope at high speed, clearing
the brush and boulders by some 10 feet. After diving some 100
yards the bats leveled off and circled near the colony-site for a
short time before flying off. A female Eumops from this colony
was launched by hand near the base of the colony-rock in the day-
time; she dove rapidly far down the slope and was lost to view.
The character of the terrain adjacent to the roosting-site seems to
affect the way in which Eumops takes flight; bats usually emerged
from roosts beneath which there was relatively little space by
diving only five or 10 feet and then pulling sharply upward.
Howell (1920a:112) thought that Eumops could not take flight
unless able to launch itself from a considerable height. Krutzsch
(1955:408) agrees that this bat is reluctant to take flight when low
to the ground, but he observed an individual to fly across a room
30 feet in length from the top of a laboratory table that was only
two and a half feet above the floor, and had another bat fy when
dropped from six feet in the air. Individuals that I tested were
never able to take off from the ground or from other flat surfaces,
and were also unable to maintain flight after launching themselves
from the tops of tables. On several occasions, however, I was
able to launch mastiff bats into flight by throwing them some 15
feet into the air, whereas bats thrown half this distance did not fly.
On one occasion a male Eumops, which I had recently captured,
was able to take flight from a granite bench with roughly a 20 de-
gree slope. Judging from my experience with Eumops, and from
the published observations of others, this bat is totally unable to
take flight from a level surface. Probably five or six feet from the
ground represents the minimum height from which the bat can
launch into sustained flight.
Eumops emits a single high-pitched, piercing cry of short dura-
tion, which can perhaps best be described as a staccato “chip,”
16 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
every two or three seconds as it flies high above the ground, away
from obstacles. The intervals between the cries are not always
constant, and as the bat approaches the ground the intervals de-
crease, until as the bat nears some obstacle or makes its approach to
the roosting-crevice the cries no longer are separate but blend to-
gether in a high buzz.
Characteristically this bat forages at a considerable elevation.
Most of my estimates of the height at which Eumops flies are the re-
sult of inference, but on several occasions, just at dusk, this bat
was watched with binoculars as it gained altitude and flew out of
sight above the colony. At a deserted granite quarry six miles west
of Riverside, Riverside County, on the night of July 9, 1954, from a
vantage point on the west slope of a ridge above the quarry, I
watched two Eumops leave their roosts while it was still fairly light.
The first bat emerged at 7:42 P. M., flew over the floor of the canyon
below the roost, and climbed in large circles out of sight to the
northwest. This bat was watched partly with the aid of binoculars
and was an estimated 1000 feet overhead when lost to view. For
some time after it could no longer be seen this bat’s strident cries
could still be heard. Another Eumops took flight at about 7:50
P. M. and was watched briefly as it circled upward. On this eve-
ning it seemed that the cries these bats made could be heard clearly
when the animals were some 1000 feet overhead. At another colony
a bat under observation disappeared from view when an estimated
800 feet away, but its cries could still be heard a short time there-
after.
Indirect evidence also indicated that Eumops forages, or at least
habitually flies, high above the ground. On the basis of the above
observations, it seems that the cry emitted by these bats as they
fly can be heard clearly at a distance of approximately 1000 feet.
Therefore, when these bats are heard at night directly overhead a
rough estimate can be made of their distance above the observer
by taking into account the faintness or loudness of the cries. These
estimates are admittedly not accurate, and must be influenced
strongly by the differences in the acoustical characteristics of the
air from evening to evening (Knudsen, 1931, 1935), but this is
the only method I found for estimating the height at which Eumops
flies in the hours of darkness.
While watching roosting sites of Eumops at night, I often heard
these bats fly nearby, and many opportunities to gain a general idea
of the height at which they fly were thereby afforded. At the
FUNCTIONAL MorRPHOLOGY OF THREE BATS 17
colony near El] Cajon, San Diego County, I repeatedly heard indi-
vidual bats or small groups pass over the colony-site at such an
elevation that their cries were barely heard. Eumops was com-
monly heard also as it flew immediately above the colony-site, but
rarely was it heard below the site. At this same colony, on an eve-
ning when high fog enveloped the area by about 9:00 P. M., few
Eumops were heard and those that did fly over were near the
limit of audibility. These bats may have been as much as 2000 feet
overhead and more than 3000 feet above most of the surrounding
countryside. In August of 1954 several roosts in Los Angeles
County were observed periodically, and bats in this area were also
heard at night, many times high above their colony-sites. From a
cliff that faced north from the crest of the eastern end of the Santa
Monica Mountains, Los Angeles County, on a summer night when
fog covered the lowlands by early evening and pushed up the slopes
of the mountains to the level of the crest by shortly after midnight,
I heard groups of Eumops far overhead, their cries sounding distant
and faint; occasionally small groups flew close to the cliff. Al-
though it often flies extremely high, this bat may regularly be
heard as it flies only 100 to 200 feet from the ground. Over the
coastal parts of San Diego County, and over the valleys of San
Bernardino and Riverside Counties Eumops seemed to fly com-
monly within several hundred feet of the ground. At Hetch Hetchy
Dam, Tuolumne County, in the summer of 1952, these bats were
heard many evenings as they flew within 100 to 200 feet of the
surface of the water. There, observations were made from a granite
bench that rose high above the water, and estimates of the elevations
at which the bats flew were fairly accurate because the height of
the observer above the water was known.
Both the character of the terrain and the weather conditions
probably influence the elevation at which these bats forage. Over
broken country they seem to fly higher than over level sections.
On foggy nights, in several localities in southern California, I was
impressed by the apparent absence of mastiff bats during part of
the night and their return to the roosts in the morning. Possibly
this bat flies high, or flies many miles away from the colony-site,
to avoid weather unfavorable for successful foraging. The noc-
turnal presence of Eumops on the western borders of the Mojave
Desert of California seems to be sporadic, and the bat’s occurrence
in this area conceivably is governed by weather in adjacent coastal
areas. On many nights in August and September of 1954, on the
18 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Mojave Desert six miles west and one mile north of Lucerne Valley,
San Bernardino County, small groups of mastiff bats repeatedly
were heard overhead starting at roughly 10:00 P. M., whereas on
other nights the bats seemingly were not present in the area. It
was not determined whether or not their presence was correlated
with the occurrence of fog in the nearby. coastal sections. Bad
weather as such, however, does not seem to keep Eumops from
flying. Individuals have been heard on rainy nights in the vicinity
of Claremont, Los Angeles County. In the early morning of July
13, 1954, at the colony site near Perris, Riverside County, a violent
thunderstorm lasted roughly two hours; many times in the storm
mastiff bats were heard overhead.
On the basis of much indirect evidence, the foraging range of
Eumops seems to be extensive. On numerous occasions, late at
night, these bats have been heard flying over areas supporting con-
tinuous tracts of chaparral, or over level desert flats five to ten miles
from the nearest places that a mastiff bat could roost. Groups of
these bats that were heard at various localities on the Mojave
Desert of western San Bernardino County were probably no less
than 15 miles from their roosts. In these instances searches for
colonies of Eumops in the closest desert hills always failed to pro-
duce evidence of the bat’s presence, and their cries never indicated
that they were leaving roosts in these hills in the evening. Almost
without exception the direction of the first cries of mastiff bats that
were heard in the evening in this area suggested that the bats were
coming from the west, from the coastal sections. Although localities
that provide suitable roosting sites for Eumops are by no means of
continuous occurrence in southern California, these bats are, as
indicated by their cries, widespread late at night, occurring from
the seacoast to the western parts of the deserts. From the above
kinds of evidence, I conclude that Eumops generally has a nightly
foraging radius of at least five miles, and that these bats may forage
regularly as far as 15 miles or more from their roosts.
The flight of this bat is strong and fast. Because of its late
emergence this bat can seldom be seen in flight except at close
range by moonlight or by flashlight. Most of my observations of
Eumops in flight were made of bats emerging from their roosts or
of individuals released in daylight. The flight is more direct and
less erratic than that of most other bats; the wing-beats appear to
be slower, and, at least in level flight, shallower. On a still evening
the sharp swishing of the wings can be heard up to a distance of
FUNCTIONAL MORPHOLOGY OF THREE BATS 19
roughly 100 feet. Using the sound of the wing-beat as these bats
flew by colony-sites as a guide, I estimated on several occasions
that there were approximately four beats per second in level flight.
During sudden turns, or when climbing rapidly, the wings appear
to beat much faster than this rate. Often, when diving, the wings
are held partly closed and rigid. When diving rapidly down a
slope after emerging from its roost Eumops looks somewhat like
a large, heavy-bodied swift. In level flight it resembles a medium-
sized charadriid shore bird.
Often, late at night and early in the morning, mastiff bats dive
repeatedly past a colony-site. This behavior was noted at three
different localities and the maneuver is illustrative of some of the
aerial capabilities of this bat. These dives are always heralded by
a sudden increase in the rate of the cries given by a passing
Eumops. Then, with the cries merged into a continuous buzz, the
bat dives at high speed toward a point below the roosting-crevice,
pulls sharply upward just before reaching the cliff, and performs a
half loop which takes the animal out away from the cliff and well
above the level of the roosting-crevice. The dive is usually made
at an angle of at least 25 degrees and at the start of the dive, in a
manner reminiscent of the beginning of a falcon’s stoop, the wings
give several powerful strokes to send the bat slanting rapidly
downward on partly closed wings. At the bottom of the dive the
wings are spread suddenly, and as the bat pulls up and away from
the cliff, the wing membranes produce a penetrating swishing
sound like canvas being lashed through the air. The wing strokes
resume at the top of the half loop and the sounds they make then
recede into the distance. A bat may dive repeatedly at the cliff
and then leave the vicinity of the roost; or after several dives a bat
occasionally enters the crevice. On one evening at the colony three
miles east of El Cajon, San Diego County, a bat made more than
20 dives at the colony rock before flying off. This “buzzing” of the
colony-site is not always done in the same way. Sometimes the
bats approach the cliff from the side and instead of swooping up-
ward at the last minute simply turn sharply away from the cliff and
slant upward at a shallow angle. During their dives the bats are
traveling at a speed that appears to me to be comparable to that of
a stooping sparrow hawk (Falco sparverius), yet the bats are able
to turn abruptly upward at the last instant and avoid the cliff. This
suggests the flight of mastiff bats is fairly maneuverable despite its
speed.
20 UnrIversiry OF Kansas Pusts., Mus. Nat. Hist.
When alighting at the mouth of the crevice Eumops makes a
dive that resembles that described above. In fact, when the diving
behavior was first observed, I assumed that the bats had to dive at
the cliff several times in order to judge their landings properly.
Early in the morning, when bats are returning to the roost, how-
ever, they generally enter the crevice on the first dive, while bats
usually leave the vicinity of the roost after making many dives.
The dive executed immediately before alighting seems to be made
at a fairly shallow angle. The bat dives for a point some 10 or 15
feet below the roost-entrance, then swoops upward just short of
the cliff and seems literally to fly into the crevice. Close observa-
tion indicates that the bats are nearly at the stalling point when
they reach the mouth of the crevice, but have enough momentum to
help them scramble rapidly into the deep part of the crevice. I
have never seen Eumops alight in any other way. Members of a
large colony of Brazilian free-tailed bats (Tadarida brasiliensis) at
the eastern end of the Santa Monica Mountains, Los Angeles
County, entered a hole in a cliff after a maneuver that resembled
that employed by Eumops. The choice of a roosting site by
Eumops may be determined as much by this bat’s need for space
to allow it to swoop upward and alight as it is by the animal’s
need for adequate launching space.
Mastiff bats have a long foraging period. Howell (1920a:117)
stated that these bats have a pre-midnight foraging period that
lasts roughly 45 minutes, but Krutzsch (1955:411) concluded from
his experience with this bat that Ewmops does not return to its
roost until some time after midnight. I have maintained a number
of all-night vigils at or near four roosting-sites of Eumops, and my
observations indicate that this bat usually flies continuously from
shortly after dark until early in the morning. Krutzsch (loc. cit.)
suggests that Eumops practices night-resting, as do many other
bats, but to me, this seems unlikely for two reasons. One is that
mastiff bats require a special type of roost that is not of common
occurrence, and although many roosts known to be occupied during
the daytime were visited at various times of the night, Eumops was
seldom observed to return to a roost to rest. The longest time a
bat remained at the roost after returning at night was less than
half an hour. Secondly, it is usually possible to hear the cries of
flying Eumops in every hour of the night in suitable areas; rarely
does there seem to be a time of peak activity and then complete
absence (except occasionally on foggy nights).
FUNCTIONAL MorPHOLOGY OF THREE BATS oT.
The time of emergence varies, but is characteristically at dark,
roughly one hour after sunset. Generally these bats begin leaving
their roosts 40 to 50 minutes after sundown, but many leave later,
from roughly one hour to one and one-half hours after sunset. By
one and one-half hours after sunset most mastiff bats have left their
daytime retreats. In the summer of 1954 the time of the first
emergence of Eumops in the evening varied from 40 minutes to
one hour and five minutes after sundown. On one evening a colony
was watched for more than two and one-half hours after sunset and
although seven bats left the colony several adult bats still were in
the crevice.
Mastiff bats return to their roosts early in the morning. The oc-
casional individuals that return to the crevice in the night and stay
for a short time are usually females with young in the roost. In
midsummer small groups of Eumops generally begin returning to
the immediate vicinity of the colony-site about 2:00 A. M., two and
one-half hours before sunrise. The groups circle around the area
and bats periodically dive near the crevice; often after staying
near the roost for a few minutes the group will leave the immediate
area. The first bats generally enter the crevice shortly after 2:00
A.M. Aerial bat activity near the colony-site usually reaches its
peak sometime between 2:30 and 3:00 A. M., and most of the bats
enter the roost in this period. In the summer of 1954 the earliest
time at which a colony began entering its roost was approximately
2:00 A. M., two and one-half hours before sunrise, and the latest
arrival recorded was at 4:06 A. M., 45 minutes before sunrise. The
following are excerpts from notes taken on June 20, 1954, at the
colony three miles east of E] Cajon, San Diego County; they give
an idea of the pattern of nocturnal activity at a Eumops colony.
6:00 P.M. Bats chattering and chirping, noise continuing nearly unin-
terruptedly.
7:25 P.M. Loud “smacking” noises at mouth of crevice and much move-
ment in crevice.
7:30 P.M. Two bats left roost; they dove down steep slope below
colony-rock, leveled off and flew nearby for a short time.
7:45 P.M. Nine bats have left crevice; some dropped from crevice and
flew out from rock in shallow arc, others dove down slope.
7:50 P.M. One bat left crevice.
7:55 P.M. Roughly six or eight bats left since 7:50; all bats (except
several young far back in crevice) have emerged.
8:12 P.M. Two or three bats swooping past crevice; bats left area after
five minutes.
8:30 P. M. Several bats back flying over colony-rock.
8:45 P.M. One bat swooped by crevice several times, then entered
crevice.
22, UNIvERSITY OF KAnsAS Pusts., Mus. Nat. Hist.
M. Bat that just entered crevice (8:45) left crevice.
M. Two bats back diving past crevice, then left area.
M. Two bats entered crevice after several preliminary swoops.
. One bat swooped by crevice more than 20 times, then left.
. Two bats (that entered crevice at 10:00) left.
Small group of bats circling rock; one swooped by crevice
several times. Group left after five minutes.
Group of bats flying overhead.
Single bat flying near rock; made 16 dives but did not alight;
left in few minutes.
Between 11:20 P.M. and 1:10 A.M. several groups of bats
came near colony-rock and dove by crevice.
Several bats swooped repeatedly by crevice.
Two bats entered crevice.
Large group of bats circling around rock; individuals oc-
casionally diving by rock.
One bat entered crevice.
No flying bats within hearing.
Group of bats near rock; several swooping by crevice.
One bat entered crevice.
Two bats entered crevice.
One bat entered crevice.
Group flying high nearby (have been overhead since 2:25).
Between 2:45 and 3:30 twelve bats entered crevice.
Sky fairly light; seemingly all bats back in crevice; much
amiable squeaking.
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On the basis of the above and many similar observations I con-
clude that in the summer Eumops has a single, long, foraging
period per night. This period seems to vary from night to night
and between individual bats on a given night. Generally the
foraging period is roughly six and one-half hours in length, and
throughout this time most Eumops fly continuously. For bats, this
is an unusually long foraging period, and indicates that mastiff
bats are not only fast, but enduring fliers.
Eumops is insectivorous. Droppings invariably consist of small
fragments of insects. This bat normally forages between several
hundred and 2000 feet above the ground, and the question arises
as to what insects occur at these levels, and how these insects get
there. Insects are not known to fly regularly at such elevations,
but little is known of this aspect of insect ecology. It is known,
however, that during the daytime rising masses of air take insects
up the slopes of mountain ranges or lift insects high ‘above the
surface of the ground. Many of the areas inhabited by mastiff
bats are fairly hot in the summer and are characterized by surface
topography of high relief; such areas promote turbulence of the
air, and the thermals and convection currents produced in the day-
FUNCTIONAL MORPHOLOGY OF THREE BATS 23
time probably carry many insects high above the ground and well
away from their normal habitats. How long insects carried by air
currents to high elevations stay at these levels has not been de-
termined, but it is possible that considerable numbers remain
throughout the night at the elevations at which Eumops usually
forages. Because it has a wide foraging range and is a speedy
flier, this bat could probably catch an adequate number of insects
even if the insects were far less numerous within the elevational
belt at which this bat forages than they are close to the ground.
Temperature inversions are prevalent in the parts of California
inhabited by Eumops. Thus, when mastiff bats forage above a
dense low-lying fog on a summer night they are often in consid-
erably warmer air than that near the ground. Possibly some in-
sects tend to concentrate in this warmer air. In southern Cali-
fornia on cool winter nights these bats are sometimes heard flying
high overhead. On these occasions, due to the temperature in-
version that often prevails for many nights in this region, the bats
may be in air warmer than that surrounding the observer at
ground level. It cannot be said that the bat finds insects, but the
fact that these bats fly on such nights suggests that they are feeding.
Myotis velifer
Roosting Habits and Terrestrial Locomotion
Field studies on the cave myotis were carried out chiefly in the
vicinity of the Riverside Mountains of Riverside and San Bernar-
dino Counties of southeastern California. The main axis of the
range extends from roughly 30 miles to 37 miles north of Blythe,
Riverside County. The colonies of bats studied were in mine
tunnels in the sides of several large canyons that cut deeply into
the eastern slope of the range. Many of the tunnels were part of
the old “Mountaineer” mining operation, now bearing the name
“New-era Mine.” The Riverside Mountains rise approximately
1500 feet above the surrounding arid desert.
Myotis velifer roosts in a variety of situations; its roosting habits
have been well described by Twente (1955). In my study area the
daytime roosts were all in deserted mine tunnels. Stager (1939:226)
found this bat to be absent from this region in the winter and early
spring. I found these bats in clusters of from several to more than
one hundred individuals in crevices, drill holes and irregular or
hollowed-out areas on the ceilings. Each of several tunnels con-
tained roughly 1000 cave myotis, and each of other tunnels was
24 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
inhabited by several hundred individuals. These bats usually were
most abundant in the deeper parts of the tunnels, beyond roughly
60 feet from the mouth, where they tended to occur in clusters on
the ceilings; in the shallower parts of the tunnels the bats generally
roosted in crevices or holes, and seldom were found less than 25
feet from the mouth of the tunnel. This species has a strong
tendency to cluster, and individuals seem to be ill at ease when
apart from others of the species. On several occasions, when I
was photographing the flight of M. velifer in a deserted cabin, four
or five of these bats were observed to alight near one another and
immediately cluster together in a tight mass.
This clustering behavior of the cave myotis is thought by Twente
(1955) to be associated with temperature regulation both when
the bats are active and when they are hibernating; the clustering
habit may also enable groups of these bats to exploit roosting sites
that cannot be used by single individuals. Considering resting
animals, the posture of the hind limbs of M. velifer is different from
that of the hind limbs of Macrotus. The latter is adapted to hang-
ing free from the ceilings of caves; when it hangs pendant the
hind limbs and feet are relaxed and project straight behind the
animal. The structure of the tarsus and the lengths of the flexor
and extensor tendons of the shank are such that the foot is extended,
even when relaxed, and thus the foot can retain a grip on some
irregularity in the surface of the ceiling when the bat is relaxed
or even torpid. Myotis velifer, in contrast, normally roosts with
its ventral surface resting against something; the hind limbs re-
main partly flexed with the femur projecting craniolaterad and
more or less dorsad and the shanks project posteroventrad and
usually slightly mediad. If M. velifer were to hang supported by
its feet only, the long axis of the body would assume an angle to
the ceiling of more than 90 degrees. When relaxed the feet are
not extended. Were the bat to hang pendant, the hind limbs and
feet would be unable to relax and still maintain a grip on the
ceiling. If the body is held in a position somewhere between hori-
zontal and vertical, with the ventral surface and head downward,
the hind limbs and feet are able to relax and still cling to the sub-
strate. When forced to hang alone from a sloping ceiling M. velifer
generally attempts to help support its body by pressing the wings
against the slanting surface. I have never seen this bat hang singly
from a ceiling, and Stager (1939:227) states that he has never seen
a myotis of any species hang from a ceiling by its claws only. Seem-
FUNCTIONAL MorRPHOLOGY OF THREE BATS 25
ingly, because all the members of a cluster of bats have their ventral
surfaces in contact with other bats or some other supporting surface,
their bodies are held in such positions that the hind limbs and feet
may relax and still maintain a grip. This attempt to keep the ven-
tral surface in contact with a supporting surface may be one reason
why peripheral members of a cluster of bats always have their bellies
facing toward the middle of the cluster.
In their usual daily cycle of activity, when roosting during the
daytime in a cave or mine tunnel, these bats seem to use quad-
rupedal locomotion but little. They move over the rock walls of
a cave only occasionally to seek better roosting sites and craw] short
distances within a roosting crevice. I have never seen M. velifer
crawl long distances under natural conditions.
Despite the seeming unimportance of quadrupedal locomotion
in this species, it is able to crawl and climb fairly well by using its
forelimbs and hind limbs in approximately the same fashion as that
described for Eumops. When crawling, M. velifer raises the an-
terior part of the body farther off the substrate than the posterior
part of the body, and the tail is curled ventrad and forward. When
in contact with the substrate, the forearms incline ventrad more
than do those of Eumops. When the animals are crawling, the body
of M. velifer is at a considerable angle to the substrate, whereas
that of Eumops is almost parallel to the substrate. This same dif-
ference has been noted by Orr (1954:200) between the postures of
crawling Antrozous pallidus and Tadarida brasiliensis, and may be
a consistent difference between the family Vespertilionidae and the
family Molossidae. Judging from the literature and from my own
observations, vespertilionid bats crawl less rapidly than molossid
bats, probably owing to the lack of freedom of the hind limbs from
the flight membranes and the relatively shorter, less rapid strides of
the forelimbs in vespertilionid bats. The uropatagium in most
vespertilionids is large, and incapable of sliding proximad along
the caudal vertebrae, and leaves less of the tail free than in molos-
sids in which the tail is mostly free when the bat crawls. In ves-
pertilionids, accordingly, the hind limbs are not able to move so
freely during terrestrial locomotion as do those of molossids. As-
suming that M. velifer and E. perotis possess the types of quad-
rupedal locomotion characteristic of the family Vespertilionidae and
Molossidae respectively, the posture of the forelimbs of vesper-
tilionids is probably more efficient in terms of energy necessary for
the stride because the limbs are oriented more nearly vertically to
the substrate than the ventrolaterally directed forelimbs of molos-
26 UNIVERSITY OF Kansas Pusxs., Mus. Nat. Hist.
sids. Because of the slightly upright posture of the body and the
partially vertical action of the forelimbs, however, quadrupedal
locomotion in vespertilionids is not so efficient within the confines
of a narrow crevice, where, judging from my observations on E.
perotis, Tadarida molossa and T. brasiliensis, the locomotion of
molossid bats is remarkably effective.
Foraging Habits and Flight
Cave myotis take flight and alight in the manner common to most
vespertilionids. When launching itself M. velifer drops from its
roosting place with its wings partly open, spreads its wings and
starts beating them rapidly when clear of obstructions, and pulls
upward into level flight at from a few inches to a foot or two be-
neath the roosting place. These bats can easily launch into flight
from a level surface by pushing off with their wings and feet. Land-
ings are usually made on steeply sloping ceilings or vertical sur-
faces, and the landing surface is grasped with the thumbs and hind
feet. The landing is made with the head upward. The uropatagium
is lowered and spread just before the landing and the wings are
fully spread; the bat slows down rapidly and when near the stalling
point comes into contact with the landing surface roughly simul-
taneously with both the forelimbs and the hind limbs. When a
solid foothold is gained the bat releases its hold with the thumbs
and one foot and moves into a head-downward position. Often,
instead of finding a foothold and assuming a head-downward posi-
tion immediately, the bat feels about with its feet while bracing
its body with its forelimbs, and slowly shifts into the head-down-
ward position.
The flight of the cave myotis is stronger and more direct than
that of most members of the genus Myotis, but is highly maneuver-
able. On many evenings when western pipistrelles (Pipistrellus
hesperus), California myotis (Myotis californicus) and M. velifer
were foraging at the same time over common ground, the more
rapid, less fluttery flight of M. velifer was accentuated by the erratic
flight of the smaller species. When foraging this species flies fairly
steadily until an insect is perceived, then the flight often becomes
extremely erratic. After the insect is captured the bat continues
on its relatively straight course until another insect is chased. The
foraging flight of the cave myotis thus alternates between direct,
steady flight, and abrupt twists and turns. When sudden changes
of direction are made the uropatagium and wings are normally
spread fully, and the bat uses the full area of its flight membranes
FUNCTIONAL MORPHOLOGY OF THREE BATS 27
as a braking surface. From level flight these bats turn upward,
downward, or to the sides to chase insects, but seem not to chase
their prey upward as often as in the other directions. The pinnae
of M. velifer face forward and slightly downward and to the side
when the animal is in flight; insects are perhaps perceived more
often when they are in front, below, or to the sides of the bat than
when they are flying above the bat. Hall and Benson, while ob-
serving bats in Nevada (Hall, 1946:142), noted a positive correla-
tion between the directions in which the pinnae of the different
species faced and the directions in which insects were pursued. In
the Riverside Mountains area, after leaving their daytime retreats,
cave myotis usually flew directly down the eastern slope of the
range to the floodplain of the Colorado River where they foraged.
The flight at such times was steady and fairly rapid, and when
flying down steep slopes the bats occasionally partially closed their
wings, dived rapidly, and skimmed not over five feet above the
large creosote bushes and jutting outcrops of rock. On the other
hand, when this bat is in a cave or mine, or other restricted area, it
travels more slowly and the wing-beat is shallower and seems more
rapid.
In the Riverside Mountains area these bats forage mostly over the
floodplain of the Colorado River, where they pursue foraging beats
over low vegetation, along the files of dense vegetation that line the
oxbows and main channel of the river, between the scattered thick
patches of vegetation that dot the floodplain, or above bodies of
water. The dense linear stands of screw bean (Prosopsis pubes-
cens), tamarisk (Tamarix), catclaw (Acacia Greggii) and mesquite
(Prosopsis sp.) that border the still water of the oxbow ponds seem
to constitute optimal foraging habitat, and in such localities just
before dark on a summer evening there are usually six or eight
M. velifer in view at all times. These bats do most of their foraging
between roughly six and 15 feet of the ground, and prefer to forage
close to vegetation. When foraging near large patches of vegetation
these bats often fly within a few inches of the foliage; in chasing
insects the bats often fly through small spaces between branches
or through narrow corridors between adjacent plants. Under some
circumstances, these bats flew back and forth over definite foraging
beats estimated to be from 50 to 70 yards in length.
The character of the vegetation affects the mode of foraging of
M. velifer; near mesquite or tamarisk, or other plants that normally
grow to at least nine to 12 feet in height, this bat flies near the
plants, and stays six to 12 feet from the ground, usually keeping to
28 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
the level of the main mass of foliage rather than foraging above it.
Near patches of arrow weed (Pluchea sericea), the bat forages
mostly five to eight feet above the ground, just above the vegetation.
While chasing insects under these conditions these bats sometimes
fly through spaces between the wandlike arrow weeds and descend
to within three or four feet of the ground. Also, when foraging
over low cattails (Typha sp.) or open water this bat is apt to fly
lower than it does around tall vegetation. Although seemingly not
so common in these areas, M. velifer often forages over dry washes
where it tends to fly from ten to 15 feet above the ground, at the
level of the thickest parts of the larger mesquite, catclaw and palo
verde (Cercidium floridum). In open desert, dominated by the
creosote bush (Larrea divaricata), where I have observed this bat
foraging on relatively few occasions, it flew just above the level of
the bushes, four to seven feet above the ground. Within the gen-
eral elevational zone at which M. velifer forages, then, it usually
flies close to foliage, and the shapes of the plants and the level at
which the densest vegetation occurs seems to govern the elevation
at which the bat flies.
In the evening M. velifer usually emerges from its daytime roost
well before dark, and for a short period forages with, but at a lower
elevation than, the western pipistrelle. In June of 1953, the times
of emergence of these bats from their daytime retreats varied from
7:08 P. M. to 7:50 P. M. (17 minutes to one hour and four minutes
after sunset), and the average time of emergence was 37 minutes
after sunset. In late June of 1954, these bats emerged somewhat
later, appearing at from 42 to 55 minutes after sunset. The
temperature was higher in late June of 1954 than in early June of
1953, and may have affected the time of emergence of the bats.
I gained the impression that this species started foraging later on
hot nights than on cooler nights. All of the individuals of a colony
of cave myotis do not leave their daytime roost together in the
evening. As an example, at a mine tunnel in the Riverside Moun-
tains, on June 6, 1953, it took 47 minutes for the emergence of a
colony of roughly 200 of these bats. To an observer stationed on
their foraging grounds, however, this species seems to appear sud-
denly in large numbers. Twente (1955) gave a good description of
the emergence of large colonies of M. velifer from caverns in south-
central Kansas.
The pre-midnight foraging period of this bat lasts until nearly
midnight. Individuals with full stomachs have been caught enter-
- FUNCTIONAL MORPHOLOGY OF THREE BATS 29
ing night-roosting places at from 8:10 P.M. to 11:41 P.M. (one
hour and 22 minutes to four hours and 53 minutes after sunset).
Each bat probably forages for considerably less than an hour in
this period; individuals captured near midnight probably were
moving from one roost to another or were doing a limited amount
of foraging after their main foraging period. Almost without ex-
ception, cave myotis that were obtained after 8:15 P. M. (one hour
and 27 minutes after sunset) had full stomachs. Considerable
evidence suggests that in summer M. velifer fills its stomach within
roughly one-half hour after emerging in the evening. For example,
on August 22, 1957, two individuals were shot as they foraged over
an oxbow of the Colorado River 57 minutes after sunset and roughly
30 minutes after the first M. velifer was seen foraging on that even-
ing; the stomach of one bat was nearly full, and that of the other
was slightly more than half fuJl. As evidenced by sight records of
foraging cave myotis and tabulations of numbers of individuals
entering night-roosting places, the greatest nocturnal activity of
these bats occurs between 35 minutes after sunset and four and
one-half hours after sunset.
In the middle of the night these bats roost in some sheltered
place and rest; there seems to be no clearly defined second foraging
period in early morning. This species has been observed night-
roosting in a variety of buildings, caves and mine tunnels. When
night-roosting these bats are more likely to roost singly or in small
groups than they are in the daytime. Night-roosting has been
observed in every hour between one and one-half hours after
sunset and one and one-half hours before sunrise, and the bats
have been recorded returning to their daytime roosts between
1:30 A. M. and 3:30 A.M. (roughly three hours to one hour before
sunrise). The records of cave bats taken at the entrances of day-
time and nighttime roosts in early morning hours indicate that there
is no long or well defined early morning foraging period, and that
the bats do not attempt to fill their stomachs just before daybreak;
in fact M. velifer often goes to roost for the day with an empty
or only partially full stomach. Many specimens were taken at their
daytime roosts immediately after sunrise, and the stomachs of most
of them were either empty or only partially full. Between June 6
and 15, 1953, 15 cave myotis were taken as they entered roosts
between 12:50 A. M. and 3:28 A.M.; the stomach of none of these
bats was more than half full, and most of the stomachs were one-
quarter full or less. From evidence of this sort, I conclude that
80 UNrversity oF Kansas Pusts., Mus. Nat. Hist.
these bats rest during most of the early morning hours, and that
there is no regular foraging period after midnight, but that many
individuals do a limited amount of foraging between midnight and
sunrise. It further seems likely that counting the time necessary
for the bats to travel from their daytime roosts to their foraging
areas and back M. velifer is on the wing no more than one and
one-half hours per night.
My field studies have given little basis for a general knowledge of
the foraging range of this species. In two localities foraging areas
of these bats were slightly more than a mile from the nearest roosts.
M. velifer is a strong flier, and I expect its nightly foraging range
under some circumstances is more than this distance.
Macrotus californicus
Roosting Habits and Terrestrial Locomotion
Macrotus was studied in the field mainly in the Riverside Moun-
tains of California, in the same localities at which Myotis velifer
was studied.
In the study area Macrotus roosts in the daytime exclusively in
caves, deserted mine tunnels and deep grottos. In the Riverside
Mountains many of the deserted mine tunnels in the steep sides of
the rocky canyons were inhabited by groups of from several to
100 or more of these bats. They usually were within 30 to 80
feet of the entrance of a tunnel, and seemed not to require dark
retreats. On many occasions leaf-nosed bats roosted in short tun-
nels less than 20 feet deep and fairly brightly lit. Small groups of
these bats roosted in some of the deeper natural grottos in the
walls of canyons.
In order to be suitable for occupancy by Macrotus a retreat must
be mostly inclosed and have overhead protection from the weather.
Although this bat is perfectly capable of flying in limited spaces
and has been observed on many occasions to fly through a small
opening to enter a mine shaft, occupied roosting chambers are
usually large enough to provide considerable ceiling surface and
flying space. Most of the tunnels inhabited by Macrotus were from
five to seven feet high and roughly the same width, and were from
15 to more than 100 feet deep. One grotto was inhabited by ap-
proximately 15 Macrotus; the entrance was roughly 12 feet high
and 25 feet wide, and the chamber extended some 30 feet into the
side of the canyon. This bat may prefer a large roosting chamber
because it provides adequate space for flying when the animal is
searching for a suitable place to hang. Another factor governing
FUNCTIONAL MORPHOLOGY OF THREE BATS Sl
the choice of roosts may be temperature, for most of the occupied
roosts were fairly cool in summer. For example, on August 15,
1953, a small group of Macrotus was hanging in a temperature of
84° F. 45 feet inside a tunnel in the south-facing slope of a narrow
canyon in the Riverside Mountains, whereas the temperature out-
side the tunnel was slightly above 110° F. in the shade. Approxi-
mately the same differences in temperatures prevailed at all other
roosts at which temperature records were kept.
This bat always rests by hanging pendant, by means of one or
both feet, from the ceiling of its roosting place. The wings are
loosely folded. Although these bats often roost together in small
groups, their bodies usually do not touch each other, and the bats
become restless when their bodies are in contact with other bats.
They prefer to hang from rough or sloping parts of the ceiling where
the irregularity of the surface of the rock enables them to find a
solid foothold with a minimum of effort. While settling itself im-
mediately after having alighted, when adjusting its foothold, or
when steadying its body in preparation to launching into flight,
the ventral surfaces of this bat’s feet usually face ventrad and
slightly mediad and appear to be actively gripping the rock. At
such times the hind limbs may be partly flexed, thus pulling the
body toward the ceiling, and the head is often raised while the bat
looks about. When Macrotus is relaxed the hind limbs are straight,
the soles of the feet face ventrad, and the head is pointed down-
ward, This bat seems to rest much of the time while hanging onto
the rock with only one foot; the other foot and leg are relaxed and
dangle, partly flexed, down to the side of the animal. The free foot
is often used for scratching and for grooming the fur, and when the
bat is engaged in these activities the body usually swings gently like
an erratically disturbed pendulum.
Although it is apparent that Macrotus actively grips the surface
of a ceiling at certain times, the feet seem able to retain their hold
on a rough surface when they are completely relaxed. The strongly
recurved claws gain purchase on small irregularities in the rock and
help prevent the relaxed foot from loosing its grip; as an added help,
the force of the tonus of the muscles of the shank and the build of
the tarsus tend to extend the foot slightly when the limb is relaxed,
thus pressing the digits against the surface of the rock. Because of
the large size of the hamstring group of muscles and the mechanical
advantage afforded by their attachments, under the control of the
tonus of the hind limb musculature the shank flexes when the hind
limb is relaxed.
32 UNIVERSITY OF Kansas Pusxs., Mus. Nat. Hist.
Macrotus seems unable to perform any type of quadrupedal loco-
motion. I have tried many times to force this bat to crawl, but
always without success. If its wings are free, the bat always
launches into flight immediately by pushing off with the wings.
The structure of the hind limbs is such that an effective stride can
not be accomplished unless the bat is hanging. This bat occasion-
ally “walks” bipedally. The animal releases its hold on the ceiling
with one foot, the body swings forward (dorsad in relation to the
animal’s body ) and the free foot flexes, reaches dorsad, and gains a
new hold on the ceiling. The other foot then lets go and the body
swings forward again at the start of a new stride. In this fashion
the bat progresses across a ceiling by a series of short strides of the
hind limbs, assisted by the swinging of the body. The plane in
which the body swings is roughly parallel with the direction of
movement, and the dorsal surface of the body faces the direction in
which the bat is moving. The head is lifted dorsad while the animal
is moving, and the bat seems to be looking about. Although this is
a peculiar-looking type of locomotion, it is fairly rapid, and by it
Macrotus can travel a foot or two in a few seconds. I have never
seen Macrotus under natural conditions move more than two and
one-half feet at one time by “walking.” This bipedal locomotion
seems to be used when a bat is seeking a more suitable roosting
place, but this type of locomotion seems to be used for this purpose
less often than is flight.
On several occasions groups of 20 to 80 individuals were ob-
served for periods of one to two hours in mid-afternoon. If the
observer remained still the bats seemed not to be disturbed. Even
so, colonies are restless. When alert the bat holds its head up with
the occiput against the anterior part of the interscapular area and
the large eyes look about. In the daytime, there is much shuffling
of wings and adjusting of feet. Often one bat will make a flight of
several feet and alight amid several bats hanging close together,
whereupon the group of bats will become active and in a few
seconds all of the members of the group will be in flight. Usually
they assemble in small groups at various places on the ceiling of a
roosting chamber; the members of a group usually stay far enough
apart so that their bodies are not in contact with each other.
Foraging Habits and Flight
The flight of Macrotus is remarkable chiefly for its extreme
maneuverability. This bat flies fairly rapidly on occasion, but the
usual foraging slight is slow and buoyant, and more nearly silent
than that of mdst bats. While watching bats at the entrances of
FUNCTIONAL MORPHOLOGY OF THREE BATS 83
caves at night I was able to identify several of the most common
species by the sounds of their wing-beats. In level flight the wings
of Macrotus make a soft fluttering sound that is less sharp and
carrying than the sounds made by the wing-beats of most bats.
The method of landing was first mentioned by Howell (1920b:
172) and later by Hatfield (1937:97), but has not been described in
detail. High-speed photographs show that the alighting maneuver
is remarkably intricate. The bat usually makes a level approach
to the intended roosting place. At a point some six to eight inches
below the ceiling the wings make a deep downstroke that is directed
nearly straight forward, and the hind limbs and uropatagium are
lowered. These movements cause the bat to swoop sharply upward
toward the ceiling. As the bat nears the ceiling the wings are pulled
back in an upstroke while the bat rolls over 180 degrees so that its
back is facing downward and the long legs reach for the ceiling.
At the peak of the upstroke of the wings the feet are pulled far
ventrad and come in contact with the ceiling while the body is
almost upside-down with the head downward. After the feet
grasp the ceiling the wings make a last downstroke that steadies the
body and cushions the impact of landing. Stated briefly, then, the
alighting maneuver consists of an upward swoop and a half-roll,
at the end of which the feet swing rapidly toward the ceiling,
seize it, and the wings give a final beat to steady the bat. Usually
no parts of the animal but the feet touch the ceiling when the bat
alights, but sometimes the wings touch the ceiling when they are
adducted rapidly after the feet have clutched the ceiling. The ears
are erect and the head faces toward the intended landing point
during the entire maneuver. Often these landings must require
remarkably precise judgement of speed and distance, as many
landings are made in the midst of a fairly closely-spaced group of
bats. It is not necessary for these bats to approach the ceiling
rapidly in preparation for alighting; they perform the same landing
maneuver after hovering close to a ceiling for several seconds. This
half-roll method of alighting was the only one Macrotus was ob-
served to use.
Macrotus has two main methods of launching into flight, by
dropping from the ceiling and taking flight after a short downward
swoop, and by taking flight directly from the roosting place. The
first type usually is used when an individual launches itself from a
group of bats that are close together, and enables the bat to avoid
bumping its neighbors. The bat releases its grip on the ceiling by
flexing the hind limbs and feet and extending the digits, and drops
2—4357
84 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
clear of the surrounding bats. After dropping a foot or less the
bat pulls upward in a shallow arc and begins level flight. This
method often is used by a bat that is hanging alone. Instead of
beginning level flight after dropping clear of the bats hanging
nearby, an individual sometimes hovers immediately beneath the
group for a few seconds and then alights at a point only a few
inches away from its former roosting place. The second method of
launching is generally used when a bat makes a short flight during
which it remains only a few inches below the ceiling. While the
feet retain their grip on the ceiling and the head is elevated, the
wings are spread and make several rapid strokes; this swings the
body in the direction of the bat’s dorsal surface and tends to bring
the body nearly parallel to the ceiling. The bat then releases its
hold on the rock and immediately begins nearly level flight. The
bat drops only slightly away from the ceiling, and to an observer
seems suddenly to begin level flight from a hanging position.
Frequently when Macrotus takes flight in this way the wings can
be heard rustling against the ceiling.
This bat often hovers, both when foraging and when flying in its
daytime retreat. Macrotus seems to hover easily, and is able to
hover for several seconds at a time. On many occasions, while I was
watching groups of leaf-nosed bats at their daytime roost, individuals
flew down to within a foot or two of my face and hovered there for
several seconds as they did also at night. Hovering is accomplished
by flying upward with just enough force to counteract the pull of
gravity. The body is nearly vertical to the ground, the legs are
pulled laterad and the uropatagium is spread as fully as possible,
while the wings make rapid but shallow beats. The ears are erect
and the head is pointed toward whatever occupies the bat’s
attention.
In early June, 1953, two miles south of Vidal, Riverside County,
in a deserted cabin on the border of a dry wash supporting scattered
examples of palo verde, catclaw, smoketree, ironwood and mesquite,
Macrotus, together with pallid bats (Antrozous pallidus), big-eared
bats (Corynorhinus townsendii), cave myotis and Yuma myotis
(Myotis yumanensis) roosted at night. In the three-day period of
my stay there, the locality was a favorite foraging area for numbers
of leaf-nosed bats, and they could be observed easily by flashlight
as they flew over the wash or the nearby desert. These bats regu-
larly foraged here as elsewhere within three feet of the ground.
They often dropped down nearer the surface of the ground and
occasionally hovered there for a few seconds. They frequently
FUNCTIONAL MoRPHOLOGY OF THREE BATS 35
foraged close to vegetation and seemed not to slacken speed when
approaching obstacles, but avoided them easily and gracefully.
Even bats released in the daytime flew fairly close to the ground.
One individual liberated in this area in the daytime flew roughly
300 yards before being lost to view and was never seen to fly higher
than approximately ten feet above the ground.
Leaf-nosed bats seem to be totally insectivorous, and their food
clearly reflects the bats’ foraging habits. Some insects regularly
eaten by Macrotus are almost certainly taken from the ground or
from vegetation. The stomachs of these bats taken in the summers
of 1953 and 1954, in the Riverside Mountains, contained fragments
of orthopteran insects, noctuid moths and caterpillars, and beetles
of the families Scarabaeidae and Carabidae, together with uniden-
tified material. The wings of sphinx moths (Sphingidae), butter-
flies and dragonflies were found beneath several night-roosts of
Macrotus. Beneath roosts in Imperial County, California, Huey
(1925) found remains of grasshoppers, sphinx moths, noctuid moths
and beetles. The flying insects mentioned above could possibly have
been caught while they were on the wing, but more likely were
taken while resting. This supposition is supported by Huey’s (op.
cit.) finding willow leaves (Salix sp.) beneath the roost of leaf-nosed
bats and Stager’s (1943) finding a dead leaf-nosed bat on a spiny
desert shrub ( Eucnide urens). H.W. Grinnell (1918:257) mentions
that a leaf-nosed bat was captured in a mouse trap set on the open
desert near Mecca, Imperial County, California. The lists of food
items of Macrotus contain a preponderance of insects that seldom
fly, are flightless, or that fly in the daytime; this constitutes strong
evidence that this bat takes mostly insects that are on the ground
or on vegetation. Assuming that Macrotus does so forage, its mode
of foraging can be seen to be particularly effective in arid regions
where there is little ground cover and where most of the vegetation
has sparse foliage. Nocturnal insects moving along the open sandy
or gravelly floor of the desert would be vulnerable to predation by
a bat flying just above the surface of the ground, and insects cling-
ing to sparse foliage would not be well protected.
Macrotus frequently alights to eat its prey and therefore probably
does not forage continuously for long periods of time. Because
some of the food items taken by this bat are fairly large, a small
number of captures may suffice to fill its stomach. On several nights
I watched a number of Macrotus returning to their daytime roost in
order to eat insects they had just captured. On June 11, 1953, I
observed this behavior at a large grotto used as a daytime roost
36 UNIVERSITY OF Kansas PUBLS., Mus. Nat. Hist.
and nighttime roost by leaf-nosed bats. From 8:08 P. M., by which
time all of the bats in the grotto had begun to forage, until 9:00
P. M. there was an intermittent traffic of bats flying in and out of
the grotto. It was my impression that during this period bats were
foraging near the grotto and returning to the roost to eat the larger
insects, and then leaving to do more foraging. At the cabin two
miles south of Vidal, San Bernardino County, these bats often
alighted beneath the eaves of the cabin where they stayed for only
a short time, probably to eat recently-caught prey.
Indirect evidence suggests that Macrotus has a small foraging
range. They foraged over the floodplain of the Colorado River
roughly one mile from the nearest place where Macrotus could
roost, and at a cabin approximately one and one-half miles from
the nearest known daytime roost. I saw none at the town of Vidal,
about three miles from the nearest known roost. On a number of
evenings, constant activity around a daytime roost suggested that
Macrotus was foraging nearby, perhaps within a few hundred yards
of the roost.
At all localities where records were kept of the times of emergence
in the evening of the various local bats, Macrotus emerged last. The
first individuals of this genus to begin foraging usually left their
roosts no earlier than at dark, roughly one hour after sunset; this
was usually about an hour and a half after the first western pip-
istrelles (Pipistrellus hesperus) appeared. The earliest recorded
emergence for Macrotus was 7:40 P.M., 51 minutes after sunset,
when a group of six left a grotto in the north-facing wall of a canyon.
Because of the depth of the canyon and the steepness of the slopes
the illumination was considerably lower here than at many other
roosts at a comparable time in the evening. The members of small
groups of Macrotus often left their daytime retreats together, or
nearly so, but taking all of the records into account, it seems that
the time of emergence of Macrotus in the evening is spread out over
approximately three hours. On a number of nights I spread a silk
“mist net” across the mouths of various caves and mine tunnels and
kept record of each bat captured, its time of capture, and, when
the animal was killed and saved for a specimen, the amount of food
in its stomach. Because the net was taken down periodically to
allow the passage of large groups of Myotis velifer, the stomach of
each Macrotus was checked before it was counted as having just
emerged. Some of the bats that hit the net from inside the caves
probably were simply making short flights preparatory to leaving
FUNCTIONAL MORPHOLOGY OF THREE BATS 37
the roost, but in general the records of times of emergence obtained
by this method agreed with those gained by direct observation of
colonies. Leaf-nosed bats generally began striking the nets slightly
after one hour after sunset, and the frequency of captures was
greatest from roughly one hour and fifteen minutes after sunset to
two hours after sunset (approximately 8:05 P.M. to 8:50 P.M.).
Many leaf-nosed bats with empty stomachs were taken as late as
two hours and fifteen minutes after sunset, and one bat having only a
trace of food in its stomach was caught three hours and four minutes
after sunset (9:51 P.M.). The pre-midnight foraging period lasts
until about four hours after sunset (11:00 P.M.). That is to say,
most of the leaf-nosed bats forage sometime between one hour after
sundown and four hours after sundown, and then retire to a night-
roosting place. Actually, each bat seems to have a pre-midnight
foraging period of roughly one hour. Only part of this time is spent
foraging, because the bats alight to eat the larger insects.
In the middle of the night Macrotus hangs in some retreat and
rests. It is less selective in choosing nighttime roosts than in select-
ing daytime roosts; adequate overhead protection seems essential,
but a wide range of structures are occupied at night—old adobes,
deserted wooden buildings, cellars, porches, and a wide variety of
caves, mine tunnels and grottos.
In the early morning there is a second foraging period beginning
shortly after midnight, probably around 1:00 A.M., and not so
clearly defined as the pre-midnight period. Although some bats
become active at about 1:00 A. M., some four and one-half hours
before sunrise, many rest until around 3:00 A. M., one and one-half
hours before sunrise. The greatest activity in early morning seems
to occur between two and one-half hours before sunrise and thirty
minutes before sunrise. Bats generally begin returning with full
stomachs to their daytime roosts about two hours before sunrise,
and the last bats usually return approximately twenty minutes
before sunrise. A pronounced increase in Macrotus activity occurs
roughly forty-five minutes before sunrise, and suggests that many
bats wait until fairly late to fill their stomachs preparatory to re-
turning to their daytime roosts. Light-tolerance of this bat possibly
differs in the morning from that in the evening. Usually there is
some activity of leaf-nosed bats through the early morning hours,
but I doubt that the early-morning foraging period of each bat is
longer than forty-five minutes. Many bats probably forage for
considerably less time than this, for in the summer when large prey
88 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
items such as grasshoppers and large crickets are common, a part of
the total time spent by Macrotus in filling its stomach must be
occupied by eating the large food items while the bat hangs in some
sheltered place. It is, of course, difficult to estimate the amount of
time spent by this bat in traveling from. its daytime roost to its
nocturnal foraging area, and this would add to the length of the
activity period. Considering all of these points, one hour and forty-
five minutes is a reasonable estimate of the maximum amount of
time spent on the wing by Macrotus on a summer night.
AERODYNAMIC CONSIDERATIONS
According to Savile (1957:212) “The form of a bird’s wing is so
basically important to the successful exploitation of an ecological
niche that it inevitably yields many instructive examples of adap-
tive evolution.” The same could be said of the chiropteran wing,
and as a basis for the anatomical discussions to follow it is necessary
to consider briefly the aerodynamic significance of certain forms and
functions of the wings of the bats under consideration here.
Poole (1936) showed that most bats have lower wing loadings
than do birds, and he mentioned that this difference may be due to
the way in which many bats forage. (Poole used the number of
square centimeters of wing surface per gram of body weight in his
measurements of wing loadings; following standard engineering
practices, I have measured the loadings in pounds per square foot,
and have assumed that the wings were continuous through the
TABLE 1. AERODYNAMIC CHARACTERISTICS OF THE WINGS OF THE
Bats STuDIED.*
Hae -
: 0 ro-
ee proxi- | patagial
Weicht Wing (oun ap Aspect | mal to | loading
Species ey span P ne ratio distal | (pounds
= (mm.) iY A ae (span?/ | segment per
f Gt) area) of square
° wing foot)
(area)
Eumops perotis...... 55.0 (5)| 516 (4)| .51 (4)} 11.9 (4)} 68.8 (4)] 4.5 (38)
Myotis velifer....... 8.6 (20)} 249 (6)| .19 (6)| 6.8 (6)! 78.8 (6)} 1.4 (3)
Macrotus californicus | 14.1 (12)| 295 (6)| .23 (6)} 6.8 (6)| 80.3 (6)} 2.1 (3)
* The measurements from which the above data were obtained were taken from speci-
mens in alcohol; thus, although the table shows in general the comparative aerodynamic
characteristics of the wings it does not necessarily give the values that would be obtained
from fresh specimens. The figures are all averages, and the numbers of specimens upon
which the averages are based are given in parentheses.
FUNCTIONAL MORPHOLOGY OF THREE BATS 89
bodies.) Most insectivorous birds forage by making repeated, short
flights. Bats, on the other hand, remain on the wing for most if
not all of the time they are foraging, and insects are captured by
virtue of the bat’s ability to maneuver rapidly. A light wing load-
ing enables bats to fly with fair economy of energy for long periods
of time at low speeds, and the large flight surfaces are of great help
in turning abruptly and in maneuvering while capturing insects.
The small sizes of most insectivorous bats favor low wing loadings
because the ratio of mass to surface area is small (the volume and
mass vary as the cube of the linear dimensions, whereas the surface
area varies as the square). If bats had as great a size range as
birds, the wing loadings of the two groups probably would more
nearly resemble each other. In birds the slots formed by the alula
and primary flight feathers allow these animals to have high wing
loadings and fairly low stalling speeds. The unslotted wings of
bats do not yield such refinements in flight characteristics, but within
the range of size and wing loadings found in bats there is little need
for increasing lift. In one respect the flight of bats is clearly less
efficient than that of birds. The upstroke consumes relatively more
power in bats than in birds because the wing-surfaces in bats are
continuous and do not allow the passage of air as do the spaces be-
tween the primaries of a bird. It is probable, however, that differ-
ences between the wing loadings of birds and insectivorous bats
reflect primarily the demands of the foraging habits of bats.
Wings of most bats are airfoils of high camber (anteroposterior
curvature) and lateral camber (decrease of the dihedral from the
base to the tip of the wing). In terms of lift, this type of wing is
efficient at low speeds. The wing membranes are irregular in shape
and the part of the wing composed of the propatagium (the mem-
brane anterior to the humerus and radius) plus the plagiopatagium
(the membrane posterior to the humerus and radius and between
the fifth digit and the body and hind limb) has a greater surface
than the chiropatagium (the membranes between the digits), which
narrows distally and has its leading edge gently swept back. Cam-
ber of the wing is produced by flexion of the digits, particularly the
fifth, the position of the hind limb, which anchors the posterior edge
of the plagiopatagium, and by the angles that the propatagium and
dactylopatagium minus (the membrane between the second and
third digits) make with the posterior parts of the wing membranes.
During flight the air pressure against the ventral surfaces of the
wing membranes is greater than that against the dorsal surfaces;
this helps to maintain a smooth camber. The wings seem makeshift
40 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
and inefficient due to their irregular shape, but this irregularity is
the source of increased efficiency in certain parts of the wing. The
tapering distal part of the wing gives the wing a fairly high aspect
ratio (ratio of the length of a wing to its width; in wings of irregular
outline the aspect ratio is considered as the ratio of the square of
the wing span to the total wing area) and serves to reduce the
turbulence and loss of lift at the end of the wing as it passes through
the air (this loss of lift is called “end effect”). The proximal part
of the trailing edge of the plagiopatagium merges fairly evenly into
the calcar portion of the uropatagium, thereby producing a smooth
transition from wing-base to tail membrane and tending to avoid
turbulence and drag at this point of junction. Bearing in mind the
basic pattern of the wing of the bat, it is instructive to compare the
wings of the three bats under discussion using as a basis certain
simple aerodynamic principles plus what is known of the flight of
birds having similar wings.
The wing of Eumops is long and slender, and has many of the
characteristics known in birds to be associated with rapid and en-
during flight. Compared to the other two bats the wing of
Eumops has low camber. In the proximal segment of the wing the
low camber results in part from a relatively narrow propatagium that
is inclined downward at only a moderate angle, and in the chiro-
patagium the low camber is caused by the small size of the dactylo-
patagium minus, which is inclined downward at only a slight angle
to the rest of the chiropatagium. A fairly low camber is typical of
high-speed wings in birds and serves to reduce drag. The chiro-
patagium is less broad than the proximal segment of the wing and
tapers to a narrow rounded tip. The tip of the wing, starting from
the end of the fourth digit, is shaped strikingly like the distal part
of the notched primary wing feather of a bird. By reducing the
end effect, the notched primaries of birds increase the lift produced
by a wing. Likewise, the slender wing-tip of Eumops reduces the
loss of lift at the wing-tip due to the end effect. The shape of the
posterior edge of the plagiopatagium is such that it blends smoothly
with the uropatagium and probably little turbulence is created at
this point. The anterior edge of the propatagium curves forward
near its base, creating a fairly smooth junction with the body. The
remarkably high aspect ratio of the wing of Eumops is strong
evidence indicating that this bat is adapted to fly for long periods
of time at a moderately low energy output. The aspect ratio of the
wing of the mastiff bat is considerably higher than that of the
herring gull (Larus argentatus) and is thus probably above that of
FUNCTIONAL MORPHOLOGY OF THREE BATS Al
most gulls. A long wing of high aspect ratio has been developed by
various sea birds noted for their ability to soar (gulls, gannets,
albatrosses). It is interesting to note that like Ewmops some birds
with high-aspect-ratio wings have difficulty taking off from flat
surfaces, and need to take off into a wind or launch themselves from
a cliff. Certain birds capable of rapid flight, such as the golden
plover (Pluvialis dominica) and chimney swift (Chaetura pelagica)
have high-aspect-ratio wings (Savile, 1957:215). While it is doubt-
ful that Eumops does any soaring, it is known that this bat’s flight
is rapid, and the aerodynamic efficiency of its wings is probably
correlated with the animal’s ability to fly for prolonged periods.
Although their description does not logically belong in a study of
appendicular morphology, the axial parts of the bats deserve men-
tion here because of their aerodynamic importance. Eumops has
large ears, and at first glance it seems that they would serve as
funnels and would produce considerable drag during flight. Upon
close examination, however, the ears can be seen to be crude airfoils
that function as lifting surfaces during flight. Compared to most
bats the head of Eumops is extremely large and the dentition is
robust, and yet the neck, which would be expected to support the
head during flight, is not particularly heavily built and seems hardly
adequate to support the head throughout long flights. When this
bat is in flight, in fact whenever the animal is not alert to danger or
looking at nearby objects, the thick, flat keels at the ventral bases of
the ears cover the eyes and lie along the sides of the head (Fig. 1),
and the broad ears face ventrolaterad with the pinnae extending
laterad. In this position, the ears resemble short, stubby wings,
even to the dorsal arching (camber) of the pinnae and the con-
tinuous leading edge. Because of their shape and position with
respect to the direction of flight, the ears produce lift during fight,
and this tends to raise the head. The lift supplied by the ears
probably has the important function of tending to hold up the head,
thereby supplementing the action of the muscles that elevate the
head. To a bat that has a large head and flies for long periods of
time any such conservation of energy would be advantageous.
The position of the ears in Eumops is characteristic of many of the
members of the family Molossidae; in this position the ears produce
a minimum of drag during flight, this being of importance in a group
of bats that fly rapidly and many species of which fly for long periods
of time.
The head, the dorsally arched body and the uropatagium of
Eumops form a crude airfoil of high camber. In this bat the fairly
42 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
flat head, the long shallow body and short but moderately broad
uropatagium seem to form a more conventional airfoil than that
formed by the corresponding parts of the other genera. The heads
and bodies of Myotis and Macrotus are fairly deep and the
uropatagial membranes are larger than that of Eumops. The airfoil
formed by the head, body and uropatagium of Eumops is a more
nearly flattened section than the airfoils formed by the corresponding
parts of the other genera, and is better adapted to rapid flight. It is
known that the body of an airplane often supplies considerable lift;
in some cases this amount of lift is greater than the amount that
would be produced by an equally broad section of wing. Taking
this into account, it is probable that in a bat the head-body-
uropatagial section contributes a significant proportion of the lift
produced by the animal in flight.
The uropatagium is important as a braking surface as well as a
lifting surface, and, other things being equal, the larger the
uropatagium in relation to the total weight of the bat the greater
the ability to maneuver. In Eumops the loading of the uropatagium
(ratio of weight of bat in pounds to surface area of uropatagium in
square feet) is higher than in the other two genera. Compared
to these bats the wing loading of Eumops is also higher, and this
demands greater flying speeds and supplies less surface for braking
in connection with sudden changes of direction. Thus it would be
supposed that Eumops flies more rapidly and less maneuverably than
the other two bats. Observations in the field support this sup-
position.
Relative to Eumops, the wings of Myotis velifer are not as long
and slender, and they have the characteristics usually associated
with a low-speed, high-lift wing. The camber of the entire wing
is high because of the large propatagium that is pulled downward
at a fairly sharp angle to the plagiopatagium, the relatively large
dactylopatagium minus that extends downward at an angle to
the rest of the chiropatagium, and the long fifth digit, the phalanges
of which are flexed fairly strongly ventrad during flight. The tip of
the wing is tapered abruptly and probably tends to avoid the end
effect, but seemingly considerably more lift is lost at the wing-
tip of Myotis than at the slender wing-tip of Eumops. Because
of the pronounced taper of the distal half of the chiropatagium the
wing has a fairly high aspect ratio. The high camber of the entire
wing and the breadth of the proximal segment of the wing clearly
indicate that the wing is adapted to low flying speeds. Myotis
velifer has a lower wing loading and the uropatagium is much
FUNCTIONAL MORPHOLOGY OF THREE BATS 43
larger relative to the body weight than in the other genera; thus it
has large flight surfaces that allow high maneuverability. The wing
loading of this bat is below that of any bird on which I can find
data available in the literature. Even allowing for the inefficiency
of the upstroke of the wing-beat cycle in bats, the characteristics of
the wing of M. velifer indicate that it is probably better adapted
than any bird to low-speed, highly maneuverable flight at a mod-
erate level of energy output. The mode of foraging of this small
bat would seem to demand such adaptation.
The wings of Macrotus are relatively shorter and broader and of
higher camber than the wings of the other two bats, and are adapted
to low-speed flight and high maneuverability. The propatagium
and dactylopatagium minus are large, and during flight are pulled
sharply downward at an angle to the more posterior parts of the
wing membranes. The fifth digit is longer relative to the other
digits than in the other genera, and the phalanges retain consider-
able ventral flexion during flight. The wing loading is far lower
than in Eumops and but slightly higher than in Myotis velifer, and
the aspect ratio is the same as that of M. velifer. Thus, in terms of
general aerodynamic characteristics, Macrotus californicus closely
resembles Myotis velifer. In the former the shape of the distal part
of the wing is such that more lift is lost due to the end effect than
in the other genera. The large ears, which face almost directly for-
ward during flight, the short, deep body, and the posture of the
hind limbs make these parts of the animal form a far less efficient
airfoil than is formed by the corresponding parts in the other two
bats. Macrotus californicus seems to spread its uropatagium widely
(by spreading the legs apart) only when hovering or performing
other maneuvers requiring a departure from straight level flight.
The loading of the uropatagium is greater than in Myotis velifer but
is less than half that of Eumops perotis. Judging from aerodynamic
considerations of Myotis velifer and Macrotus californicus, the for-
mer might be expected to be more maneuverable than the latter.
This is known not to be the case, and the extreme maneuverability of
M. californicus is probably largely due to the specializations of its
sensory equipment.
The proportions of the segments of the wings of the three bats
show considerable variation. The area of the chiropatagium is
larger relative to that of the proximal segment of the wing in
Macrotus than in the other genera. This ratio may simply be a re-
sult of the greater degree of taper in the wings of Eumops and
Myotis, and hence their greater efficiency in the utilization of lift.
Relative to the proximal segment of the wing, the chiropatagial seg-
44 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
ment is much longer in Eumops than in the other bats. Perhaps this
reflects a reduced dependency on flapping flight in Ewmops, for in
birds, as a general rule, elongation of the distal segments of the
wings relative to the total length of the wing and of the body is
associated with soaring flight, whereas relatively short distal seg-
ments are found in species using predominantly flapping flight
(Fisher, 1955:84). This generality has many exceptions among the
birds; applied to the bats here considered, however, it may find
some support, since observations of these bats in the field indicate
that Eumops does beat its wings more slowly than do the other bats,
and, at least when landing and taking off, performs dives and rapid
glides during which the wings are set and do not beat.
OSTEOLOGY
Introductory Remarks
Bats normally hold their forelimbs more or less out from the sides
of the body and not vertical to the substrate as do cursorial mam-
mals. Therefore, the dorsal surface of the humerus in bats cor-
responds to the lateral surface in most mammals. The carpus of
bats is so modified that the manus is always in a partly supinated
position; the palmar surface is directed mediad (or ventrad, when
the wing is extended to the side). Hereinafter, descriptive terms
are applied to the forelimbs of the bats under study as these terms
are applied to the forelimbs of other mammals, by assuming that
the forelimb is oriented vertically to the substrate. Regarding the
manus, however, the descriptive terms are applied as if the limb
were held outstretched laterally as it is when extended for flight.
Thus, the pollical side of the manus is termed the anterior side and
the opposite side, the posterior side; the palmar surface is termed
the ventral surface and the opposite surface, the dorsal surface.
The hind limbs of cursorial mammals are carried vertical to the
substrate with the plane of movement of the joints directed antero-
posteriorly. This is not the posture of the hind limbs in bats; in
fact, none of the three bats under study is even able to bring its
hind limbs into this position. When Eumops and Myotis are in level
flight the hind limb is held out to the side, with the femur extending
laterad and slightly dorsad at an angle of roughly 90 degrees to the
long axis of the body. The shank extends caudad and slightly ven-
trad, but is approximately parallel with the long axis of the body.
In Macrotus the position of the hind limbs during level flight differs
even more from that in most other mammals. The hind limbs are
held behind the bat in a spider-leg-like posture, with the femur ex-
FUNCTIONAL MORPHOLOGY OF THREE BATS 45
tending dorsad and caudad, and with the shank partially flexed and
extending caudad and more or less downward. In Eumops and
Myotis, the hind limb during flight is as if it has been rotated 90
degrees from the position of the limb in cursorial mammals. In
Macrotus the rotation amounts to nearly 180 degrees. Thus, the
lateral surface of the shank in most mammals is homologous with the
medial surface of the shank in Macrotus. In Eumops and Myotis,
and in other bats that crawl with any facility, the hind legs are used
in reptilian fashion, being held out to the side of the body rather
than beneath it; the femur extends laterad and more or less dorsad
from the body, and the shank is flexed at right angles to the thigh
and is held roughly vertical to the substrate. The descriptive termi-
nology will be applied in the same way to the hind limbs of the bats
considered here as it is to the hind limbs of other mammals. It
should be stressed, however, that these terms apply only when the
hind limbs of bats are put in the posture typical for terrestrial mam-
mals, and that normally the limbs of bats are not in these positions.
The discussions of the functional significance of many of the
osteological specilizations found in bats are reserved for the section
on conclusions.
The osteology of Eumops is described in some detail, whereas the
osteological descriptions of Myotis and of Macrotus are essentially
comparisons between the elements in those genera and in Eumops.
When a bone is said to be larger in one bat, the bone is larger rela-
tive to the size of the body than is the corresponding bone in the
other bat, or bats, used in comparison.
Vertebral Column
The bodies of most bats are short and thick as a result of the anteroposterior
compression of the individual vertebrae and the marked dorsal arching of the
thoracolumbar section of the vertebral column. In the three genera studied
the arching is most pronounced in Macrotus and least so in Eumops. Owing
mostly to the difference in arching, Macrotus is relatively short, broad and
deep, Eumops is long, narrow, and shallow, and Myotis is intermediate.
The numbers of postcervical vertebrae in the three genera are as follows:
Eumops, thirteen thoracic, six lumbar, five sacral, ten caudal; Myotis, eleven
thoracic, five lumbar, five sacral, ten caudal; Macrotus, twelve thoracic, six
lumbar, five sacral, seven caudal. Only thoracic, lumbar and sacral vertebrae
are discussed; on them many appendicular muscles have their origins.
Thoracic Vertebrae
Eumops.—These vertebrae are roughly one half as long as high. The first
is the largest of the thoracic series and each succeeding vertebra is smaller
to the seventh thoracic, from which point caudad all the thoracics are approxi-
mately the same size. The centra of the first twelve thoracics are flat anteriorly
46 UNIVERSITY OF KANnsAs Pusts., Mus. Nat. Hist.
and posteriorly; those of the last two are procoelous. The centra are dorso-
ventrally compressed, and the dorsal surfaces are slightly concave. The ver-
tebral foramen is largest in the first several thoracics, becoming progressively
smaller toward the posterior end of the thorax. The laminar parts of the neural
arches are nearly flat through the anterior half of the series, but become more
strongly arched toward the posterior end of the column.
The neural spine on the first thoracic vertebra is prominent, broad, and
knoblike, and provides part of the origin for the large anteriormost part of the
trapezius group of muscles. On the second thoracic vertebra, a low medial
neural ridge merges with two weak lateral ridges. They give rise to a pair of
low tuberosities at the posterior edge of the laminar part of the neural arch.
From thoracics two to six the medial ridge diminishes in size but the lateral
ridges persist, joining medially to form a low neural ridge on the seventh
thoracic. This ridge increases in breadth and height throughout the rest of
the thoracic series.
In dorsal view, the anterior edges of the neural arches are concave, the
posterior edges are convex, and there are spaces between the laminar parts of
adjacent neural arches. These spaces are largest at the level of the fifth thoracic
and become smaller both anteriorly and posteriorly from this point.
The short transverse processes extend dorsad and slightly laterad from the
lateral edge of the nearly flat laminar part of the neural arches, rise above the
highest parts of the arches in thoracics one to five, and become smaller toward
the posterior end of the series. There are no transverse processes on the last
four thoracics.
The zygapophyses of the first three thoracics are short and broad and the
articular surfaces of the anterior zygapophyses face dorsolaterad. (The pos-
terior articular surfaces always face in the direction reciprocal to that of the
anterior articular surfaces.) The zygapophyses of thoracics four to eight are
long and fingerlike. A small pointed tuberosity (metapophysis) extends
craniad from the anterior surface of the top of the transverse processes of
thoracics one to eight. The anterior zygapophyses project progressively more
sharply dorsad toward the caudal end of the thoracic series, until, on the tenth
thoracic, they unite with the metapophyses to form a large anterior articular
process. The cranial articular processes of the last four thoracics, then, are
formed partly by the anterior zygapophyses and partly by the metapophyses.
The anterior articular surfaces are concave and the posterior surfaces convex,
thus forming a strong but incomplete tongue-in-groove joint that probably
limits dorsoventral bending of the vertebral column. The cranial articular sur-
faces face dorsolaterad in thoracics one to seven, roughly dorsad in thoracics
eight and nine, and directly mediad in the rest of the thoracics. The last three
thoracics bear accessory processes.
The articular facets for the tubercula of the ribs face laterad. The caudal
demifacets are large, face caudolaterad, and are surrounded by bone much
thicker than that comprising the rest of the pedicle. The cranial demifacets
are smaller and less well defined than the anterior demifacets, but are similarly
braced by thick bone. The ribs make contact not only with the main articular
facets but also with the anterior parts of the pedicle to which the medial sur-
faces of the ribs between the tubercula and capitula are bound by fascia.
The caudal intervertebral notches are large at the anterior end of the thorax
and extend craniad farther than the level of the middle of the centrum, but
FUNCTIONAL MORPHOLOGY OF THREE BATS 47
become progressively smaller toward the posterior end of the thorax. The
cranial intervertebral notches are small and are covered by the ribs.
The first thoracic vertebra is the largest of the thoracic series and merits
special consideration. Its centrum and anterior zygapophyses are fused with
the centrum and posterior zygapophyses of the last eervical vertebra leaving
no trace of the former points of articulation. A few gaps remain between the
laminar parts of the vertebrae; the intervertebral foramen is unconstricted. The
large, dorsoventrally elongate facet for the articulation of the head of the first
rib extends ventrad to the level of the slightly concave ventral surfaces of the
fused vertebrae. The facet for the articulation with the tuberculum of the
first rib is on the ventrolateral surface of the short, thick transverse process.
The proximal part of the greatly enlarged first rib is in contact with most of
the lateral surface of the first thoracic vertebra. The transverse processes of
the first thoracic vertebra extend more nearly laterad than those of the rest of
the thoracics and provide origin for the tendons of the anteriormost part of the
trapezius group of muscles.
Myotis.—The spaces between the neural arches are relatively smaller; the
transverse processes project more nearly laterad and persist throughout the
thoracic series; the first thoracic and last cervical are not fused; fingerlike
projections (metapophyses) extend anteriorly from the dorsal edge of the cranial
articular processes of the last four thoracics and partly enclose the tops of the
concave anterior articular surfaces. Although there is no bony fusion of the
first thoracic and last cervical vertebrae, there is little movement possible be-
tween them. The intervening intervertebral disc is thin and the anterior
articular surface of the first thoracic faces anteromediad and makes a tight
junction with the posterior articular surface of the last cervical. Two bony
projections from the posterior margin of the lamina of the last cervical make
contact with the anterior margin of the lamina of the first thoracic and provide
additional reinforcement.
Macrotus.—The thoracic vertebrae of this bat differ considerably from those
of the other two genera. The spaces between adjacent neural arches are nar-
row and these, except for the last four, are bridged by two bony projections
from the posterior edges of the lamina. The first thoracic is not fused with
the last cervical and they fit together approximately as they do in Myotis. A
pair of ridges, one on each side of the midline, occur on thoracics one to nine.
These lateral ridges become smaller posteriad and converge to form a low
medial ridge on thoracics eleven and twelve. The articular surfaces of the
anterior zygapophyses face dorsomediad throughout the thorax and there is
no tendency toward the development of a tongue-in-groove articulation anterior
to the last thoracic. From the dorsal part of the anterior zygapophyses of this
vertebra, fingerlike metapophyses extend over the dorsolateral part of the
posterior zygapophyses of the eleventh thoracic.
Lumbar vertebrae
Eumops.—There are six lumbar vertebrae in this genus. In general they
resemble the last four thoracics but are more robust and relatively longer.
The lumbar centra are all procoelous. The intervertebral discs posterior to the
last three lumbars are appreciably thicker than those of the thoracic vertebrae,
and the intervertebral foramina become progressively larger toward the caudal
end of the lumbar series.
48 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
The lateral surfaces of the centra of lumbars three to five bear low ridges
that terminate caudally on each vertebra as small tuberosities. These ridges
are inclined ventrad and caudad and considered together, form a ridge from
the level of the top of the centrum on the anterior edge of the third lumbar
to the ventral surface of the posterior edge of the centrum of the fifth lumbar.
Dorsal to this ridge the lateral surfaces of lumbars three to five are flattened
and give origin to the M. psoas major and M. ‘iliacus. From the ridge itself
the Mm. psoas major and minor take origin.
When viewed dorsally the tops of the neural arches are notched anteriorly
and posteriorly, and only narrow spaces are present between adjacent arches.
Broad, low neural ridges occur on all the lumbars; on the last two the ridges
become spinelike and are inclined caudad.
The anterior zygapophyses of the lumbars are much like those of the last
four thoracics but are larger, extend dorsolaterad, and their articular surfaces
face dorsomediad. The posterior zygapophyses are broader and more robust
than those of the last four thoracics.
Myotis.—This genus has five lumbar vertebrae. The lumbars are pro-
coelous, but the curvature of the anterior and posterior surfaces of the centra
is slight. No definite ridges are present on the lateral surfaces of the last few
lumbars along the line of origin of the iliacus group of muscles, The ventral
surface of the centrum of the fourth lumbar has a pair of low, rounded ridges,
and the pedicle has a small tubercle that probably represents a transverse
process, The last lumbar has a short transverse process that projects craniad
and bears an extremely reduced neural ridge. Only the first lumbar bears
accessory processes. The thickest intervertebral disc of the thoracolumbar
series separates the centra of the last lumbar and first sacral vertebrae; the
articulation is between two convex surfaces.
Macrotus.—This genus has six lumbar vertebrae and, as in Myotis, the
centra are only weakly procoelous. In comparison to the other two genera,
there are several characters of the lumbar vertebrae peculiar to this genus.
A pair of ventral ridges occurs on the centra of lumbars four and five and
the sixth lumbar bears a short, broad mid-ventral spine that projects craniad.
Lumbars three to five have small transverse processes. The metapophyses
are pointed and unusually large. The top of the neutral arch of the sixth
lumbar is flat and shows no indication of a neural spine, but a cranially
projecting transverse process is present. The sixth lumbar is strongly com-
pressed anteroposteriorly; the posterior surface of its centrum and the anterior
surface of the first sacral centrum are both convex and produce a joint that
allows considerable movement.
Sacral Vertebrae
Eumops.—The four sacral vertebrae are completely fused. They remain
roughly the same breadth as the posterior lumbars, but, starting with the
first sacral, are abruptly compressed dorsoventrally in the parts beneath the
transverse processes; as a result the last sacral is approximately three fifths
as high as the lumbars. The diameter of the vertebral canal is greatly re-
duced toward the caudal end of the sacrum. The sacral foramina are distinct,
becoming smaller caudally. The neural spine of the first sacral is low and
thick and resembles, in general, the spine of the last lumbar; the spines of the
FUNCTIONAL MORPHOLOGY OF THREE BATS 49
rest of the sacrals are thin but are higher and longer than those of the lumbars.
Only the bases of the neural spines are fused. The zygapophyses of the first
sacral are roughly the same as those of the lumbars but bear no metapophyses.
The transverse processes are fused to form a broad lateral mass that is thick
in sacrals one and two and thin in the last two sacrals. The thick part of the
lateral mass is fused to the ilium; the fusion is continuous and extends from
the crest of the ilium to a point approximately 2 mm. anterior to the level
of the acetabulum.
Myotis.—There are four sacral vertebrae. Neural spines on the first two
are more nearly fused and the lateral masses of these vertebrae are relatively
thinner than in Eumops.
Macrotus.—The sacrum of this genus differs from that of the other two
genera as follows: composed of five vertebrae the last of which is incom-
pletely fused with the fourth; in dorsal view the last three sacrals are much
narrower than the first two; intervertebral foramina are more strongly reduced;
neural spines of the first four sacrals fused, forming a low, thin crest from
which the neural spine of the last sacral is separate; sacrum is more lightly
built and is arched slightly ventrad.
Sternum
Eumops.—The sternum is composed of three bones separated by discs of
cartilage. The longest element is the body of the sternum, the manubrium
is slightly shorter, and the xiphoid process is the shortest part.
Viewed ventrally the manubrium looks like an asymmetrical cross. The
manubrium has two short lateral arms, a slightly longer anterior arm, and a still
longer posterior arm. The anterior arm is clavate as viewed from the side
and is laterally compressed. It is inclined ventrad and forms an angle of
approximately 30 degrees with the long axis of the posterior arm. The lateral
arms are short and thick, roughly triangular in cross section, with one flat
surface facing ventrad. The lateral arms curve gently dorsad; their distal
ends are expanded and terminate in flat articular surfaces that face antero-
laterad and against which lie the sternal ends of the clavicula and first costal
cartilages. The posterior arm of the manubrium is roughly circular in cross
section; its dorsal surface is flattened and the posterior end is expanded and
truncate. A low, inconspicuous ridge extends the length of the ventral sur-
face of the manubrium.
The body of the sternum is straight and nearly circular in cross section
anteriorly, becoming increasingly dorsoventrally flattened caudally. The an-
terior end is expanded and smooth, and meets the smooth posterior articular
surface of the manubrium. A heavy band of connective tissue binds together
the ventral part of this joint, whereas the dorsal part of the joint is free to
gape open slightly. This creates a hinge joint that allows the posterior part
of the sternum to swing ventrad but does not allow it to move dorsad beyond
the point at which it is in line with the posterior arm of the manubrium.
There are small tubercles on the lateral surfaces of the body of the sternum
to which the costal cartilages of ribs three to seven attach. A low, poorly
defined ridge extends the length of the ventral surface of the body. The
anterior half of the xiphoid process is narrower than the posterior end of the
body of the sternum and is approximately square in cross section. The xiphoid
50 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
process becomes progressively broader and more dorsoventrally flattened cau-
dally; the posterior third is cartilaginous. The cartilaginous extension is broad
and flat and the posterior margin is rounded. The connection between the
body of the sternum and the xiphoid process allows but little movement.
Myotis.—Two separate elements make up the sternum of this genus, an
anterior part, the manubrium, and a posterior part consisting of the fused
body of the sternum and the xiphoid process.
The manubrium is approximately half as long as the posterior part of the
sternum. As seen ventrally the manubrium is T-shaped. The anterior arm
is roughly as long as the posterior part and is laterally compressed; it projects
ventrad at nearly right angles to the posterior arm and therefore is seen end-
wise when the manubrium is viewed ventrally. The lateral arms are broad,
dorsoventrally flattened, and project slightly dorsad. The distal ends are
thickened and bear flat articular surfaces that face craniolaterad. The pos-
terior arm is broad and dorsoventrally compressed. A thin ridge extends from
the posterior surface of the anterior arm of the manubrium to the posterior
end of the ventral surface of the posterior arm.
In general shape the posterior part of the sternum in Myotis and Eumops
is nearly alike. In Myotis costal cartilages three to six attach to the lateral
surfaces of the body of the sternum. A low ridge extends along the entire
ventral surface of the body of the sternum and along most of the ventral
surface of the xiphoid process. The posterior end of the xiphoid process has
a short, cartilaginous extension.
Macrotus.—The sternum in this genus consists of a manubrium and a single
posterior part which represents the fused body of the sternum and the xiphoid
process.
Discounting the cartilaginous caudal extension on the xiphoid, the manu-
brium is less than half as long as the posterior part of the sternum. Viewed
ventrally the manubrium is approximately twice as broad as high and is shaped
like a “T,” with the lateral arms flared upward and the vertical bar shortened.
The part of the manubrium that extends craniad in Eumops, and that here is
termed the anterior arm, extends ventrad in Macrotus at approximately right
angles to the long axis of the posterior arm and is therefore seen endwise when
the manubrium is viewed ventrally. The anterior arm is short and broad an-
teroposteriorly, but is laterally compressed. From the middle of the ventral
edge of the anterior arm, a thin keel extends directly posteriad to the caudal
end of the posterior arm of the manubrium. The lateral arms arch gently
dorsad and craniad; they become broader distad and are dorsoventrally com-
pressed. The articular surfaces for the bases of the clavicula face anterolaterad
from the expanded ends of the lateral arms. The posterior arm of the manu-
brium is short, not so broad as the other arms, and is dorsoventrally com-
pressed. It has a prominent keel on the ventral surface. The posterior end of
the posterior arm is truncate and by means of a movable joint articulates with
the anterior end of the body of the sternum.
The body of the sternum is narrow and straight, flattened on its dorsal
surface, and bears a high ventral keel that extends along the entire posterior
part of the sternum. The keel is highest at the middle of the posterior part
of the sternum and gradually diminishes in height craniad and caudad from
this point. Costal cartilages three to seven connect onto the lateral surfaces
FUNCTIONAL MORPHOLOGY OF THREE BATS 51
of the body of the sternum. The xiphoid process becomes only slightly
broadened at its posterior end and gives way to a large, bifurcate cartilaginous
extension.
Ribs
In the bats under study the rib cages are large compared, for example,
to those of rodents of comparable sizes, and are broader than deep. The rib
cage of Eumops is long and shallow, whereas that of Macrotus is relatively
short and deep. In terms of shape, the rib cage of Myotis is roughly midway
between these extremes. Eumops has thirteen ribs; the first seven are verte-
brosternal, the next four are vertebrocostal, and the last two are vertebral.
The corresponding arrangement in Myotis is eleven, six, three, and two; in
Macrotus it is twelve, seven, three, and two.
The ribcages of all three genera have certain characteristics in common.
The first rib is short but is considerably thicker and more strongly built than
the other ribs. It is anteroposteriorly compressed and the entire medial surface
of the elongate proximal end of the rib, from the tuberculum to the capitulum,
is in contact with the lateral surface of the anterior part of the first thoracic
vertebra. The first costal cartilage is the broadest and thickest of the series
and attaches to the lateral arm of the manubrium. The first three ribs are
more or less anteroposteriorly compressed, whereas the rest of the ribs are
flattened in the opposite plane. When viewed laterally, the shafts of the ribs
seem to be broadened by thin margins of bone that extend craniad and caudad
from the thicker central part of the shaft. In Eumops the inner surfaces of ribs
four through eleven are reinforced by a thin ridge of bone; in cross section, the
anterior ribs are roughly T-shaped but the posterior ribs become triangular.
In all three genera the ribs are two headed excepting the last two or three,
in which the heads tend to merge. The tubercula articulate with the transverse
processes and the capitula contact the articular facets of the centra; in addition,
the entire medial surfaces of the proximal ends of the ribs, from the tuberculum
to the capitulum, are in contact with the vertebrae.
Pectoral Girdle and Limb
Scapula
The scapulae in these bats lie dorsal to the ribcage with their long axes
roughly parallel to the vertebral column.
Eumops.—The scapula is long (breadth approximately 40 per cent of
length) and the post-spinous part is irregularly trapezoidal as viewed dorsally.
The round anteromedial border of the supraspinous fossa gives way laterally
to a deep suprascapular notch. The scapula is broadest anteriorly, at the level
of the glenoid fossa, and becomes narrowed caudally. The supraspinous fossa
is roughly one third as large as the infraspinous fossa. The entire rim of the
scapula is thick, but the intervening surfaces are mostly thin and semitranspar-
ent. The axillary border and the part of the scapula between the glenoid
fossa and the bases of the acromion and coracoid processes are especially
heavily reinforced by thick bone.
The glenoid fossa faces almost directly laterad. It is irregularly shaped
and is elongate anteroposteriorly. The broad part is posterior and the narrow
part anterior; the dorsal edge of the fossa is concave and the ventral edge is
straight. A large, blunt-pointed supraglenoid tuberosity projects laterad from
52 UNIVERSITY OF KANSAS Pusts., Mus. Nar. Hist.
the anterior rim of the fossa. There is a smooth depression in the dorsal surface
of the scapula just medial to the anterior part of the glenoid fossa against which
the greater tuberosity of the humerus rests when this bone is flexed and raised.
A low infraglenoid tubercle arises from the thick axillary border of the scapula
2 mm. posterior to the glenoid fossa.
The supraspinous fossa is inclined ventrad so that its flat dorsal surface
faces anteromediad and is at an angle of roughly 145 degrees to the plane
of the post-spinous part of the scapula. A narrow rim extends ventrad from
the lateral border of the fossa. This rim enlarges anteriorly into a flat flange
that extends ventrad at approximately right angles to the plane of the post-
spinous part of the scapula. The flange increases in breadth and thickness
anteriorly and is roughly triangular as seen from the front.
The low scapular spine is not perpendicular to the plane of the post-
spinous part of the scapula but inclined slightly craniad. The lateral edge of
the spine is thick and bears a long acromion process. A ligamentous sheet
spans the gap from the medial edge of the spine to the dorsal part of the
medial end of the acromion process and provides additional surface for the
attachment of the Mm. supraspinatus and infraspinatus. The acromion is
thick proximally and narrows distally; from the middle of its lateral surface
projects a short metacromion process. The rounded distal tip of the acromion
is flattened on its ventral surface where it contacts the distal end of the
clavicle.
The long coracoid process arises from the anterolateral angle of the scapula.
The coracoid projects ventrad for roughly one third of its length, then it turns
and the distal two thirds extends ventromediad and slightly caudad. The tip
of the coracoid is ventral to the lateral part of the suprascapular notch. The
flat proximal third of the coracoid is broad as seen from the front; the distal
two thirds is thinner, roughly elliptical in cross section, and the tip is expanded.
From the lateral apex of the triangular flange of the supraspinous fossa a liga-
mentous sheet extends to the medial surface of the distal half of the acromion
and the medial base of the coracoid. This sheet increases the area of origin
of the M. supraspinatus.
The surface of the infraspinous fossa is divided into three facets that
are set at angles to each other; the bone at the intersection of these facets is
slightly thickened. This faceting serves to increase the area of origin of the
Mm. infraspinatus and subscapularis. Working from the vertebral border
laterad, the anteromedial facet faces caudolaterad and is tilted downward;
the intermediate facet faces craniomediad and is tilted upward; the postero-
lateral facet faces caudolaterad and is tilted downward. As seen from above
the intermediate and posterolateral facets are slightly concave and the antero-
medial facet is convex. Viewed dorsally, the medial part of the infraspinous
fossa is a broad trough formed by the anteromedial and intermediate facets;
seen ventrally, the lateral part of the infraspinous fossa is a trough formed
by the intermediate and posterolateral facets and a second trough is formed by
the anteromedial facet of the infraspinous fossa and the entire supraspinous
fossa. Extending caudad from the posterior end of the scapula is a carti-
laginous extension that tapers to a point caudally. This extension provides
extra surface for muscle attachment.
Myotis.—The scapula of this genus is slightly shorter and broader (breadth
approximately 43 percent of length) and less strongly built. The supraspinous
FUNCTIONAL MORPHOLOGY OF THREE BATS So
fossa is roughly one quarter as large as the infraspinous fossa. The glenoid
fossa faces laterad and slightly craniad; the supraglenoid tuberosity is less
prominent. The surface of the supraspinous fossa is concave as viewed dorsally,
is inclined only slightly downward, and the lateral rim is small. The scapular
spine it low. The acromion is slightly shorter and bears no metacromion.
The coracoid process is approximately the same breadth throughout its length;
the proximal three quarters curves gently laterad and the distal quarter bends
sharply laterad. The tip of the coracoid is ventral to the glenoid fossa. The
posterior cartilaginous extension of the scapula is small.
Macrotus.—The scapula is less specialized than those of Eumops and Myotis
and differs from them in many details. The scapula is shorter and broader
(breadth approximately 50 per cent of length) and irregularly ovoid. The
supraspinous fossa is considerably more than one third as large as the infra-
spinous fossa and is bounded medially by a small suprascapular notch. The
glenoid fossa faces craniolaterad and the supraglenoid tuberosity is small. The
supraspinous fossa has only a weakly developed rim and no anterior flange.
The distal third of the acromion is flat and its finger-shaped tip extends
ventrad to within roughly 0.5 mm. of the dorsal surface of the base of the
coracoid. The broad, flat coracoid process curves steadily laterad and its
tip extends beyond (lateral to) the glenoid fossa. There is no posterior
cartilaginous extension on the scapula.
Clavicle
This element articulates proximally with the lateral arm of the manubrium
in all three genera. In Eumops and Myotis the distal end of the clavicle is
bound by ligaments to the dorsal base of the coracoid process and the ventral
surface of the tip of the acromion process of the scapula; in Macrotus the
clavicle is attached distally to the dorsomedial surface of the base of the
coracoid process.
In Eumops the proximal third of the clavicle is nearly straight and is
directed slightly forward, whereas the distal two thirds curves caudad. As
seen from the front it is slightly S-shaped, with the proximal half curving
ventrad and the distal half curving dorsad. The sternal base is large, nearly
circular in cross section, with a smooth articular surface. As seen from above
the clavicle is broad at its base, has a slight constriction just distal to the base,
and becomes broader distally throughout the proximal third. The distal two
thirds of the shaft becomes progressively narrower. The shaft is dorsoventrally
compressed; the dorsal surface is slightly convex and most of the ventral
surface is concave. The distal end is knoblike and inclined posteriad; its
posterior surface is flattened.
The clavicle is more slender in Myotis. As viewed from above the shaft is
is bow-shaped; it is curved throughout its length, the curvature increasing
distally. The bone is S-shaped as seen from the front, but the curvature is
less pronounced than in Eumops. The shaft remains nearly constant in breadth
from end to end and is roughly elliptical in cross section, having the ventral
surface slightly flattened.
The clavicle in Macrotus is similar to that in Myotis, but is more robust and
is dorsoventrally compressed.
54 UNIVERSITY OF KAnsAS PuBLis., Mus. Nat. Hist.
Humerus
In the bats considered here the humerus is long and slender and the ridges
for muscle attachment are on the proximal fourth of the bone. The humerus
is shorter and thicker in Eumops than in the other two bats. The diameter
of the shaft is approximately 36 per cent of the width of the proximal epiphysis
and 53 per cent of the width of the distal epiphysis; the diameter of the
humerus is approximately 3.2 per cent of its length. Corresponding percent-
ages for the other two genera are as follows: Myotis, 34, 37, 4.3; Macrotus,
33, 30, 4.3.
Eumops.—The head of the humerus is large and extends caudad from the
posterior edge of the shaft a distance roughly equal to the diameter of the
shaft. There is no neck; the head merges into the posterior part of the
shaft and the tuberosities. If the humerus is viewed from the proximal end
the head is ovoid and inclined mediad; the thickest part lies immediately
posterior to the dorsal surface of the shaft, and the head extends craniad and
mediad from this point to merge into the lesser tuberosity.
Immediately anterior to the head is a deep pit that is bounded anteriorly
by the converging anterior ridges of the greater and lesser tuberosities and the
proximal part of the pectoral ridge. The supraglenoid tuberosity fits into this
depression when the humerus is extended to approximately a right angle with
the lateral edge of the scapula. This arrangement serves as a locking mecha-
nism to limit the forward movement of the humerus and prevent it from being
extended beyond roughly a right angle with the body.
The greater tuberosity projects dorsolaterad well beyond the head of the
humerus, is slightly flattened on its medial surface, and is broad and rounded
as viewed laterally. Its anterior edge forms a ridge that continues distad to
join the proximal end of the pectoral ridge. The entire medial surface of the
greater tuberosity is smooth and forms an articular surface that merges pos-
teriorly with the head and distally continues into the depression for the supra-
glenoid tuberosity. When the wing is flexed and raised, as it is in the up-
stroke of the wing-beat cycle, the medial surface of the greater tuberosity
makes contact with a smooth-surfaced depression just medial to the anterior
half of the glenoid fossa. This contact is made when the humerus reaches
an angle of approximately 25 degrees above the plane of the scapula and
locks the humerus at this angle. This tends to stop the upstroke by transfer-
ring its force to the scapula and the muscles binding the scapula to the body
(see discussion of action of posterior division of M. serratus anterior). Miller
(1907:13) thought that in some bats the greater tuberosity formed a secondary
articulation with the scapula, thereby creating a strong joint at which move-
ment was limited to a single plane. If movement were thus limited, the
plane of movement of the humerus would be anteroposterior; this movement
could not produce the wing beat. I am unable to understand Miller’s reason-
ing on this point.
The lesser tuberosity does not extend beyond the head of the humerus, is
thick proximally, becomes narrower toward its tip, and is connected by a thick
ridge to the proximal end of the pectoral ridge. Distally the lesser tuberosity
gives way to a prominent medial ridge that is turned slightly posteriad along
its edge so that its posterior surface is concave. The lateral head of the
triceps muscle takes origin partly from this concavity.
FUNCTIONAL MORPHOLOGY OF THREE BATS 55
The pectoral ridge is strongly developed and projects craniad from the
anterolateral surface of the humerus. The ridge starts at the distal base of the
greater tuberosity and ends at a point slightly less than one quarter of the
way along the humerus. The width of the ridge approximately equals the
diameter of the proximal part of the shaft of the humerus. Viewed laterally
the ridge is rounded anteriorly and reaches its greatest width roughly half
way from its proximal to its distal end. The lateral surface is flat distally but
slightly concave proximally; the anterior edge is broad and flat and extends
mediad as an acute ridge posterior to which the medial surface of the ridge
is concave and forms the lateral part of the bicipital groove. There is no
lateral ridge on the humerus.
The shaft of the humerus is nearly straight, but its distal half curves gently
craniad and dorsad. The lateral and medial surfaces of the distal half of
the shaft become increasingly flattened distally. The distal quarter of the
anterior surface of the shaft is flat. In the distal quarter of the posterior
surface of the shaft a broad, shallow longitudinal depression is bounded lat-
erally by a small ridge. The tendons and the distal sesamoid of the triceps
muscles lie in this depression,
The distal epiphysis is displaced slightly anterior to the shaft of the
humerus. The articular surface formed by the trochlea and capitulum is in
line with the main axis of the shaft when viewed anteriorly. The capitulum
is composed of two rounded ridges that are inclined laterad, slightly out of
line with the main axis of the humerus, and separated by a groove. A second
groove separates the medial ridge of the capitulum from the trochlea. This
system of ridges and grooves extends from the shallow radial fossa around
the anterior part of the distal articular surface. On the posterior side the
grooves and ridges give way to a single broad groove with a low medial ridge;
along this groove rides the pointed proximal end of the radius and the ventral
part of the proximal end of the ulna. The lateral rim of the capitulum is
prominent and is interrupted at the anterior edge of its proximal surface by
a small tubercle. Immediately proximal to the tubercle is a shallow depres-
sion bounded posteriorly by a small ridge that extends along 1.5 mm. of the
distal part of the shaft. From this ridge the Mm. extensor carpi radialis
longus and brevis take origin. The medial epicondyle bears a curved spinous
process, which extends distad beyond the trochlea. From the tip of this
process originates the M. flexor carpi ulnaris. There is a fossa at either end
of the pulley-shaped distal articular surface. The lateral fossa, which is sur-
rounded by the rim of the capitulum, gives origin to the tendon of the M.
supinator, The medial fossa is deep, extending nearly half way through the
distal epiphysis, and serves as the surface of attachment for a thick ligament
that extends from the fossa to the medial knob on the anterior end of the
radius.
Myotis.—A careful description of the humerus of Myotis is given by Law-
rence (1943). The humerus of Myotis differs from that of Eumops mainly in
the following ways. The shaft is relatively thinner and the distal part curves
more strongly craniad. The head is more nearly round, less ovoid, and pro-
jects caudad to the posterior surface of the humerus a distance greater than
the diameter of the shaft. The pit for the supraglenoid tuberosity is shallower.
Because of the placement of the glenoid fossa and the small size of the supra-
56 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
glenoid tuberosity, the humerus must be extended beyond an angle of 90
degrees with the lateral edge of the scapula before it is locked. The greater
tuberosity does not extend so far beyond the head, and does not lock until
an angle of roughly 30 degrees above the plane of the scapula is reached. The
pectoral ridge is long and low; the anterior edge is straight and reaches its
greatest height just short of its distal end. A low lateral ridge extends from
the distal part of the greater tuberosity to just beyond the level of the distal
end of the pectoral ridge. A small tubercle rises from the middle of the lateral
ridge. The distal part of the shaft is not flattened on any surface. The distal
articular surface is displaced farther out of line with the shaft of the humerus.
The grooves on the distal articular surface of the humerus are shallower, as
is the radial fossa. The notch formed by the depression immediately proximal
to the tubercle on the rim of the capitulum is deeper. The spinous process
of the medial epicondyle is shorter, extending only slightly beyond the distal
articular surface, The fossae at the lateral and medial ends of the distal
epiphysis are shallower.
Macrotus.—The humerus of this bat differs in many particulars from the
humeri of the other two bats and seems less specialized than either. As
seen laterally the distal part of the shaft curves gently forward; as viewed
posteriorly it has a barely noticeable S curve, with the proximal end of the
shaft curving mediad and the distal end laterad. The head is more nearly
round; its longest axis extends from anterior to posterior. The head extends
farther caudad from the posterior surface of the humerus. The pit for
the supraglenoid tuberosity is shallower and does not limit the movement of
the humerus until this element is extended well beyond an angle of 90 de-
grees to the lateral border of the scapula. The greater tuberosity extends only
slightly beyond the head, and the humerus does not lock until it is raised to
an angle of roughly 35 degrees with the plane of the scapula. The lesser
tuberosity has a thicker, more nearly knoblike end. Near its distal end the
ventral ridge bears a small thin tubercle. The shape of the pectoral ridge is
similar to that in Myotis, but the ridge is lower; the anterior edge is thinner
and does not help enclose the bicipital groove. The low lateral ridge angles
across the humerus from the posterior edge of the dorsal surface of the
greater tuberosity to a point immediately anterior to the middle of the dorsal
surface of the humerus opposite the middle of the pectoral ridge. The distal
part of the shaft is not flattened. As viewed anteriorly the distal articular
surface is displaced so far laterad that the medial rim of the trochlea is nearly
in line with the main axis of the shaft. There is no radial fossa. The ridges
and grooves on the distal articular surfaces are only weakly developed. There
is no depression at the medial end of the distal epiphysis. The medial epi-
condyle is large, irregularly anterioposteriorly flattened, and bears a short
spinous process that does not extend to the distal edge of the distal articular
surface.
Radius
Eumops.—tThe radius is long and thin and the proximal epiphysis is nearly
twice as thick as the shaft. As viewed anteriorly the proximal epiphysis is
triangular, and its most acute angle is directed proximad from the caudolateral
surface of the radius. The triangular articular surface is inclined strongly
FUNCTIONAL MORPHOLOGY OF THREE BATS 57
craniad and is deeply concave. It is marked by a central longitudinal groove
that is bordered by two low ridges and a lateral and medial depression. Into
the central groove fits the medial ridge of the capitulum of the humerus.
There is a shallow smooth-surfaced depression in the posterior surface of the
pointed proximal end of the radius; the proximal end of the ulna fits into
this depression, Immediately distal to the anteromedial part of the distal rim
of the articular surface is a deep slit that extends proximad and into which
the tendons of the Mm. triceps brachii and brachialis insert.
The shaft arches forward gently, the curvature being greatest in the
proximal half of the bone. Viewed anteriorly, the shaft curves slightly ventrad;
this helps give horizontal camber to the wing. The shaft is largely circular
in cross section and becomes slightly narrower distally; the distal half is
flattened on its dorsal and ventral surface. The anterior surface of the distal
three fifths of the radius is marked by a broad longitudinal depression in
which lie the large tendons of the Mm. extensor carpi radialis and brevis. A
second and narrower groove angles across the distal half of the radius from
the posterolateral to the anterior surface. Along this groove passes the tendon
of the M. abductor pollicus longus.
The distal epiphysis is slightly narrower than the proximal epiphysis. The
articular surface of the distal epiphysis is deeply concave and is bordered an-
teriorly by two pointed processes; the lateral one probably represents the
styloid process and the medial one an accessory process (pseudostyloid proc-
ess). At the posteromedial rim of the articular surface is a broad, prominent
process, and from the posterolateral rim a high, thin ridge extends proximad
roughly 3 mm. from the articular surface. There is an elongate hole in the
middle of the ridge and the distal end of the ridge forms a posterolateral con-
tinuation of the articular surface. This ridge provides a barrier between the
tendons of the flexor and extensor muscles. On the anterior surface of the
distal epiphysis, between and immediately proximal to the styloid and pseudo-
styloid processes, is a short broad ridge with concave sides and a weakly
concave anterior surface. In the deep groove between this ridge and the
pseudostyloid process lies the tendon of the M. abductor pollicus longus;
over the anterior surface of the ridge passes the tendon of the M. extensor
carpi radialis longus; through the groove between the ridge and a small
tubercle on the anterodorsal surface of the epiphysis extends the tendon of
the M. extensor carpi radialis brevis.
Myotis.—The proximal articular surface is broader and less clearly tri-
angular, is inclined less strongly craniad, and is not so deeply concave. The
slit into which the tendons of the M. biceps brachii and the M. brachialis
insert is not as completely closed; seen ventrally the slit is a large irregularly
triangular depression that is deepest proximally and is bordered by an anterior
and a posterior ridge. The shaft is slightly thinner and is mostly round in
cross section, but becomes progressively more anterioposteriorly compressed
distally and is not flattened on its dorsal or ventral surface. The distal
epiphysis is elongate dorsoventrally. The distal articular surface is bounded
anteriorly by two low knobs and posteriorly by a broad low process. The
ridge on the dorsal surface of the distal epiphysis is slightly larger and is not
perforated. A small tubercle arises from the anteroventral rim of the articular
surface. On the anterior surface of the radius are three ridges; the small
58 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
central ridge is bordered on each side first by a depression and then by
a low ridge.
Macrotus.—The proximal epiphysis is broad and is inclined craniad to
roughly the same degree as it is in Eumops. The depression in which the
M. biceps brachii and the M. brachialis insert is not enclosed medially. The
entire shaft is anteroposteriorly compressed and is flattened on neither the
lateral nor medial surface. The distal epiphysis is elongate laterally and its
articular surface is directed slightly caudad. The styloid process and anterior
pseudostyloid process are represented by two small tubercles immediately in-
side the rim of the distal articular surface; the posterior pseudostyloid process
is more strongly developed. In addition there is a well-developed process at
the posteromedial rim of the articular surface. No ridge is present on the
lateral surface of the distal epiphysis, but a stout process, which is in the same
position as the distal end of the ridge in the other genera, forms a postero-
lateral extension of the articular surface. Two short, low ridges arise from
the anteromedial rim of the distal articular surface. There are three ridges
on the anterior surface of the distal epiphysis; they are similar to those in
Myotis but are medial to the center of the anterior surface of the epiphysis.
Ulna
In Eumops the part of the ulna that lies against the proximal epiphysis of
the radius is robust; the shaft of the ulna is thin, becomes threadlike distally,
and terminates opposite roughly the middle of the radius. The body of the
proximal part of the ulna is irregularly fusiform; the proximal end is truncate
and the anterior surface is convex and smooth. The smooth surface fits in the
shallow depression on the posterior surface of the proximal tip of the radius
and extends slightly proximal to it. This small proximal extension of the ulna
represents the olecranon process and on it inserts the M. triceps brachii. From
the fusiform body of the ulna projects a broad, square-ended medial articular
process. It is convex posteriorly and concave and smooth anteriorly and ex-
tends medial to the proximal tip of the radius where the medial articular process
contacts the medial ridge of the trochlea.
Compared to Eumops the proximal part of the ulna is more symmetrical
and less angular in Myotis. The articular surface of the body of the ulna is
broader and slightly concave, and the medial articular process is smaller and
does not extend as far mediad. The thin shaft ends within the proximal third
of the forearm.
In Macrotus the articulation with the radius is less steady than in the other
genera because the articulating surfaces are flat. The medial articular process
is small and projects only slightly medial to the body of the ulna. The shaft
is thin and fuses with the posterior surface of the radius approximately two-
fifths of the way along the forearm.
Manus
A detailed description of each element of the manus is beyond the scope
of this report. Nonetheless, a description of some of the more striking special-
izations and a discussion of their functional significance is worthwhile. A
general idea of the carpi of the bats under consideration may be gained from
figure 7.
FUNCTIONAL MORPHOLOGY OF THREE BATS 59
Seemingly all the unique modifications evident in the bat’s carpus serve
the end of limiting movement of the wrist and digits to one plane. One of the
basic prerequisites for efficient flight is the development of flight surfaces
that can be held rigid against the force of the air stream. One way is which
this rigidity is attained in bats is by limiting movement at the elbow, wrist and
carpometacarpal joints (except the first) to the anteroposterior plane. The
manus of bats has been rotated (supinated) 90 degrees from the position of
the carpus in cursorial mammals; the digits are arranged one behind the other
in the anteroposterior plane with the thumb foremost. When the digits are
fully flexed they lie together next to the posterior surface of the radius. When
the wing is completely extended the second and third digits project straight
to the side and are nearly parallel with the distal part of the shaft of the
radius; the fourth digit extends caudad at an angle of roughly 15 degrees to
the third digit in Eumops. This angle varies considerably from one kind of
bat to another; the angle is approximately 20 degrees in Myotis and 35 degrees
in Macrotus. The fifth digit projects almost directly posteriad at an angle of
roughly 90 degrees to the third digit.
The following descriptions are based on Eumops and indicate the types of
specializations evident in the carpus. The lunar and cuneiform fit against
each other and their rounded proximal surfaces fit into the deeply concave
and laterally elongate distal articular surface of the radius. In the dorsal part
of the anterior surface of the lunar is a broad groove along which the styloid
process of the radius slides and a depression into which the process locks
when the manus is fully extended. In the ventral part of the articular sur-
face of the lunar is a second groove; this groove accommodates the anterior
pseudostyloid process of the radius. In the posteroventral surface of the
lunar is a broad groove bordered medially by a ridge. Along this groove
moves the posterior pseudostyloid process of the radius. This system of
tongue-in-groove articular surfaces limits movement at the radiocarpal joint
to the anteroposterior plane, or when the wing is outstretched, to the horizontal
plane.
The ventral surface of the carpus is spanned and reinforced by the pisi-
form. The bone is large and irregular in shape; its proximal end is attached
by fascia to the ventral surface of the trapezium and its distal end is bound to
the proximal part of the ventral surface of the fifth digit.
The proximal ends of all metacarpals except the first are modified so as
to rest tightly against each other and brace each other. As additional
strengthening the distal parts of some of the carpals are narrow and pointed
and fit tightly between or against the proximal ends of the metacarpals. The
dorsal part of the trapezoid is laterally compressed and pointed distally; it
rests in a deep groove in the dorsal surface of the base of the second meta-
carpal and locks into a depression at the distal end of the groove when the
metacarpal is fully extended. The dorsal part of the magnum is flattened on
its posterior surface and lies closely against the flat proximal end of the third
metacarpal; the distal edge of the ventral part of the magnum is narrow and
fits in a space between the proximal ends of metacarpals two and three.
Distally the unciform is divided into two lobes; the anterior lobe is thin and
extends between the bases of metacarpals three and four. The flattened
proximal end of metacarpal four extends into the depression between the two
lobes of the unciform. The second and posterior lobe is bluntly pointed and
60 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
fits into an anteroposterior groove in the proximal end of the fifth metacarpal.
In Myotis and Macrotus the carpus is not so specialized as in Eumops, but
in general the modifications are the same. The lunar is grooved, but the
processes of the radius are not so well developed and the joint seems to be
less strongly braced. The carpometacarpal joints are complex and although
they differ in many morphological details from those in Eumops they achieve
the same functional result.
The proximal ends of all metacarpals except the first in all three genera
are highly specialized for strong articulations with the carpals and for move-
ment in only the anteroposterior plane (Fig. 7). The digits are greatly
elongated and are curved ventrad slightly giving the distal part of the wing
horizontal camber. The metacarpals are the longest segments and the
phalanges become shorter distally (except for the third digit in Macrotus in
which the first phalanx is shorter than the second). The phalangeal formulae
of the genera are as follows (the superscript “c” means that the distalmost
phalanx is cartilaginous): Eumops and Myotis, 2-1-3°-3°-3°; Macrotus,
2-1-3-3°-8°.
In all three genera the thumb is allowed free movement and is the only
clawed digit. The phalanges are slightly laterally compressed and the claw
strongly so. In all three genera the second digit is nearly round in cross
section and terminates opposite the third metacarpophalangeal joint. In
Eumops the end of the second digit is bound by connective tissue to the third
metacarpophalangeal joint; the dactylopatagium minus is narrow and ends at
this point. In Myotis and Macrotus the dactylopatagium minus is broad and
ends at the joint between the first and second phalanges of the third digit.
The third metacarpal is nearly round in cross section in Myotis, slightly
dorsoventrally compressed in Macrotus, and considerably compressed in
Eumops. The phalanges of this digit are round in Myotis and Macrotus; in
Eumops the first phalanx is compressed and the second is round. The fourth
digit is round in the three genera. The third and fourth digits of Eumops are
remarkable for the direction of flexion of their phalanges. In Myotis and
Macrotus these digits flex ventrally. In Eumops, in contrast, the first
phalanges of the third and fourth digits flex posteriorly, and the second
phalanges flex anteriorly. Due to this arrangement, when these digits are
flexed the first phalanges lie along the distal parts of the posterior surfaces of
the metacarpals and the second phalanges rest next to the anterior surfaces
of the first phalanges. This accordionlike folding of the phalanges allows the
long distal part of the wing to be folded into a bundle that does not project
beyond the distal end of the third metacarpal. In all three genera the
phalanges of the fifth digit flex ventrally, but are not allowed to flex as far
as those of the other digits. This movement is most restricted in Eumops.
The length of the fifth digit varies considerably; it is slightly shorter than the
third metacarpal in Eumops, approximately 1.4 times as long in Myotis, and
1.6 times as long in Macrotus. The fifth metacarpal is interesting because of
the specializations that strengthen the shaft of the metacarpal and reduce
dorsoventral bending. Among the bats studied, the fifth metacarpal is most
highly specialized in Eumops. In this genus the shaft is curved ventrad more
abruptly than are the shafts of the other metacarpals. The proximal two
thirds of the shaft is strongly laterally compressed and is broadest (dorso-
ventrally) at roughly the end of the first quarter; the greatest breadth of the
FUNCTIONAL MORPHOLOGY OF THREE Bats 61
shaft is approximately 4.3 per cent of its length. As seen in cross section,
the proximal two thirds of the shaft has a large, rounded dorsal part that
blends into the central biconcave section; the ventral part is expanded
slightly and is rounded ventrally (Fig. 7). In Myotis and Macrotus the shaft
of the fifth metacarpal is laterally compressed, elliptical in cross section, and
not nearly so robust as in Eumops (greatest breadth of shaft roughly 1.7 per
cent its length in Myotis, 1.6 per cent its length in Macrotus).
Pelvic Girdle and Limb
Innominate Bone
Eumops.—This bone is heavily built and has strongly developed ridges and
tuberosities. The pelvis is set on the sacrum at an angle; the dorsalmost part
of the iliac crest is dorsal to the level of the neural spine of the first sacral
vertebrae, whereas the center of the acetabulum is ventral to the level of the
ventral surface of the centrum of this vertebra.
The ilium is tilted so that the gluteal fossa faces dorsad whereas the part
of the ilium that is the dorsal rim in most mammals is directed mediad and
contacts the sacrum. The sacroiliac joint involves that part of the thick medial
portion of the ilium between the iliac crest and a point 2 mm. anterior to the
acetabulum and the expanded lateral masses of the sacral vertebrae one to
three. In adult individuals the ilium and the sacrum are solidly fused and the
line of junction is difficult to trace. The anterior third of the ilium broadens
abruptly and merges anteriorly with the broad, thick iliac crest. As seen
anteriorly the ilium is roughly triangular; the most acute angle is represented
by the iliac ridge that points laterad. The thickest part of the ilium is the
medial part. The surface of the broad gluteal fossa is deeply concave; the
ventral surface of the ilium faces ventrad and slightly laterad and is almost flat.
The acetabulum is large, faces dorsolaterad and somewhat caudad, and is
situated immediately posterior to the middle of the innominate. Attached
to the anterior cartilaginous rim of the acetabulum is a sesamoid bone from
which the M. rectus femoris takes origin. The acetabulum is open posteroven-
trally.
The dorsal rim of the ischium is short and robust and at its posterior end
the thick, rounded dorsal ischial tuberosity projects dorsad. The ascending
ramus of the ischium is flat and moderately broad and is arched outward.
The rami converge ventrally and from immediately anterior to the ventral
ischial tuberosities bars of bone extend mediad and form a symphysis. I have
available no skeletons of the bats here reported on that retain traces of sutures
between the three bones that comprise the innominate bone. Probably the
part of the innominate that surrounds the obturator fenestra and extends from
the ventral rim of the acetabulum to the symphysis is the pubis; for descriptive
purposes it will be so regarded. From the symphysis the pubis extends cranio-
laterad; the tip of the large pubic spine extends well anterior and slightly
lateral to the acetabulum. The dorsal ramus of the pubis is broad and thick
and from the ventral rim of the acetabulum is directed ventrolaterad. Probably
due to the reptilian posture of the hind limbs in this bat, the pubes flare so
sharply laterad as they extend forward from the symphysis that the width of
the post-acetabular part of the pelvis as measured between the centers of
the ventral borders is greater than the width between the dorsal borders.
This is the reverse of the usual proportions in cursorial mammals.
62 UNIVERSITY OF Kansas Pus3s., Mus. Nat. Hist.
Myotis.—The pelvic girdle of this genus is more lightly built and is not at-
tached to the sacrum at so great an angle. The sacroiliac joint is not fused
and involves only the first two sacral vertebrae. The lateral border of the
ilium is straight and nearly parallel to the midline, or anteriorly converges
slightly toward the midline in some specimens; the anterior end of the ilium is
truncate and nearly round in cross section. The posterior two thirds of the
ilium is roughly elliptical in cross section (the long axis is the lateral axis)
and there is neither a gluteal nor an iliac fossa. There is a small tubercle
immediately anterior to the acetabulum on which the M. rectus femoris
originates. The dorsal ischial tuberosity projects more nearly caudad, and
the ascending ramus angles more markedly caudad from the tuberosity. The
ventral ischial tuberosity is larger. The symphysis is broader; the pubes ex-
tend less abruptly laterad and at a greater angle dorsad so that the post-
acetabular part of the pelvis is not nearly so broad ventrally. The pubic
spine is shorter and, in relation to the acetabulum, does not extend so far
craniad or laterad. The dorsal ramus of the pubis is shorter and broader.
Macrotus.—In general the pelvis is more lightly built than those of the
other two bats and is unusually small. The length of the pelvis is nearly the
same in Macrotus californicus and Myotis velifer although the weight of the
former bat is nearly twice that of the latter. The sacroiliac joint is not
ankylosed and involves the first two sacral vertebrae. The ilium is rotated
so far that the gluteal fossa faces dorsomediad and the iliac ridge points
dorsolaterad. The anterior quarter of the ilium is expanded and is broader
and thicker than the rest of the bone. There is a moderately well-developed
gluteal fossa and a faint iliac fossa. A large sesamoid bone that gives origin
to the M. rectus femoris is attached to the anterodorsal part of the cartilagi-
nous rim of the acetabulum. The pelvis is narrow as measured across the bodies
of the ischia; the ischia lie close to the lateral masses of the last sacral vertebrae
and are connected to them by ligaments. The ascending ramus of the ischium
is lightly built. The symphysis is narrow and the pubes are less widely
spread apart than in Eumops, but, because the bodies of the ischia are so
close together, the post-acetabular part of the pelvis is broader ventrally than
dorsally. The pubic spine is extremely long, extending craniad beyond the
level of the middle of the ilium. The dorsal rami of the pubes are longer
than in Myotis and extend ventrolaterad from the ventral rim of the acetab-
ulum.
Femur
Compared with the femora of most mammals those of bats are thin and
long; also, they are long relative to the length of the shank.
Eumops.—tThe diameter of the shaft is approximately 4.7 per cent the length
of the femur, and the femur is slightly longer than the shank. In cursorial
mammals the head of the femur projects mediad from the proximal end of
the bone; in this bat, however, the head angles craniad and only slightly
mediad, being offset to the extent that the posterior surface of the head is
approximately in line with the long axis of the shaft. The articular surface
of the head extends around almost the entire head and the head projects well
beyond the trochanters. These features suggest that a wide range of move-
ments is possible at the hip joint.
FUNCTIONAL MORPHOLOGY OF THREE BATS 63
The greater trochanter is broad, rounded distally, and extends proximolaterad
from the short neck of the femur to a point opposite the middle of the head.
The end of the greater trochanter bears two small marginal tubercles. A low,
thin lateral ridge extends from the posterolateral surface of the greater tro-
chanter onto the proximal part of the shaft. The lesser trochanter is slightly
lower and narrower than the greater trochanter and gives rise distally to a low,
broad medial ridge on which insert certain adductor muscles. The lesser tro-
chanter projects proximomediad to roughly the level of the base of the head.
The shaft is nearly straight, but curves gently craniad and laterad. Roughly
three fifths of the way along the lateral surface of the shaft is a short ridge to
which the M. gluteus maximus and M. caudofemoralis attach.
The distal epiphysis is narrow (breadth of distal epiphysis approximately
2.3 times that of shaft) and extends posterior to the shaft a distance slightly
less than the diameter of the shaft. The patellar groove is broader than either
the lateral or medial condyle and the intercondylar fossa is deep. The center
of the medial surface of the medial condyle is marked by a shallow pit, and
the proximal part has a low prominence. The lateral condyle has at the
center of its lateral surface a shallow depression. The articular surfaces extend
around the posterior surfaces of the condyles to the point at which the condyles
meet the posterior surface of the shaft. This placement of the articular sur-
faces suggests that the shank is normally held in a flexed position.
Myotis.—The femur is slightly less robust (diameter of shaft approximately
4.3 per cent length of femur) and is shorter than the shank. Both trochanters
extend to a point roughly opposite the middle of the head. The lesser tro-
chanter merges distally with a moderately high medial ridge; the lateral ridge
is represented by an elongate prominence at the distal base of the greater
trochanter. The shaft is straight. The distal epiphysis is slightly broader
(breadth of distal epiphysis approximately 2.7 times that of shaft), is cleft
by a deeper intercondyloid fossa, and extends posterior to the shaft of the
femur a distance greater than its diameter. The patellar groove is less clearly
defined and, relative to the condyles, is narrower.
Macrotus.—The femur is longer than the shank and is slenderer than in the
other two genera (breadth of shaft approximately 3.6 per cent length of femur).
The head extends more nearly proximad than in the other genera. The head
in Macrotus lies slightly anterior and lateral to the long axis of the shaft of the
femur. The trochanters are small and knoblike. The lesser trochanter extends
proximad and posteromediad immediately beyond the middle of the head. The
greater trochanter projects laterad and does not extend beyond the base of
the head. The medial ridge of the femur is thin and high and arises from the
medial surface of the proximal fifth of the femur, starting immediately distal
to the medial base of the lesser trochanter. The lateral ridge extends for
roughly 1 mm. along the lateral surface of the second fifth of the femur. Im-
mediately beyond the level of the trochanters the shaft turns abruptly laterad
and then slightly mediad throughout the rest of its length. The proximal
epiphysis, therefore, makes an angle of roughly 160 degrees with the distal
four fifths of the shaft. The distal epiphysis is narrow (breadth of distal
epiphysis approximately twice that of shaft) and extends only slightly posterior
to the distal part of the shaft. There is a prominent tubercle on the lateral
surface of the lateral condyle.
64 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Tibia and Fibula
Eumops.—tThe tibia is heavily built and is shorter than the femur. The
proximal epiphysis is broad and extends posterior to the shaft. The proximal
articular surface is at right angles to the long axis of the proximal] half of the
shaft and is bilobed; each lobe is marked by a broad, shallow, anteroposteriorly
aligned groove that articulates with a femoral condyle. The lateral condyle
of the tibia is larger than the medial condyle and extends farther beyond the
corresponding adjacent surface of the shaft. The proximal quarter of the
shaft is roughly circular in cross section. At the end of the proximal eighth of
the posterior surface of the shaft the heavy tendon of the M. semimembranosus
inserts on a broad, elongate, rough-surfaced tubercle. A broad ridge ap-
proximately 1.5 mm. long arises roughly one third of the way along the posterior
surface of the femur. The tendon of insertion used in common by the Mm.
gracilis and semitendinosus attaches on the rough top of the ridge. This
ridge gives way distally to a lower, thinner ridge that extends slightly beyond
the middle of the posterior surface of the shaft. The distal two thirds of the
shaft is laterally compressed, roughly elliptical in cross section, and curved
caudad. The anterior edge of the distal half of the shaft bears a narrow
crest that reaches its greatest height three quarters of the way along the
tibia and gives origin to part of the M. tensor plagiopatagii. The distal
epiphysis is expanded and is elongate anteroposteriorly. The distal end and
lateral surface of the medial malleolus articulate with the medial and ventral
surfaces of the astragalus. The lateral part of the distal articular surface faces
distad and articulates with the dorsomedial part of the proximal half of the
astragalus. The tendon of the M. plantaris passes along a depression in the
posteromedial surface of the distal epiphysis of the tibia.
The fibula is slender but complete in Eumops. As viewed posteriorly the
bone forms a gentle S-shaped curve; the proximal part of the shaft curves
mediad and the distal part laterad. The medial surface of the head of the
fibula contacts the lateral surface of the lateral condyle of the fibula. The
proximal end of the fibula rests against a small bone that articulates distally
both with the fibula and the lateral condyle of the tibia and articulates
medially with the lateral condyle of the femur. The distal epiphysis is large
and its broad lateral malleolus articulates with the dorsolateral part of the
proximal end of the astragalus. The posteromedial part of the distal articular
surface articulates with the ventral part of the proximal end of the astragalus.
Myotis.—The tibia is more slender and longer than the femur. The lateral
condyle is extended caudolaterad as a broad, bluntly pointed projection that
serves to increase the area of the articular surface. Only a small elongate
prominence on the posterior surface of the tibia immediately distal to the
proximal epiphysis marks the point of insertion of the hamstring muscles.
The shaft of the tibia is nearly round in cross section, becomes progressively
narrower distally, and curves slightly mediad. The medial malleolus is short
and narrow.
In this genus the fibula is thin and incomplete; it extends from the proximal
end of the carpus to within approximately 2 mm. of the lateral projection of
the lateral condyle. The distal end of the fibula is expanded and contacts the
dorsolateral part of the proximal end of the astragalus.
FUNCTIONAL MorRPHOLOGY OF THREE Bats 65
Macrotus.—The tibia is slightly shorter than the femur and is considerably
thinner than in the other two genera. The lateral condyle is extended laterad
as a pointed process that turns sharply distad at its tip. A short ridge extends
onto the shaft from the medial condyle. A broad, rough tubercle, which
serves as the point of attachment for the hamstring muscles, arises from the
posterior surface of the tibia immediately distal to the proximal epiphysis.
Distal to this point, the shaft is slightly laterally compressed and has low crests
along its anterior and posterior borders. The shaft becomes progressively
narrower distally and curves gently caudad. The distal epiphysis is ex-
panded, and inclined posteriad, and the articular surface faces distad and
slightly laterad.
The fibula in Macrotus extends along roughly the distal half of the shank
and is thinner than in the other genera. It becomes broader distally and
articulates with the dorsolateral part of the head of the astragalus.
Pes
In general the feet of these bats are unspecialized. They retain the primi-
tive mammalian phalangeal formula (2-3-3-3-3) and in all three genera the
foot is short, amounting to roughly 25 per cent of the total length of the hind
limb. Lateral as well as anteroposterior movement is possible at the ankle-
joint. The major specializations of the pes are as follows: elongation of the
astragalus and calcaneus; development of the calcar; lateral compression of
the phalanges and claws; elongation of the first phalanx of the first digit thus
making all the digits nearly equal in length.
Eumops.—The foot is broader and the individual bones are more massive
in this bat than in the other genera. The astragalus is irregular in shape
and is roughly two thirds as long as the calcaneus. The astragalus articulates
proximally with the tibia and fibula and distally with the navicular and cuboid.
The fibula articulates with the broad, rounded proximal surface of the astraga-
lus and the tibia contacts the middle of the dorsomedial surface. The distal
end of the astragalus is rounded and elongate dorsoventrally. The astragalus
is not in exact alignment with the plane of movement of the knee-joint, but
extends slightly mediad. Thus, when relaxed, the foot “toes” inward slightly.
The tendons of the M. tibialis posterior and the M. flexor digitorum fibularis
pass along the grooved ventral surface of the astragalus. Roughly one third
of the way along its length the dorsomedial surface of the calcaneus contacts
the ventrolateral surface of the proximal end of the astragalus. The dorso-
lateral surface of the distal end of the astragalus and the dorsomedial surface
of the distal end of the calcaneus are also in contact, and these bones are
bound together by connective tissue throughout most of their lengths. The
distal part of the calcaneus is laterally compressed; the distal articular surface
is deeply notched and fits against the cuboid. The partly ossified base of the
calcar articulates with a large concavity in the lateral surface of the proximal
half of the calcaneus. The calcar is circular in cross section, mostly cartilag-
inous, and approximately 27 mm. long. Its proximal end is pointed, laterally
flattened, and notched. A hollow in the posterior surface of the base of the
calear accommodates the rounded lateral part of the proximal end of the
calcaneus and allows the calcar to be drawn caudad, toward the posterior
surface of the shank, when the bat is crawling. The calcar is pulled laterad
38—4357
66 UnIvERSITY OF KAnsAS Pusts., Mus. Nat. Hist.
during flight by the M. depressor ossis styliformis and serves to spread the
uropatagium and reinforce its lateral edge. The medial tarsal (see Hill, 1937:
100) is large and crescent-shaped and lies at right angles to the long axis of
the tarsus. The medial end of this bone rests against the ventral surface of
the internal cuneiform. The thick, fleshy pad of the ventral surface of the
tarsus is attached to the medial tarsal; this bone probably serves to protect
the ventromedial part of the tarsus when the bat is crawling. The distal
surface of the medial tarsal gives origin to the M. abductor hallucis brevis.
The remainder of the tarsal bones present no unusual features. The base of
the fifth metatarsal is broad and dorsoventrally flattened; from its ventral
surface arises the M. abductor digiti quinti. In digits two to five, the segments,
arranged in order from longest to shortest, are: first phalanges, metatarsals,
second phalanges, third phalanges. In the first digit the first phalanx is the
longest element and the second phalanx, the shortest.
Myotis.—The foot of this bat is more lightly built, but the basic pattern
of articulation between the bones closely resembles that in Eumops. The
astragalus angles more sharply mediad and is rotated slightly laterad; hence,
when relaxed, the foot turns inward more sharply and is slightly supinated.
The astragalus and calcaneus are in closer contact with each other and less
movement seems to be possible between them. The flat, entirely cartilaginous
calcar is truncate proximally and articulates with the ventrolateral surface of
the proximal end of the calcaneus. The calcar is roughly 14.5 mm. long.
The medial tarsal is small. There is a round sesamoid on the ventral base of
the fifth metatarsal. The base of this bone is flattened and broad, but not
so much so as in Eumops. The segments of digits two to five in Myotis be-
come progressively shorter distally. The first phalanx is the longest segment
of the first digit and the second phalanx, the shortest.
Macrotus.—The astragalus and calcaneus are shorter relative to the lengths
of the digits than in the other genera, but the scheme of articulation is the
same, The tibia and fibula both articulate with the proximal end of the
astragalus and this bone extends almost directly distad in line with the tibia.
The round proximal articular surface faces proximodorsad and the truncate
distal articular surface, distad and slightly ventrad. The foot is slightly ex-
tended when relaxed, while in the other genera the foot remains partly flexed
when relaxed. In Macrotus the foot is slightly supinated when relaxed. The
cartilaginous calcar is oval in cross section, has a simple truncate base, is
roughly 10 mm. long, and articulates with the proximolateral surface of the
proximal end of the calcaneus. The dorsal part of the distal third of the
calcaneus is enlarged and has two grooves in its dorsal surface through which
pass the tendons of the Mm., peroneus longus and brevis. The tendons of the
Mm. tibialis posterior and flexor digitorum fibularis pass along the depression
between the medial surface of the proximoventral part of the calcaneus and the
ventral ridge on the proximal part of the astragalus. The medial tarsal is
small and lies on the ventral surface of the internal cuneiform. From the base
of the fifth metatarsal a fingerlike tubercle projects proximolaterad. The
relative lengths of the digits are as in Myotis.
FUNCTIONAL MorPHOLOGY OF THREE BATS 67
MYOLOGY
Introductory Remarks
In the arrangement of the muscles I have followed almost en-
tirely the system used by Hill (1937). Rinker’s arrangement (1954)
has been followed for muscles not mentioned by Hill. These
systems were employed primarily because they are convenient to
use in a study of functional morphology.
In connection with specific anatomical points, only in a few cases
are the conclusions of early workers given. My findings are oc-
casionally in disagreement with those of early authors, but it would
add little to discuss these points of difference. Errors made by
early workers may well have been due to the lack of adequate
optical equipment for use in dissecting.
Except with regard to the hand, descriptive terms are applied
to the bats considered here as they are to other mammals (see
page 44). In the following accounts the brief description given
under the headings “origin” and “insertion” are based on Eumops.
In instances where no description is given for a muscle in Myotis or
Macrotus the reader is to assume that the muscle does not differ
significantly from the corresponding muscle in Eumops. The in-
nervations of most of the muscles are given in parentheses next to
the names of the muscles. Where no innervation is given this
means that it was not determined.
Muscles Unique to Bats
M. occipito-pollicalis
Oricin.—From lambdoidal crest just lateral to midline.
INsERTION.—Along distal part of anterior surface of second metacarpal.
ReMarKS.—This muscle is similar in all three genera. The origin is fleshy
and the belly of the muscle passes over the anterior surface of the shoulder,
to which it is bound by connective tissue. Between the shoulder and thumb
the muscle gives way to a tendon that passes along the anterior edge of the
propatagium, ventral to the base of the first phalanx of the thumb, and along
the membrane on the leading edge of the wing to insert on the distal part of
the anterior edge of the second metacarpal. The manner in which the muscle
is bound to the shoulder varies. In Eumops a slip of the M. clavodeltoideus
is reflected outward and, instead of inserting on the pectoral ridge of the
humerus, ends in a short flat tendon intersecting the distal part of the belly
of the M. occipito-pollicalis. The slip is slightly more than 1 mm. wide, and
its distal part normally lies in a depression immediately proximal to the in-
sertion of the M. clavodeltoideus. Continuing from the distal end of the belly
68 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
of the M. occipito-pollicalis in Eumops is a heavy band of elastic cartilage
that becomes tendinous near the carpus. In Myotis a small strand of connective
tissue arises from the distal surface of the M. clavodeltoideus and intersects
the occipito-pollicalis. In Macrotus a flat tendon extends to the muscle from
a slight depression extending at right angles across the fibers of the pectoralis
muscle 2 mm. short of its insertion on the humerus. Seemingly the depression
is caused by the tautness of the band of fascia that gives rise to the tendon.
This muscle was given the name occipito-pollicalis by Kolenati (1857:9),
and the name was adopted by Macalister (1872:129). The phylogenetic
origin of the muscle was disputed by early workers. The muscle was found
by Macalister (loc. cit.) to be innervated by the spinal accessory nerve, and
for this and other reasons he concluded that the muscle was a derivative of
the occipital part of the trapezius. I was unable to trace the innervation of
the M. occipito-pollicalis. Because the distal part of the muscle continues
beyond the thumb to the second metacarpal, the name occipito-pollicalis is
not descriptively correct. Because of the long usage of the name, however, I
am retaining it.
ActTion.—The muscle increases the area and camber of the plagiopatagium
by pulling the propatagium craniad and ventrad. This action greatly im-
proves the effectiveness of the air foil of the wing. Photographs of Myotis
and Macrotus in flight demonstrate that in these genera, when the wing is
fully extended, the propatagium is stretched taut between the shoulder and
the base of the first phalanx of the thumb; the leading edge of the membrane
curves only slightly posteriad. In Eumops the propatagium narrows rapidly
toward the middle of the radius and even when fully spread the membrane
curves sharply toward the elbow. The breadth of the propatagium has an
important effect on the camber of the wing.
The placement of the origin of the tendon binding the belly of the M.
occipito-pollicalis to the front of the shoulder seems to be correlated with the
aerodynamic differences between the wings of the three genera. In Eumops,
which has a high speed, narrow wing, low camber is achieved by the high
placement of the origin of the binding tendon and the resulting shallow angle
of the propatagium. In Macrotus the placement of the origin of the binding
tendon is low—near the insertion of the M. pectoralis—and the propatagium
is pulled sharply downward; this helps to produce a high-lift wing of high
camber. Photographs of Macrotus in level flight show that when the wing is
extended the propatagium is drawn downward at an angle of roughly forty
degrees to the plagiopatagium. In Myotis the shoulder tendon originates on
the M. clavodeltoideus, but not so high as in Eumops.
M. coraco-cutaneus
Oricin.—From posterior surface of distal part of medial ridge of humerus.
INSERTION.—Into plagiopatagium.
ReEMARKS.—In Eumops this muscle is relatively larger than in the other
species. The long slender tendon of origin gives rise to a fusiform muscle
bundle roughly 9 mm. long that extends posteriad and laterad into the proximal
part of the plagiopatagium and terminates as a group of cords of elastic
cartilage that pass into the network of elastic strands reinforcing the posterior
FUNCTIONAL MORPHOLOGY OF THREE BATS 69
part of the plagiopatagium. In Myotis the origin is from the tip of the cora-
coid process. In Macrotus the muscle takes origin from the fascia on the dorsal
surface of the belly of the coracoid head of the biceps.
AcTIon.—This muscle helps reinforce the plagiopatagium by anchoring the
supporting network of elastic fibers in this membrane to the axilla.
M. humeropatagialis
Oricin.—From fascia over medial epicondyle of humerus and _postero-
medial surface of ulna, and by thin fascial sheet from medial surface of humerus.
INSERTION.—Into the plagiopatagium.
Remarxs.—Of the genera under study, this muscle is present only in
Eumops, in which it is cylindrical, approximately 20 mm. long and 1.5 mm.
wide, and extends distad and slightly caudad into the plagiopatagium. The
muscle was found in all the molossid bats that I examined. I have found no
reference in the literature to this muscle, and describe it here under the new
name Musculus humeropatagialis.
Action.—This muscle braces and tenses the anterodistal part of the plagio-
patagium.
M. depressor ossis styliformis
Oricin.—From dorsolateral surface of calcaneus and dorsal part of base of
fifth metatarsal.
INsERTION.—Along roughly 12 mm. of anterior surface of calcar.
REMARKS.—This muscle is approximately the same in all three genera, but is
somewhat shorter in Myotis and Macrotus (roughly 6 mm. long in each). The
name used here for this muscle was first proposed by Macalister (1872:159).
Action.—By pulling the calcar laterad this muscle spreads the uropatagium
and keeps it taut.
M. tensor plagiopatagii (tibial nerve)
Oricin.—In two parts. First part from distal three quarters of medial
surface and anterior crest of tibia and also from fascia on ventral surface of
tarsus (a sheet becoming progressively thicker distally; when relaxed, fibers
lie along medial surface of shank and extend proximad from their origin).
Second part on medial surface of internal cuneiform and base of first metatarsal
(band approximately 2.5 mm. wide at base and 10 mm. long) becoming
narrower distally.
INsERTION.—First part into that part of plagiopatagium attaching to anterior
surface of shank. Fibers extend approximately 6 mm. into membrane when
plagiopatagium fully spread. Second part extending along trailing edge of
plagiopatagium and merging with elastic connective tissue that reinforces
posterior edge of plagiopatagium.
RemMarks.—Among the bats under discussion here, this muscle occurs only
in Eumops, but in the Family Molossidae is present and varies but little in all
species that I have examined. I find no mention in the literature of the muscle
concerned and here propose for it the new name Musculus tensor plagiopatagii.
Action.—This muscle helps maintain the tautness of the posterior part of
the plagiopatagium when the wings are spread. An equally important function
70 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
may be to strengthen the connection between the plagiopatagium and the
lower leg, as the fast flight characteristic of many molossid bats demands strong,
well-braced flight membranes.
Muscles of the Pectoral Girdle and Limb
Trapezius Group
M. acromiotrapezius and clavotrapezius (spinal accessory nerve )
Oricin.—Along mid-dorsal line from level of last cervical to fifth thoracic
vertebra.
INsERTION.—On medial surface of distal end of clavicle, entire media] sur-
face of acromion process, and rim of scapular spine.
ReMarKs.—In all three genera the muscle becomes thicker anteriorly, and
the anteriormost part, which originates on the first thoracic vertebra, is more
or less separated from the rest of the muscle. The anteriormost part is prob-
ably a remnant of the clavotrapezius that has lost its cervical connections. In
Eumops the anteriormost part is only indistinctly separated from the rest of
the muscle, and the muscle fibers give way to a strong raphe, which is at-
tached to the knoblike neural spine of the first thoracic vertebra. From this
raphe large flattened tendons extend downward and laterad and attach to
the dorsal surfaces of the transverse processes of the first thoracic vertebra.
As a result an X is formed; the upper arms are the muscle fibers descending to
the raphe, and the lower arms are the tendons passing ventrad and laterad
to the first thoracic vertebra. Thus, an unusually strong origin for the anterior
division of this muscle is created.
In Myotis the anterior part takes fibrous origin on the dorsal surface of the
first thoracic vertebra. In Macrotus the origin closely resembles that in Eumops.
The insertion is on the medial surface of the acromion and medial edge of
the distal third of the clavicle, and the two parts of the muscle are more
nearly separated than in the other two genera.
Action.—This muscle pulls the clavicle and scapula mediad and tips the
vertebral border of the scapula downward. The remarkably strong origin of
the anterior part of this muscle and the large size of the entire muscle suggest
that it is important in steadying the clavicle and anterior part of the scapula,
against the pull of the powerful ventral flight musculature. Because of its
insertion on the prominent acromion process, this muscle is mechanically
well situated to aid in the upstroke of the wing by steadying the medial edge
of the scapula against the forces resulting from the contraction of the deltoid
muscles.
M. spinotrapezius (spinal accessory nerve)
OriciIn.—From the mid-dorsal line over thoracic vertbrae eight to thirteen.
INsERTION.—Along middle third of medial border of scapula adjacent to
intersection of spine and medial border.
ReMarks.—In Eumops all but the fleshy anterior part of the origin is
fibrous; the insertion is fibrous anteriorly becoming fleshy posteriorly. In the
other two genera the origins and insertions appear fleshy.
In Myotis the origin is from thoracic vertebrae seven to ten. In Macrotus
the muscle takes origin from thoracic vertebrae ten to thirteen and inserts along
FuNCTIONAL MorpPHOLOGY OF THREE BATS 6b
the medial border of the scapula from a point immediately anterior to the
junction of the spine and medial edge to a point slightly anterior to the pos-
terior end.
Action.—This muscle pulls the scapula posteromediad and tips the medial
edge ventrad, thus acting with the M. clavotrapezius and M. acromiotrapezius
to brace the scapula against the pull of the ventral flight muscles.
Costo-spino-scapular Group
M. levator scapulae (dorsal scapular nerve )
Oricin.—By four large slips, from transverse processes of cervical vertebrae
four to seven.
InseRTION.—Along vertebral border of scapula from posterior end of
anteromedial flange to junction of spine and vertebral border of scapula.
Remarxs.—In Myotis the origin is from the transverse processes of cervical
vertebrae three to six, by four slips. In Macrotus the origin is by three slips
from the transverse processes of cervical vertebrae four to six.
Action.—This muscle pulls the anteromedial border of the scapula forward
and ventrad; when the scapula is braced by other muscles, the M. levator
scapulae lifts the head and neck. While a bat is in flight the scapulae rock
back and forth on their long axes with each wing stroke. Intermittent con-
tractions of this muscle, associated with bracing the scapulae, would necessitate
reciprocal contractions of antagonistic muscles in order to steady the neck.
It seems likely that this bracing action, at least in steady flight, is not this
muscle’s major function. Also it is difficult to see how this muscle would exert
a steady enough pull to hold up the head and neck while the bat is in flight.
Perhaps this muscle functions mainly to brace or move the scapula in sudden
maneuvers demanding a departure from the standard wing stroke.
M. serratus anterior (dorsal scapular nerve )
Anterior Division
Oricin.—From broad band along middle sections of ribs one to four, by
four slips.
INSERTION.—On anteromedial surface and rim of anteromedial flange of
scapula.
RemMarks.—The origin in Myotis is by five slips that cover the distal end
of the first rib and a 3 mm. wide band along the distal surfaces of ribs two to
five. In Macrotus, the muscle originates on the first rib and costal cartilage.
Action.—This is the most effectively situated muscle to exert a direct
ventral pull on the anteromedial edge of the scapula. Working with the
trapezius group of muscles the anterior division of the M. serratus anterior
anchors the medial edge of the scapula and may help initiate the upstroke
of the wing.
Posterior Division (long thoracic nerve)
Oricin.—By eight heavy slips, from along distal surfaces of ribs one to
eight. Area of origin widens from roughly 2 mm. anteriorly to 10 mm. poste-
riorly.
INsERTION.—Along posterior two thirds of axillary border of scapula, along
adjacent lateral 3 mm. of M. subscapularis and lateral edge of M. infraspinatus
12 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
and on entire ventral surface and lateral half of dorsal surface of posterior
cartilaginous extension of scapula.
ReMaArKS.—The band of origin in Myotis extends from the short, broad
first costal cartilage, along the distal ends of ribs two to nine. The origin
broadens posteriorly, extending along roughly 5 mm. of costal cartilages eight
and nine. In Macrotus the origin extends from ribs two to 10, and is ap-
proximately 4 mm. wide posteriorly.
Action.—Second to the M. pectoralis the posterior division of the M.
serratus anterior is the largest muscle in the bats under consideration. It is
thick, divided into slips at its origin, and has a broad, fleshy insertion. In
most mammals the M. serratus anterior inserts on the vertebral border of the
scapula, but in bats the insertion is on the lateral border. Attending this
difference in insertion is a change in function. The M. serratus anterior in
cursorial mammals cradles the body between the scapulae and serves to bear
much of the weight of the anterior part of the body. This function is no
longer important to bats, as these animals use the forelimb primarily for flight
and seldom for quadrupedal locomotion. In bats the posterior division of
the serratus anterior is so situated as to be an efficient depressor of the axillary
border of the scapula; this action serves directly in the downstroke (power
stroke) of the wings.
It was mentioned in the description of the humerus that the greater tuberosity
functions as a locking device to stop the upward stroke of the humerus when
it has reached the angle to the scapula at which the greater tuberosity meets
the scapula. At this locking point the force of the stroke is transmitted to the
scapula, tending to rock the scapula by pulling the lateral border upward.
Contraction of the posterior division of the serratus anterior at this point in
the cycle would not only stop the tipping of the scapula but, by depressing the
lateral border and thus starting to push the locked humerus downward, would
initiate the downstroke without the help of the large adductors of the humerus.
The scapula rotates slightly about its anteroposterior axis, and the connection
between the clavicle and the scapula constitutes the single fairly rigid point
along this axis. The elastic effect of the tonus of the large serratus muscle is
probably sufficient to reduce greatly rocking of the scapula at the top of the
upstroke and thus the upstroke is terminated. By manipulating preserved
specimens of Eumops it was found that a full contraction of the posterior
division of the serratus anterior would swing the humerus downward through
an arc of at least twenty-five degrees. The labor of the downstroke, then, is
divided between the powerful adductors of the humerus—the M. pectoralis and
the M. subscapularis—and the posterior division of the M. serratus anterior.
The control of the downstroke is probably as follows: at the peak of the up-
stroke, when the humerus locks against the scapula, the lateral edge of the
scapula is anchored by the serratus anterior; the downstroke of the wing is
started by the contraction of this muscle with the resulting depression of the
lateral border of the scapula and the lowering of the humerus; the large
adductor muscles then take over and pull the humerus through most of its
downstroke. It should be stressed that the posterior division of the serratus
anterior may act to move the wing only when the humerus is in its locked
position. Perhaps the major importance of this muscle lies in its relieving the
adductors of the job of stopping the upstroke of the wings and beginning the
FuNCTIONAL MorPHOLOGY OF THREE BATS 73
downstroke, and of allowing the adductors to rest during the upstroke and
while the upstroke is stopped and the downstroke is begun. For a major flight
muscle this increase in the proportion of resting time to working time in the
wing-beat cycle may be extremely important when the wings are beat rapidly.
M. rhomboideus (dorsal scapular nerves )
Oricin.—From dorsal surfaces of thoracic vertebrae one to seven.
Insertion.—Along entire post-spinous part of medial border of scapula
including posterior cartilaginous extension.
Remarks.—The muscle takes origin in Myotis from thoracic vertebrae one
to five. In Macrotus the origin is from thoracic vertebrae one to six; the inser-
tion is relatively less extensive than in the other two genera because Macrotus
lacks a posterior cartilaginous extension on the scapula.
Action.—This muscle pulls the scapula mediad and tips the vertebral
border ventrad. Together with the trapezius muscles the M. rhomboideus
braces the scapula and helps control the upstroke of the wing.
M. omocervicalis (third cervical nerve )
Oricin.—From short, posteroventral spine at base of transverse process of
atlas.
INsERTION.—On tip of acromion process.
RemMarks.—In Macrotus the origin is from the ventral arch of the atlas and
the insertion on the anterior surface of the middle of the clavicle.
Action.—In most mammals this muscle draws the scapula forward and
mediad. In bats the scapula is powerfully braced by heavy muscles associated
with flight and the M. omocervicalis probably serves to draw the head and neck
caudad. In Macrotus the shift of the insertion to the clavicle may be associated
with this bat’s greater ability to move the head and neck.
Latissimus-subscapular Group
M. latissimus dorsi (subscapular nerves )
Oricin.—From mid-dorsal line over thoracic vertebrae ten to thirteen and
on lumbodorsal fascia to level of fourth lumbar vertebra.
INsERTION.—By short, heavy tendon shared with M. teres major, on distal
end of medial ridge of humerus
RemMarxs.—This muscle has a fleshy origin and tendinous insertion in all
three bats. The origin in Myotis is from thoracic vertebrae nine to eleven and
on the first two lumbar vertebrae; the insertion is on the distal end of the
medial ridge of the humerus slightly medial to the insertion of the M. teres
major. In Macrotus the origin is from thoracic vertebrae ten to twelve and
lumbar vertebrae one to four; the insertion is on the medial ridge of the
humerus just proximal to the insertion of the M. teres major.
ActTion.—This broad, long muscle is a flexor and rotator of the humerus
and can act upon the humerus nearly throughout its range of movement. Be-
cause of the rigid build of the elbow and wrist joints, allowing motion only
in the anteroposterior plane, contraction of the M. latissimus dorsi tends to
pronate the entire forelimb. The presence of the ventral ridge on the humerus
increases the effectiveness of the M. latissimus dorsi and M. teres major as
74 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
pronators by moving the insertions of these muscles away from the long axis
of the humerus. These muscles may act to control the rotational stability of
the humerus during the upstroke of the wing by acting as counter rotators
against the deltoideus muscles and the M. infraspinatus, and, in addition, in
terrestrial locomotion, help in the propulsion part of the stride of the forelimb.
M. teres major (subscapular nerves )
Oricin.—From lateral third of posterolateral facet of scapula.
INsERTION.—By common tendon with M. latissimus dorsi on distal end of
medial ridge of humerus.
REMARKS.—The entire posterolateral facet of the scapula is covered by the
origin of this muscle in Myotis; the insertion is by a fibrous sheet on the distal
end of the medial ridge of the humerus immediately lateral to the insertion of
the M. latissimus dorsi. In Macrotus the origin is over the posterolateral facet
of the scapula; the lateral fibers take origin from the dorsal surface of the
expanded rim of the scapula, and the medial fibers originate from the surface
of the M. infraspinatus. The insertion is by a broad flat tendon on the end of
the medial ridge of the humerus distal to the insertion of the M. latissimus
dorsi. The M. teres major in Macrotus is larger, relative to the latissimus dorsi,
than in the other two genera.
Action.—The function of the M. teres major — rotation and flexion of
the humerus—corresponds closely to that of the M. latissimus dorsi.
M. subscapularis (subscapular nerve )
Oricin.—From entire ventral surface of scapula, including posterolateral
(inner) surface of anteromedial flange and ventral surface of posterior carti-
laginous extension.
INSERTION.—On proximal part of lesser tuberosity of humerus.
ReMarks.—This is the third largest muscle in the bats under consideration,
being only slightly smaller than the posterior division of the M. serratus an-
terior. Macalister stated (1872:143) that “probably the largest subscapulars
in the animal kingdom are possessed by bats.” Seemingly the large antero-
medial flange of the scapula and the unusual faceting of the scapula are
modifications to increase the area of origin of the large subscapularis. The
muscle is composed of two thick, bipinnate parts, the fibers of each part in-
serting on a broad, central tendon. One tendon lies in the trough formed
by the posterolateral and intermediate facets, and the other, in the trough
formed by the anteromedial facet of the infraspinous fossa and the supra-
spinous fossa. The insertion is partly fleshy and partly by the extensions of
the two heavy, tendinous partitions. Proportionally, this muscle is largest in
Eumops and smallest in Macrotus. The lack of an anteromedial flange and
cartilaginous extension on the scapula limits the area of origin of the M.
subscapularis in Macrotus; otherwise, the attachments of this muscle seem
not to differ in the three genera.
Action.—This muscle adducts and extends the humerus. The attachments
of the muscle suggest that it has little mechanical advantage for power, but
can produce rapid action. Large size of the M. subscapularis, and the
osteological specializations associated with its origin, indicate that the muscle
is important. Its action is approximately that produced by the common con-
FUNCTIONAL MORPHOLOGY OF THREE BATS 75
traction of the anterior and posterior divisions of the M. pectoralis. The
principal function of the subscapularis is clearly to work with these muscles
and with the posterior division of the M. serratus anterior in producing the
downstroke of the wings. In addition, the subscapularis helps to support the
weight of the anterior part of the body in terrestrial locomotion by adducting
the humeri.
Deltoid Group
M. clavodeltoideus (axillary nerve)
Oricin.—From distal quarter of ventral surface of clavicle.
INSERTION.—On dorsal edge of proximal half of pectoral ridge of humerus.
Remarxks.—In Myotis the origin is from roughly the distal third of the
clavicle; the insertion is along the proximal two-thirds of the pectoral ridge
and the anterior surface of the base of the greater tuberosity of the humerus.
In Macrotus the insertion is along the dorsal edge of the proximal angle and
end of the pectoral ridge. The origin is roughly the same as in Myotis.
The M. clavodeltoideus is clearly separated from the anterior division of
the M. pectoralis only in Macrotus. Here the fibers of the muscles lie at
different angles, and the medial border of the M. clavodeltoideus overlies the
lateral border of the anterior division of the M. pectoralis. In the other two
genera these muscles appear more or less continuous with one another and
the exact division between them was not determined with certainty. A small
branch of the axillary nerve crosses the dorsal surface of the humerus, passes
between the proximal end of the pectoral ridge and anterior base of the greater
tuberosity, and penetrates the ventral surface of the M. clavodeltoideus. By
tracing this nerve into the muscle and finding where the adjacent M. pectoralis
began being innervated by the anterior thoracic nerve it was possible to gain
a rough idea of the relative extend of the M. clavodeltoideus and the anterior
division of the M. pectoralis.
Action.—This muscle is an extensor of the humerus and probably works
with the pectoralis muscles when the wing stroke is directed forward. The
M. clavodeltoideus also serves in quadrupedal locomotion to extend the
humerus at the start of the stride.
M. acromiodeltoideus (axillary nerve)
Oricin.—From lateral three-quarters of scapular spine and entire lateral
surface of acromion process of scapula.
INsERTION.—On dorsal surface of pectoral ridge of humerus.
Remarxs.—This muscle in Eumops has a fleshy origin and thick fleshy
insertion, In Myotis the fleshy origin is from the entire lateral surface of the
acromion; the fibrous insertion is on the distal half of the dorsal surface of
the pectoral ridge and along roughly one and one-half millimeters of the
dorsal surface of the humerus adjacent to the distal end of the pectoral ridge.
In Macrotus this muscle is divided into two parts; the first part takes origin
on the expanded anterior end of the acromion process and inserts on the dorsal
surface of the pectoral ridge; the second part originates along the entire
acromion process posterior to the tip, and inserts along roughly 2 mm. of the
dorsal surface of the humerus starting opposite the distal end of the pectorak
ridge.
76 UNIvERSITY OF Kansas Pusis., Mus. Nat. Hist.
Action.—This muscle elevates and rotates the humerus, and with the M.
spinodeltoideus controls the upstroke of the wing. It is difficult to estimate the
power requirements of the upstroke of the wing in bats, but some understanding
of the requirements can be gained by a comparison of the upstroke cycle of
the wing-beat in bats and birds.
In most birds—not in swifts and hummingbirds (Savile, 1950)—the up-
stroke requires little power and creates little thrust or resistance (drag) relative
to the down stroke. This economy of power during the upstroke is obtained
largely by the partial closing of the wings and the spreading apart of the
primary feathers to reduce air resistance, and by raising the wing with the
leading edge uppermost so that the air stream and muscles cooperate in its
elevation. Economy of power in the upstroke is achieved similarly in bats.
In bats also the wing is elevated partly closed and with the leading edge
uppermost. In contrast to birds, however, bats have continuous flight surfaces
and consequently are unable to “feather” the distal segment of the wing as
effectively as do birds. More drag is created, with a resultant demand for
more power. Accordingly, the elevating muscles in bats are fairly large.
Photographs of Macrotus in level flight show that the humerus moves
downward and forward during the downstroke, and is raised upward and
backward during the upstroke. This latter action can be produced by the
common action of the spinodeltoideus and acromiodeltoideus muscles, and it
is these muscles that supply much of the power needed for the upstroke.
The slight differences between the connections of the spinodeltoideus and
acromiodeltoideus muscles in the three genera of bats here considered may
have considerable functional importance. In Eumops the placement of the
origins of these muscles is such that they are posterior to the main axis of the
humerus throughout its normal range of activity; the insertions are both on
the dorsal surface of the pectoral ridge, anterior to the long axis of the humerus.
Their contraction, together or singly, would result in a fairly limited range of
movements, all acting to elevate and flex the humerus. Eumops forages in the
open, and its rapid, enduring flight probably calls for wing strokes directed
mainly in one plane, with relatively little need for a large variety of wing
actions. The restriction of the upstroke to an up and back direction probably
achieves an efficient concentration of power within this limited range of move-
ment. In Myotis, while the placement of the origin of the M. acromiodeltoideus
is roughly the same as in Eumops, the insertion is nearer the long axis of the
humerus. Thus, in Myotis, the humerus can probably be pulled nearly straight
upward. This bat forages near vegetation, and maneuverability, requiring
wing strokes in a variety of planes, is needed. The greater freedom of move-
ment possible in the upstroke may reflect this mode of flight. In Macrotus the
M. acromiodeltoideus and M. spinodeltoideus are each divided into two parts,
and each part has a separate origin and insertion. Seemingly these muscles
can control a greater range of actions in Macrotus than in either of the other
genera. Macrotus forages close to foliage or near the surface of the ground,
and extreme maneuverability is of primary importance; the wings must be
able to beat in many different planes at the expense of great efficiency in one
plane.
M. spinodeltoideus (axillary nerve)
Oricin.—From medial quarter of spine and entire post-spinous portion of
vertebral border of scapula including posterior cartilaginous extension.
FUNCTIONAL MORPHOLOGY OF THREE BATS ViTp
INSERTION.—The muscle fibers aggregate distally and emerge with those of
M. acromiodeltoideus, inserting on posterior border of M. acromiodeltoideus
and on dorsal surface of pectoral ridge of humerus.
Remarks.—In Myotis the origin is from the medial three-quarters of the
top of the ligamentous spine of the scapula and the post-spinous part of the
vertebral border of the scapula to the base of the posterior cartilaginous ex-
tension; the fibrous insertion is on the small knob in the middle of the lateral
ridge of the humerus. In Macrotus the muscle is divided into two parts. The
first part takes origin from the post-spinous part of the vertebral border and
the posterior two thirds of the ligamentous extension of the spine of the scapula;
the fibrous insertion is along roughly 2 mm. of the lateral ridge of the humerus
opposite the middle of the pectoral ridge. The origin of the second part is
from the anterior half of the ligamentous extension of the spine and the
posterior half of the acromion process of the scapula; the fibrous insertion is
on the lateral surface of the humerus at the distal base of the greater tuberosity.
The posterior edge of the second part of the muscle is overlapped by the
anterior edge of the first.
Action.—This muscle abducts (elevates) and flexes the humerus, and
probably supplies most of the power, together with the M. acromiodeltoideus,
for the upstroke of the wing. The spinodeltoideus is a strong rotator of the
humerus in Eumops, and a less effective rotator in Myotis and Macrotus.
Rotational stability of the humerus during the upstroke is probably controlled
by the deltoideus muscles’ working against the counter-rotational action of the
Mm. latissimus dorsi and teres major.
This muscle is discussed more fully under the account of the M. acromio-
deltoideus.
M. teres minor (axillary nerve )
Oricin.—From dorsolateral edge of scapula just posterior to glenoid fossa
and from above middle of origin of long head of M. triceps brachii.
INSERTION.—On distal surface of greater tuberosity of humerus.
ReMarks.—This muscle is composed of a single, short, flat band of fibers.
In Myotis it is especially thin and delicate. Otherwise, the muscle is similar
in all three genera.
AcTIon.—The muscle is a weak flexor and rotator of the humerus.
Suprascapular Group
M. supraspinatus (suprascapular nerve )
Oricin.—From entire supra-spinous fossa and medial surface of spine of
scapula and from thick ligament that extends from anteromedial flange of
scapula to tip of acromion and base of coracoid processes.
INsERTION.—On proximal part of lateral surface of greater tuberosity of
humerus.
ReMarks.—The connections of this muscle are similar in all three genera.
Relative to the other muscles taking origin on the scapula, M. supraspinatus is
larger in Macrotus than in the other genera.
AcTion.—This muscle elevates, extends and rotates the humerus. This
action is not part of the pattern of the usual wing-beat cycle, but is probably
important in rapid turns and changes of level, or in alighting, when this action
78 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
helps raise the wings and draw them forward. When the humerus is at or
above the horizontal, the structure of the humeroscapular articulation stops the
humerus from being extended beyond a position in which it forms a right angle
with the body in Eumops; the humerus may move slightly farther forward in
Myotis; in Macrotus the greatest freedom is allowed, and the humerus may be
extended forward much farther. The M. supraspinatus, therefore, can move
the humerus forward to different degrees in the three genera. The far greater
freedom of movement of the humerus and the larger size of this muscle in
Macrotus seem to be associated with the remarkable maneuverability in flight
of this bat. When Eumops and Myotis are walking this muscle helps extend
the wing at the start of each stride, and in this action probably works together
with the M. clavodeltoideus.
M. infraspinatus (suprascapular nerve )
Oricin.—From lateral surface of spine of scapula and all of infraspinous
surface of scapula but lateral third of posterolateral facet.
INsERTION.—On distal part of lateral surface of greater tuberosity of humerus.
REMARKS.—In Eumops the muscle is composed of two bipinnate parts, the
fibers of each part inserting on a broad, central aponeurosis. In Myotis the
muscle is bipinnate, with a single aponeurosis; the muscle originates from all
but the posterolateral facet of the infraspinous surface of the scapula and from
the lateral surface of the spine and ligamentous extension of the spine of the
scapula. The fleshy and fibrous insertion extends from the lateral surface to
the distal base of the greater tuberosity of the humerus. In Macrotus the
muscle is bipinnate, and the origin is from all of the infraspinous surface of the
scapula except the rim of the posterolateral facet.
Action.—This muscle flexes, abducts, and rotates the humerus. It helps
in the upstroke of the wing and in maintaining rotational stability of the
humerus, its action being similar to that of the M. spinodeltoideus.
Triceps Group
M. triceps brachii, caput lateralis (radial nerve)
Oricin.—On posterolateral surface of humerus from level of distal base of
greater tuberosity to level of distal end of pectoral ridge, and from entire con-
cave, posterior surface of medial ridge and adjacent posterior surface of
humerus.
INSERTION.—On the proximal end of the olecranon process.
ReMarks.—This muscle is similar in all three genera.
M. triceps brachii, caput medialis (radial nerve)
OricIn.—From distal three-quarters of posterior surface of humerus.
INsERTION.—On proximal end of olecranon process deep to tendons of long
and lateral heads of triceps.
Remarks.—In Eumops and Myotis this division of the triceps is distinct
throughout its course from the other two divisions. In Myotis the muscle is
thin and takes origin along the distal third of the humerus. In Macrotus the
origin is on the medial ridge of the humerus just distal to the insertion of the
teres major. The insertion is on the proximal end of the olecranon in all
three genera.
FUNCTIONAL MORPHOLOGY OF THREE BATS 79
M. triceps brachii, caput longus (radial nerve)
Oricin.—From axillary border of scapula along first 4 mm. posterior to
glenoid fossa.
INSERTION.—On proximal end of olecranon process.
ReMARKS.—The attachments of this muscle are similar in all three genera.
REMARKS ON THE TrICEPs Grourp.—Compared to the other divisions of the
triceps, the medial head is small. The lateral head is large, indistinctly divided
into a posterolateral portion and a posterior part, and fills the depression
formed between the concave posterior surface of the medial ridge of the
humerus, the posterior rim of the lesser tuberosity, and the posterior base
of the head of the humerus. The long head is the largest division of the
triceps, and is divided into a superficial and a deep portion. The origin of
the long head is partly fleshy and partly by a fibrous sheet attaching to the
tubercle immediately posterior to the glenoid fossa of the scapula.
In Eumops the tendons of the lateral head and the long head of the triceps
converge distally and are bound together in a common sheath as they extend
along the distal half of the humerus. Relative to the size of the animal,
the medial head of the triceps is largest in Eumops. It is separate from the
other divisions of the muscle, and gives rise, just short of the insertion on the
ulna, to a large sesamoid bone in the depression on the posterior surface of
the humerus between the prominent spinous process of the medial epicondyle
and the capitulum. The sesamoid is attached to the ulna by a short, broad
tendon. The tendons of the long head and lateral head of the triceps run
through the deep groove on the posterior surface of this sesamoid bone.
In Myotis the medial head is distinct from the other divisions of the triceps,
and the tendon of the medial head enlarges into a thick cartilaginous pad over
the posterior surface of the trochlea. A small sesamoid bone at the distal end
of the pad is connected to the ulna by a short tendon. The tendons of the
long head and lateral head of the triceps lie on the surface of this cartilaginous
pad. In Macrotus the tendons of the three divisions of the triceps are bound
together distally, and give rise to a single thick, padlike tendon over the
posterior surface of the trochlea. The tendon connects directly to the proximal
end of the ulna, and contains no sesamoid bone. Macalister (1872:146) in-
correctly thought that in all bats a sesamoid bone occurs in the distal part of
the triceps tendon.
Action.—The triceps group extends the forearm. The short olecranon
process indicates that these muscles produce rapid but not powerful extension
of the forearm. To achieve the maximum effect of thrust and lift from the
wing-beat cycle, the M. triceps must extend the forearm rapidly at the start
of the downstroke; as the downstroke continues the antagonistic biceps and
triceps act to keep the wing rigidly outstretched. Toward the bottom of the
downstroke the forearm is partly flexed, and remains in this posture until the
start of the next downstroke. Because the long head of the triceps originates
on the scapula posterior to the glenoid fossa, extension of the humerus by the
major flight muscles during the downstroke tends to lengthen the distance
between the origin and insertion of this muscle, thereby compensating for
the shortening of the distance due to the extension of the forearm and proxi-
mal movement of the olecranon process. Thus, the long head of the triceps
can act more effectively when the wing is extended than can the medial and
80 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
lateral heads, and is more important in steadying the forearm against the
flexing action of the biceps than are the medial and lateral divisions. The
triceps group is important also in quick manuevers calling for rapid extension
of the wing.
Extensor Group of Forearm
M. extensor carpi radialis longus (radial nerve)
Oricin.—From low ridge extending from lateral epicondyle onto distal part
of shaft of humerus.
INSERTION.—On dorsal base of first metacarpal and anterodorsal base of
second metacarpal.
REMARKS.—In Myotis and Macrotus the fibrous origin is from the proximal
edge of the lateral epicondyle of the humerus. The insertion is the same in
all three genera. Relative to other forearm muscles, this is a large muscle.
It is covered by a thick, glistening sheet of fascia. The fibers extend roughly
one third of the way along the radius and give way to a heavy, flat tendon
that passes beneath the tendon of the M. abductor pollicis longus and along
a shallow groove in the anterior surface of the distal end of the radius. After
passing beneath the transverse ligament, the tendon divides, and the two
tendons extend beneath the tendon of the M. extensor pollicis brevis to their
insertions. In Eumops the tendons of the M. extensor carpi radialis longus
pass beneath the broad fibrous insertion of the M. extensor indicis.
Acrion.—This muscle acts directly to extend the first and second meta-
carpals and indirectly to extend the entire distal part of the wing. Function
is discussed under the M. extensor carpi radialis brevis.
M. extensor carpi radialis brevis (radial nerve)
Oricin.—From distal part of ridge of lateral epicondyle of humerus.
INsSERTION.—On anterodorsal surface of third metacarpal immediately dis-
tal to its base.
ReMarks.—In Eumops the fibrous origin is immediately distal to the origin
of the M. extensor carpi radialis longus, and contains a large sesamoid bone
in the depression immediately anterior to the lateral epicondylar ridge. The
sesamoid bone has a concave articular facet that lies against the knob on the
anterior edge of the proximal rim of the lateral epicondyle. The distal tendon
of the M. extensor carpi radialis brevis contains a sesamoid bone just short
of its insertion in all three genera. In Myotis the origin lacks a sesamoid bone,
and is on the proximal edge of the lateral epicondyle immediately distal to the
origin of the M. extensor carpi radialis longus. In Macrotus the origin of the
M. extensor carpi radialis brevis lacks a sesamoid bone, and is in common
with the smaller M. extensor carpi radialis longus. In all three genera these
are the largest muscles in the forearm. Each muscle has a strong fascial
covering, which becomes thick and glistening distally. The tendons are large
and flat, and lie in shallow grooves in the anterior surface of the distal part
of the radius.
Action.—This muscle acts directly to extend the third metacarpal, and
indirectly to extend the distal part of the wing. Judging from their sizes and
functions, this muscle and the M. extensor carpi radialis longus are the most
important flight muscles in the forearm. Together they control the extension
FUNCTIONAL MORPHOLOGY OF THREE BATS 81
of all the digits and the spreading of the distal half of the wing membrane.
In terms of power requirements, this is the most demanding function of the
forearm muscles. (In connection with the actions of these muscles, it should
be stressed again that flexion and extension of all but the first digit in bats
is in the anteroposterior plane; no other kind of movement of these digits can
occur except by “give” at the joints or bending of the bones.) Because all of
the digits but the first are connected by a membrane, the extension of the
second or third fingers, which form the leading edge of the wing, will cause
the extension of all the digits and the spreading of the distal part of the wing
membrane.
The job of extending the distal part of the wing and holding it against the
force of the air stream during each downstroke of a prolonged period of flight
seems beyond the ability of these slender muscles. It is probable, however,
that when the bat is in flight these muscles function as strong, non-elastic bands,
that extend the distal half of the wing with relatively little muscular effort
when the radius is extended. There seem to be two structural details that help
these muscles accomplish this. The elasticity of the muscles is greatly reduced,
both by their large tendons that extend to within roughly the proximal fourth
of the muscles, and by the strong fascial sheets investing most of the bellies
of the muscles. Because the origins of these muscles are proximal to the
center of the lateral epicondyle of the humerus, extension of the radius tends
to increase the distance between the origins and insertions of the radial ex-
tensors. The proximal displacement of the origins of these muscles, which is
caused by the full extension of the radius, is considerable, and amounts to
roughly 3 mm. in Eumops. Thus, assuming the muscles to be non-elastic, the
extension of the radius would cause extension of the distal part of the wing by
the pull directed from the proximally shifted origin, through the muscles, to
the first three metacarpals. Strong contraction of these muscles would increase
the speed of extension and rigidity of the distal part of the wing. A con-
traction of just sufficient strength to make the muscle react as a non-elastic
cord, however, would cause extension of the distal part of the wing when the
radius was extended. By this arrangement, then, part of the burden of ex-
tending the distal half of the wing is transferred to the extensor of the radius,
the M. triceps brachii, and the slender radial extensors of the forearm are
helped to perform their actions effectively and with a minimum of effort.
M. supinator (radial nerve)
Oricin.—From the depression in lateral epicondyle of humerus.
InsERTION.—Along first 10 mm. of anterolateral surface of radius.
ReMArRKS.—In all three genera this muscle originates by a large tendon that
contains a sesamoid bone. The relationships of this short, broad muscle are
similar in the three genera.
Action.—This muscle flexes the radius and braces the elbow joint.
M. extensor pollicis brevis (radial nerve)
Oricin.—From dorsal surface of proximal part of ulna and entire anter-
odorsal surface of slender shaft of ulna, and by fibers that attach to M. ab-
ductor pollicis longus and M. extensor indicis.
INSERTION.—On first metacarpophalangeal joint and on distal end of second
phalanx of first digit.
82 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
REMARKS.—In Eumops the slender tendon divides into three parts over the
carpus; the lateral two are bound to the sides of the first metacarpal, the middle
tendon extends to the dorsal surface of the second phalanx. In Myotis and
Macrotus the tendon is bound by fascia to the first metacarpophalangeal joint
and then continues to its insertion on the second phalanx.
AcTION.—Extension of the thumb.
M. abductor pollicis longus (radial nerve )
Oricin.—Along all but first 10 mm. of proximal half of interosseous surface
of radius.
INSERTION.—On scaphoid bone of carpus.
ReMarRks.—The attachments of this muscle are approximately the same
in all three genera. The tendon passes from the dorsal surface to the anterior
surface of the radius roughly two thirds of the way along this bone. In Eumops
a well-developed groove in the radius is present along the course of this
tendon. In the other two genera the groove is not so clearly defined.
Action.—This muscle braces the ventral base of the fifth metacarpal. In
most mammals the insertion is on the first metacarpal, and the muscle func-
tions as an abductor of the thumb. In the bats under consideration here, the
insertion is on the scaphoid, which lies on the anteroventral surface of the
carpus, and the function of the muscle is completely different from that in
most mammals.
When the bat is in flight, it is important aerodynamically that the fifth digit
and hind limb hold the plagiopatagium at the proper angle of attack to de-
velop lift. By reinforcing the fifth carpometacarpal joint, this muscle helps
keep the joint from “giving” during the downstroke, and aids the fifth meta-
carpal in maintaining the plagiopatagium at the optimal angle. The action
is exerted on the fifth metacarpal via the scaphoid and the pisiform bone. The
distal end of the pisiform bone is strongly bound by fascia to the proximal
part of the ventral surface of the shaft of the fifth metacarpal; the proximal
end of the pisiform rests against the ventral surface of the trapezium, and is
bound to the connective tissue of the ventral surface of the carpus. The distal
end of the pisiform is solidly fixed, but the proximal end may move slightly.
It is by anchoring the proximal end of the pisiform that the M. abductor pol-
licis longus steadies the fifth metacarpal. This muscle attaches to the
anterior edge of the scaphoid, and this in turn is attached to the proximal
end of the pisiform by a broad ligament. Accordingly, contraction of this
muscle tends to move the scaphoid craniad, this pull being transmitted by
the strong ligament to the proximal end of the pisiform. This bone is thus
kept from being displaced caudad when the articulation between the carpus
and fifth metacarpal is subjected to the strain caused by the force of the air
pressure on the flight membranes and the fifth digit. If free dorsoventral
movement were allowed at the articulation between the carpus and fifth meta-
carpal, the M. abductor pollicis longus would act as a ventral flexor of the
fifth digit; but, because the joint allows little dorsoventral movement, this
muscle serves instead to help the joint resist forces tending to cause dorsal
extension of the fifth digit.
FUNCTIONAL MORPHOLOGY OF THREE BATS 83
M. extensor digitorum communis (radial nerve)
Oricin.—By broad aponeurosis, from lateral epicondyle of humerus.
INsERTION.—By three tendons, on dorsal surfaces of distal ends of second
phalanges of digits three, four and five.
ReMarks.—In Myotis the muscle gives rise to two tendons; the first divides
into two parts over the carpus, one part inserting on the shaft of the first
phalanx of the second digit, and the other on the distal end of the second
phalanx of the third digit. The second tendon also divides; the large branch
inserts on the shaft of the second phalanx of the fourth digit, and the small
branch extends along the dorsal surface of the shaft of the fifth metacarpal
where the tendon appears to join the tendon of the M. extensor digiti quinti
proprius. The M. extensor digitorum communis has two partially separate
bellies in Macrotus. One has a fibrous origin from the lateral epicondyle of
the humerus and lateral surface of the proximal part of the ulna, and a fleshy
origin from the posterodorsal border of the proximal half of the radius; the
insertion is on the distal end of the third phalanx of the third digit. The
second and smaller belly takes fibrous origin from the lateral epicondyle, and
fleshy origin from the surface of the first belly; the tendon divides over the
carpus, one division inserting on the distal end of the second phalanx of the
fourth digit, and the other on the distal end of the second phalanx of the
fifth digit. In Macrotus an extremely small tendon extends along the proximal
part of the second metacarpal, inserting, in most specimens, at about the
middle of the dorsal surface of the metacarpal. I found this tendon difficult
to trace to its origin, but it appears to arise from the tendon of the division of
the M. extensor digitorum communis, which inserts on the third digit. In all
three genera the tendons of this muscle are bound by fascia to the digital
joints across which the tendons extend.
Action.—This muscle extends the phalanges of digits three to five in
Eumops and Macrotus, and the phalanges of digits two to five in Myotis,
serving in each genus to spread the tip of the wing membrane. This is an
important function, for the entire distal part of the wing is kept spread while
the bat is in flight. Seemingly, however, the action demands little power;
the muscle is long and slender and looks to be weak. Perhaps while the bat
is in flight the air stream helps keep the wing membrane spread.
This muscle retains a functional connection with the second digit only
in Myotis of the three bats considered. Because in the Microchiroptera the
second digit is bound distally to the third digit and can not be extended
separately, it would seem that an extensor of the phalanges of the second digit
is unnecessary, and its loss in Eumops, and the great reduction in Macrotus,
is what might be expected. Retention in microchiropteran bats of the part
of the M. extensor digitorum communis serving the second digit, as seen in
Myotis, is probably a primitive character.
M. extensor indicis (radial nerve )
Oricin.—Opposite middle half of forearm on posterodorsal surface of ulna
and interosseus surface of radius.
INsERTION.—Mostly on dorsal base of second metacarpal.
84 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
RemMarxs.—In Eumops the tendon spreads out over the dorsal surface of
the carpus into a broad, fan-shaped aponeurosis and inserts on the anterodorsal
surface of the carpus; the thickest part attaches to the dorsal base of the
second digit. The origin in Myotis is from the distal tip of the ulna and the
middle fifth of the interosseus surface of the radius; the insertion is by a
thick tendon on the dorsal surface of the second metacarpal about 3 mm. from
the base. In Macrotus the origin is along roughly the distal two thirds of the
posterior surface of the radius; the insertion is by a large tendon on the dorsal
base of the second metacarpal.
ActTion.—This muscle extends the second digit, thus helping the radial
extensors to spread the distal part of the wing and steady its leading edge.
M. extensor digiti quinti proprius (radial nerve )
Remarks.—I failed to find this muscle in Eumops. In Myotis the muscle
arises by fibrous origin from the posterodorsal base of the ulna and by fleshy
origin from the posterior edge of the M. extensor digitorum communis opposite
the proximal third of the forearm; the insertion is by tendon on the dorsal
surface of the shaft of the second phalanx of the fifth digit. This tendon is
joined, over the fifth metacarpal, by a small tendon from the M. extensor
digitorium communis. In Macrotus the M. extensor digiti quinti proprius takes
fibrous origin from the proximal 8 mm. of the dorsal surface of the ulna and
from the lateral epicondyle of the humerus; the insertion is on the dorsal
surface of the tip of the second phalanx of the fifth digit.
Action.—In Myotis and Macrotus this muscle extends the fifth digit.
M. extensor carpi ulnaris (radial nerve)
Oricin.—Along first 8 mm. of posterior surface of ulna, and on distal half
of posterodorsal surface of radius, by fleshy attachments.
INsERTION.—By large, broad tendon, on anteromedial base of third meta-
carpal.
Remarxks.—In Eumops the tendon extends proximad through much of the
belly of the muscle, which is nearly the length of the humerus. Distally, the
tendon becomes flat and broad, and passes beneath the tendons of the M.
extensor digitorum communis to the third metacarpal. In Myotis the muscle
originates by fascia from the posterior and dorsal surfaces of the base of the
ulna, and fleshily from all but the first 3 mm. of the posterior surface of the
ulna and the posterodorsal surface of the middle of the radius. The insertion
is the same as that in Eumops. In Myotis the belly of the muscle extends
along roughly the proximal three quarters of the forearm. The muscle is
relatively smaller in Macrotus than in the other two genera, and inserts on the
dorsal base of the fifth metacarpal as in most mammals. The origin is along
the dorsum of the distal half of the ulna and the posterodorsal surface of the
distal half of the radius.
Action.—In Macrotus the fifth digit is extended by this muscle. In Eumops
and Myotis it is a powerful flexor of the third digit, and consequently of the
entire distal part of the wing, and its action is directly antagonistic to that of
the radial extensors.
In Eumops and Myotis the tendon passes over the posterior ridge on the
distal end of the radius, along the edge of the carpus posterior to the third
osteofibrous canal, and over the dorsal bases of metacarpals four and five to
FUNCTIONAL MORPHOLOGY OF THREE BATS 85
its insertion on the base of the third metacarpal. The tendon passes posterior
to the cuneiform bone, and is held there by a heavy fascial sheet. Because the
tendon approaches the third digit from the posterior edge of the carpus, the
tendon serves as an effective posterior flexor of the third digit.
It has been mentioned that when the radius is extended the distal part of
the wing tends to be spread by the M. extensor carpi radialis longus and the
M. extensor carpi radialis brevis. When the bat is using its wings for ter-
restrial locomotion the radius must be partially extended during the forward
component of the stride, but the distal part of the wing must not be allowed
to extend and must remain tightly closed. The principal function of the M.
extensor carpi ulnaris is probably to act against the radial extensors and keep
the distal part of the wing fully flexed during terrestrial locomotion. It is
interesting that, among the bats here considered, only in Macrotus, which al-
most never uses its wings in terrestrial locomotion, does this muscle retain its
insertion on the fifth metacarpal.
Pectoralis Group
M. subclavius (subclavius nerve )
OriciIn.—From flat, ventral surface of first costal cartilage.
INsERTION.—Along all but distal quarter of posterodorsal edge and posterior
third of flat, dorsal surface of clavicle.
ReMarks.—In Myotis this muscle takes fibrous origin from the anterior
part of the distal base of the first rib; the insertion is along the middle two-
thirds of the posterodorsal surface of the clavicle. In Macrotus the origin is as
in Eumops; the insertion is along the middle three-quarters of the posterodorsal
surface of the clavicle.
ActTIon.—This muscle pulls the clavicle posteriad and ventrad, and probably
serves mainly to steady the clavicle against forces pulling its distal ends to-
gether and forward.
M. pectoralis
This is a large, thick sheet of muscle that seems, superficially, to be undi-
vided from the posterior end of the sternum to the shoulder. Actually, the
muscle is composed of two major divisions, one taking origin from the clavicle
and the other from the sternum. This division is present in all three genera.
Macalister (1872:135) distinguished between the pars sternalis and the pars
clavicularis of the pectoralis major in bats. Howell has concluded (1937:457)
that “there is no ready criterion for distinguishing pectoralis minor from major
elements.” Lacking a means for assigning the two divisions of the large
pectoral muscle to either the major or minor elements, I am referring to them
as anterior and posterior divisions.
M. pectoralis, anterior division (anterior thoracic nerve)
Oricin.—From lateral surface of base, and all but distal quarter of flat,
ventral surface of clavicle.
INsERTION.—By fibrous sheet, along dorsal edge of pectoral ridge.
Remarks.—In Myotis the origin extends onto the manubrium and the
muscle itself is divided into three parts in some individuals, but the complete-
ness of the separations and the origins of the parts varies considerably. In all
specimens that I have examined the deep division is more or less separate
86 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
and usually originates as a flat fleshy sheet on the ventral surface of the first
costal cartilage and along the posterior border of the proximal two thirds of
the clavicle; the insertion is by an aponeurosis along 1 mm. of the dorsal
surface of the base of the pectoral ridge. The’ rest of the anterior division is
sometimes separable into two parts: one takes origin from the manubrium and
from the ligament that extends from the body to the spine of the manubrium,
and inserts on the distal two thirds of the pectoral ridge; the other originates
along roughly the proximal half of the ventral surface of the clavicle, and
inserts on the dorsal edge of the proximal third of the pectoral ridge. In
Macrotus the muscle is not divided, and originates from all but the distal 4 mm.
of the ventral surface and edges of the clavicle. The insertion is on the prox-
imal angle of the pectoral ridge.
Action.—This element of the pectoralis muscle pulls the humerus down-
ward and forward and rotates it so that the leading edge of the wing is
lowered.
M. pectoralis, posterior division (anterior thoracic nerve)
OriciIn.—From ventral and lateral aspects of spine and entire ventral sur-
face of the body of manubrium, ventral ridge of sternum and mid-ventral
raphe to within 9 mm. of cartilaginous tip of xiphisternum, and from band
extending 5 mm. laterally over seventh costal cartilage.
INSERTION.—On entire anterior surface of pectoral ridge, by heavy fibrous
sheet.
RemMarks.—In Myotis the origin is from the ventral surface of the posterior
end of the manubrium, the entire sternum and mid-ventral raphe to within
1 mm. of the cartilaginous tip of the xiphisternum, and from a 3 mm.-wide
band extending laterally from the bony part of the xiphisternum along the
abdominal fascia. In Macrotus the origin extends onto the manubrium and
to within 2 mm. of the bifurcate tip of the xiphisternum, thence laterad on
the abdominal fascia for roughly 5 mm.
Action.—The pectoralis muscle is a powerful adductor and rotator of the
humerus and considering the anterior and posterior divisions together is easily
the largest muscle in the bat’s body. This muscle supplies the majority of
the power for the downstroke of the wings. In Eumops the pectoralis muscle
is approximately four times as heavy as the next largest muscle, the posterior
division of the serratus anterior. Because the fibers of the pectoralis muscle
originate from the xiphisternum to near the distal end of the clavicle, the
muscle can govern adduction movements of the humerus through a wide range
of planes. The anterior division draws the humerus downward and sharply
forward, whereas the posterior part of the muscle pulls that bone downward
and backward. The direction of the downstroke can thus be varied con-
siderably by the pectoralis muscle alone. Seemingly the anterior and posterior
divisions usually work together to produce the downstroke. By this common
action the humerus is extended well forward, increasing the surface area of the
wing to its maximum during the downstroke.
The large pectoral ridge on the humerus of bats puts the insertion of the
pectoralis muscle well anterior to the long axis of this bone. Thus, this muscle
acts as a rotator of the humerus, and, because of the rigidity of the elbow
and wrist, tends to rotate the leading edge of the wing downward during the
downstroke. This rotation is important aerodynamically. In birds the forward
FUNCTIONAL MORPHOLOGY OF THREE BATS 87
propulsion for flight is supplied mostly during the downstroke when the distal
primary wing feathers twist in response to the air pressure, each functioning
as a propeller. The lift is supplied largely by the secondary feathers which
remain at a relatively constant angle of attack throughout the cycle of the
wing-stroke. These two main forces, forward thrust and lift, are produced
similarly by the bat wing. The distal part of the wing forms a propeller, with
the pitch increasing distally from the level of the last digit. Although the
digits may “give” slightly dorsoventrally, the bat carpus allows only flexion
and extension in the anteroposterior plane. In order to adjust the angle of
attack of the wing, therefore, the humerus must be rotated. Only the fourth
digit supports the trailing edge of the wing distal to the fifth digit; the second
and third digits form the leading edge of the wing. The fifth digit aids the
hind limb in maintaining the proper angle of the plagiopatagium throughout
the downstroke. Therefore, during the downstroke, while the trailing edge of
the distal part of the wing is tilted upward and is supplying forward pro-
pulsion, the fifth digit and the trailing edge of the plagiopatagium must be
held down to maintain the camber and angle of attack necessary for developing
lift. In other words, the fifth digit must be held at a different angle from the
fourth digit during the downstroke. Interestingly enough, the fifth digit is
modified in several ways in order, seemingly, to meet this requirement. Com-
pared to the fourth digit, the shaft of the fifth metacarpal is curved more
strongly downward, and the phalanges are slightly more flexed when the wing
is spread. In addition, the fifth metacarpal is heavier and is laterally com-
pressed. When not affected by air pressures, the trailing edge of the plagio-
patagium is slightly lower than the trailing edge of the distal segment of the
wing. Under the force of the downstroke this effect is magnified due to the
greater flexibility of the fourth metacarpal and its more distal position in the
wing subjecting it to greater air pressures. Photographs of bats in flight show
clearly the twisting of the distal part of the wing during the downstroke.
Probably, then, the tendency of the pectoralis muscle to rotate the humerus
gives the plagiopatagium an advantageous angle of attack, in terms of lift, and
it puts the fourth digit at an angle from which it may bend slightly and allow
the distal segment of the wing to act as a propeller and supply forward
thrust. The M. pectoralis also controls rotational stability of the wing during
the downstroke by acting against the M. biceps brachii (pages 89-90).
The pectoralis muscles are important also in quadrupedal locomation in
bats. The wings remain folded and the limb more or less flexed while the bat
is walking or running, and the humerus is not vertical to the ground but is
splayed out laterally. Accordingly, the pectoralis muscles must partly support
the weight of the front part of the body by adducting the humerus. When the
body is supported on the non-vertical forelimbs, part of the stride requires a
downward and backward push; this movement is controlled largely by the
pectoralis muscles. Thus, these muscles furnish a large share of the power
necessary for both terrestrial and aerial locomotion,
M. pectoralis abdominalis (anterior thoracic nerve )
Oricin.—From abdominal fascia along strip roughly 9 mm. long and ex-
tending from 8 mm. lateral to midline, at level of base of xiphisternum, obliquely
posteriad to approximately 15 mm. lateral to midline, at level of tip of
xiphisternum.
88 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
INsERTION.—Along middle half of ventral edge of pectoral ridge, by thin
aponeurosis.
RemMarks.—This is a long flat muscle underlying the posterior division of
the pectoralis muscle. In Macrotus the insertion extends from the pectoral
ridge onto the anterior surface of the base of the greater tuberosity of the
humerus. Otherwise the relationships of the muscle are similar in all three
genera.
Action.—This muscle is so situated as to be a strong flexor of the humerus,
and also pulls it slightly downward. The primary function of this muscle is
probably in connection with quadrupedal locomotion, for here this action pro-
duces the propulsion stroke for the front limb. Because of its length, this
muscle can act on the humerus throughout a wide arc, and can fully flex it.
Except in maneuvers demanding deviations from the usual wing-stroke cycle,
this muscle is probably not important in flight.
Flexor Group of Arm
M. coracobrachialis (musculocutaneous nerve)
Remarks.—This muscle is absent in Eumops. The fibrous origin in Myotis
is from the lateral surface of the tip of the coracoid; the fleshy insertion extends
along roughly 1 mm. of the medial surface of the humerus adjacent to the
distal end of the medial ridge. In Macrotus the muscle takes fleshy origin
from the dorsal edge of the tip of the coracoid process; the insertion is by a
thin fibrous sheet on the medial surface of the humerus approximately 5 mm.
beyond the distal end of the medial ridge. The coracobrachialis is smaller
than either of the heads of the biceps in these genera.
Actrion.—This muscle is an adductor and extensor of the humerus. It
probably helps the larger flight muscles depress the humerus; because of its
small size, however, the coracobrachialis can be only a weak adductor. The
coracoid head of the biceps seems to have taken over the function of the cora-
cobrachialis in Eumops.
M. biceps brachii (musculocutaneous nerve)
Oricin.—Short head (coracoid head): from entire expanded tip, and
all but medial surface of distal half of coracoid process of scapula, by thick,
fleshy attachment. Long head (glenoid head): from lateral base of coracoid
process of scapula, adjacent to anterior lip of glenoid fossa, by broad, thick
tendon.
INsERTION.—Into deep slit in anteromedial surface of radius just distal to
head.
ReMaARKS.—Because of the variations in the morphology of the scapula
among bats, it is difficult to apply the standard terminology to the biceps
muscles of these animals. The coracoid process turns strongly laterad in
Macrotus, gently laterad in Myotis, but swings sharply mediad in Eumops.
Accordingly, considering total lengths of muscles and tendons, the long head
of the biceps is longer than the short head in Macrotus, roughly the same
length as the short head in Myotis, whereas in Eumops the long head is
actually shorter than the short head. Because of this possible source of
confusion, it seems preferable to refer to the long head of the biceps as the
glenoid head, and the short head as the coracoid head in the following remarks.
FUNCTIONAL MORPHOLOGY OF THREE BATS 89
In Eumops the glenoid head of the biceps divides into two bellies as it
passes through the bicipital groove. Each belly gives way to a broad, thick
tendon roughly half way along the humerus; these tendons are separate to
their insertions although bound in a common fascial sheath. The coracoid
head of the biceps has a thick, fleshy origin; the belly gives way to a tendon
approximately one third of the way along the humerus. The heavy tendon
is bound in a fibrous sheath along with the tendons of the glenoid head, but
remains a separate tendon. In terms of volume, the coracoid head of the
biceps is larger than the glenoid head.
In Myotis the coracoid head of the biceps originates along the distal fourth
of the coracoid process, and becomes tendinous one quarter of the way along
the humerus. The coracoid head has roughly the same volume as the glenoid
head, The large tendons of the biceps insert in a groove on the ventral surface
of the radius immediately distal to the head of the humerus.
The coracoid head of the biceps takes origin in Macrotus from the enlarged
distal end of the coracoid, and the tendon fuses with the broad, flat tendon
of the glenoid head on the ventral surface of the latter. In contrast to the
other two genera, the coracoid head of the biceps is smaller than is the
glenoid head in Macrotus. The tendons of the biceps insert in a depression
on the ventral surface of the radius immediately distal to its head.
Action.—These muscles are flexors and rotators of the forearm, and ad-
ductors of the wing. They are important in holding the forearm rigidly out-
stretched against the opposing forces of the triceps. Photographs of Macrotus
ir. level flight, and in various stages of performing the half roll preparatory to
alighting, show that the biceps muscles are in a state of contraction through-
out much of the downstroke; in bats with wings almost fully spread the biceps
tendons are pulled well away from the anterior edge of the humerus and ap-
pear as taut cords extending from the front of the shoulder to the proximal
part of the radius. During the downstroke, for the sake of aerodynamic effi-
ciency, the wings must be held stiffly extended against the force of drag
created by the air stream. This steadying of the wing during the power
stroke appears to be an important action of both the M. biceps brachii and
the M. triceps brachii.
Throughout the upstroke the wing is partly flexed. This flexing, however,
is probably passive. Compared with the position of the insertion of the triceps
muscles, the position of insertion of the biceps tendons gives these muscles
much greater mechanical advantage; therefore, under the control of the
tonus of the biceps and triceps the forearm tends to flex.
The insertion of the tendons of the biceps is ventral to the long axis of the
radius in all three genera. Thus, the biceps group acts to rotate the radius,
tilting the leading edge of the wing upward, or, considering the action aerody-
namically, increasing the angle of attack of the wing. This action also is asso-
ciated with stabilizing the wing during the downstroke. The pectoralis muscles
tend to rotate the humerus as they adduct it, their action being to lower the
leading edge of the wing and decrease the angle of attack. Because of the
rigidity of the elbow and wrist in bats, rotation of the brachium or antebrachium
affects the entire wing. The rotating action of the biceps is in opposition to
the rotating action of the pectoralis muscles. Probably the biceps muscles not
only maintain the anteroposterior rigidity of the forearm by acting against the
90 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
triceps, but help control the rotational stability of the wing by acting as a
counter-rotator against the action of the pectoralis. Expressed in terms of
aerodynamics, the angle of attack during the downstroke may be the result
of the antagonistic action of the biceps and pectoralis muscles.
The coracoid head of the biceps is clearly a more important muscle in
Eumops than in either of the other two genera, and more effectively aids in
the downstroke of the wing. It should be stressed prefatory to the following
remarks that the long coracoid process enables the coracoid head of the biceps
to act as an adductor of the humerus by placing its origin below the line of
the long axis of the humerus, and that the greater this displacement, the
greater the mechanical advantage of this muscle as an adductor. The medially
curved coracoid process in Eumops takes the origin of the coracoid head of
the biceps out of the way of the lesser tuberosity of the humerus, allowing
more space for the origin and large belly of the muscle. In addition, the
total length of the muscle and tendon is increased. Probably due primarily
to the modifications in the coracoid process, the coracoid head of the biceps is
relatively larger in Eumops than either of the other genera. Attending the
other changes in Eumops, the function of the muscle is altered. In Myotis and
Macrotus because of the laterally curved coracoid process, the origin of the
coracoid head of the biceps is below the line of the long axis of the humerus
when the wing is at the top of the upstroke, and in this position the muscle
is effective as an adductor of the wing. The greatest mechanical advantage
of the muscle as a brachial adductor is attained as the humerus reaches the
horizontal, for here the origin of the muscle is at its maximum displacement
from the line of the long axis of the humerus. The mechanical advantage of
the muscle as an adductor is reduced, however, as the wing is fully depressed
because the origin of the coracoid head of the biceps is no longer so far below
the long axis of the humerus. Also, because of the relatively shorter length
of this muscle in Myotis and Macrotus, it probably is not effective when the
distance between its origin and insertion is shortened near the bottom of the
downstroke. In short, the adduction action of the coracoid head of the biceps
in these genera is effective at the top of the upstroke, reaches its peak efficiency
when the wing is horizontal, and is reduced when the wing is depressed. In
Eumops, in contrast, the origin of the muscle is only slightly below the line
of the long axis of the humerus at the top of the upstroke because the humerus
and the medially curved coracoid are nearly in line. At this point, then, the
muscle is probably not effective as an adductor. The position of the origin
of this muscle in relation to the line of the long axis of the humerus is lowered
continuously through the downstroke of the wing, however, and the mechanical
advantage of the muscle as an adductor is thus increased continuously through-
out the downstroke. In Eumops, because of the medial curvature of the
coracoid process, the distance between the origin and the insertion of the
coracoid head of the M. biceps is probably always great enough for the muscle
to function. Therefore, in Eumops, the coracoid head of the biceps serves
most effectively as an adductor of the wing in the lower part of the downstroke.
The modifications enabling the coracoid head of the biceps to help in the
adduction of the forelimb seem to constitute an additional refinement in the
system, characteristic of bats, of dividing the work of the powerstroke of the
wing between a number of muscles.
FUNCTIONAL MORPHOLOGY OF THREE BaTs 91
M. brachialis (musculocutaneous nerve )
Oricin.—Along anterior surface of humerus from distal end of pectoral
ridge to within roughly 5 mm. of distal end of humerus.
INsERTION.—Into deep slit in anteromedial surface of radius just distal to
head.
ReMARKS.—In Myotis and Macrotus the origin is from the third quarter of
the anterior surface of the humerus. In all three genera the insertion of this
muscle is the same as the insertion of the biceps muscles. This small muscle
is relatively more robust in Eumops than in the other genera.
Action.—This muscle is a flexor and rotator of the radius, and probably
acts with the biceps muscles to stabilize the wing during the downstroke.
Flexor Group of Forearm
M. flexor carpi ulnaris (ulnar nerve)
Oricin.—By large tendon, from tip of spinous process of medial epicondyle.
INSERTION.—On proximal end of pisiform.
ReMarks.—In Eumops the tendon of origin is 6 mm. long. The belly of
the muscle is thin and flat, and lies along roughly the proximal two thirds of
the forearm. The elasticity of the muscle is reduced by glistening fascial
sheets that extend from the large tendons of origin and insertion and envelop
most of the belly. A tendinous core passes through the belly of the muscle
in Myotis. Otherwise, the muscle is the same as that in Eumops. In Macrotus
the fleshy origin is from the anterior surface of the base of the ulna, the medial
surface of the ulna, and by a thin sheet of muscle fibers along the distal part
of the posterior border of the radius to within roughly 8 mm. of its distal
end. The insertion is by a thin tendon on the proximal end of the pisiform.
Action.—This muscle is a posterior flexor of the fifth metacarpal. Be-
cause it is attached firmly to the ventral base of the fifth metacarpal, the
pisiform serves as a proximal and ventral extension of the base of this bone.
Thus, in terms of effect, when this muscle acts upon the pisiform, it is acting
upon the fifth metacarpal.
In Eumops and Myotis two anatomical modifications seem to enable this
muscle to control the degree of extension or flexion of the fifth metacarpal with
little muscular effort. Because of the long spinous process of the medial
epicondyle in these genera the origin of the M. flexor carpi ulnaris is moved
through a wide arc when the radius is extended or flexed. In Eumops, for
example, the origin of this muscle is moved 4 mm. proximad when the radius
is moved from its position of maximum flexion to maximum extension. In
contrast, due to its ulnar origin, the position of the origin of this muscle is
not affected by movements of the radius in Macrotus. The tendons of this
muscle are thick in Eumops and Myotis and the elasticity of the bellies is
greatly reduced by connective tissue. Any proximal movement of the origin of
this muscle is therefore transmitted to the pisiform through the relatively non-
elastic muscle. Accordingly, probably with little muscular effort on the part
of this muscle, the fifth digit is flexed when the radius is flexed, and the digit
is allowed to extend to roughly a right angle to the humerus when the radius
is extended. Because this muscle does not allow the fifth digit to extend
92 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
beyond this point, the distal part of the wing membrane is kept tautly spread
when the wing is extended.
The transfer to the flexors and extensors of the radius—the biceps and
triceps groups respectively—of much of the work of controlling the distal part
of the wing serves the important end of keeping the sizes of the forearm
muscles in bats to a minimum. This transfer is well illustrated by the M.
flexor carpi ulnaris in Eumops and Myotis, and by the Mm. extensor carpi
radialis longus and brevis in all three genera.
M. palmaris longus (median nerve )
Oricin.—From anterior base of spinous process of medial epicondyle.
INsERTION.—By aponeurosis, on anterovenitral surface of carpus, on thumb
pad, and on ventral surface of M. abductor digiti quinti.
REMARKS.—In Eumops this muscle has a tendinous origin, and the narrow
belly extends along roughly the proximal half of the forearm. The slender
distal tendon broadens into a thin aponeurosis that lies superficial to all the
tendons on the ventral surface of the carpus. The aponeurosis divides into
two parts at the base of the carpus; one passes craniad to the thumb, and
the other passes caudad to the fifth digit. This latter part of the aponeurosis
is much reduced in size in some specimens. I failed to find any trace of this
muscle in Myotis. In Macrotus the fleshy origin is from the distal edge of the
medial epicondyle and on the proximal part of the belly of the M. flexor
digitorum profundus. The muscle is short and robust, and gives way to a
strong tendon roughly one quarter of the way along the radius. The tendon
divides into two parts at the carpus. The first tendon spirals around to the
anterior surface of the thumb where it inserts on the anterior surface of the
distal end of the first metacarpal; the second tendon passes to the ventral base
of the third digit where it divides into two fascial sheets that insert on each
side of the ventral surface of the proximal fifth of the third metacarpal.
Action.—In Eumops this muscle is a weak flexor of the first and fifth
metacarpals. In Macrotus the muscle is relatively larger than in Eumops,
and acts to flex and rotate the first metacarpal, and flex the third metacarpal.
Perhaps the reduction of this muscle in Eumops, and its loss in Myotis, is
correlated with the specialization of the M. extensor carpi ulnaris and the M.
flexor carpi ulnaris as efficient flexors of the distal part of the wing in these
bats.
M. flexor carpi radialis (median nerve )
Oricin.—From distal part of posterior border of M. pronator teres.
INsERTION.—Into fascia at ventral base of first metacarpal.
ReMaArKS.—In Eumops and Myotis this muscle is vestigial. In Eumops
it arises by a thin, delicate tendon from the surface of the M. pronator teres
and has no separate belly. In Myotis the muscle has a tiny belly 2 mm. long
that lies on the distal surface of the M. pronator teres. The distal part of
the tendon is extremely small, and could not be traced beyond the level of
the base of the carpus. In Macrotus the muscle is robust, being roughly the
same size as the M. palmaris longus, and takes fleshy origin from the central
tendon and belly of the middle third of the M. pronator teres. The tendon
of insertion is no smaller than the other flexor tendons of the forearm, and
inserts on the ventral base of the third metacarpal.
FUNCTIONAL MORPHOLOGY OF THREE BATS 93
Actron.—In Macrotus this muscle flexes the third digit. In Eumops and
Myotis the muscle is probably functionless; its extreme reduction in these
genera may be correlated with the development of the M. extensor carpi
ulnaris into a strong flexor of the third digit.
M. pronator teres (radial nerve)
Oricin.—By tendon, from anterior base of spinous process of medial
epicondyle.
INsERTION.—Along roughly proximal eighth of medial surface of radius.
ReMARKS.—In all three genera the muscle becomes broad and flat im-
mediately distal to its origin, has a tendinous central part, and inserts fleshily
on the shaft of the radius. In Myotis and Macrotus the muscle inserts along
the proximal third of the ventral surface of the radius. In Macrotus the origin
is on the proximal part of the medial epicondyle. The central tendon is
larger than in the other genera, and from its middle third the M. flexor carpi
radialis takes origin.
Action.—In bats, because of the nature of the elbow joint, pronation of
the forearm can not occur. Therefore, this muscle is a weak flexor of the
forearm. The most important function of this muscle in bats is that of steady-
ing the elbow joint. The relatively small size of this muscle in Eumops may
be due to the especially strong humeroradial articulation in this genus.
M. flexor digitorum profundus (median nerve)
Oricin.—From anteromedial surface of ulna and roughly proximal three
fifths of posterior border of radius, by fleshy attachments.
INsERTION.—On ventral base of second phalanx of thumb and on ventral
base of fifth digit.
ReMarks.—In Eumops this is the largest muscle of the flexor group of the
forearm. The broad tendon passes immediately medial to the proximal end
of the pisiform and deep to the scaphoid-pisiform ligament, dividing into two
parts at this point. In Myotis the origin is from the spinous process of the
medial epicondyle, from fascia on the medial epicondyle, and from the second
fifth of the posterior border of the radius; the insertion is on the ventral surface
of the second phalanx of the thumb, the ventral surface of the shaft of the
second phalanx of the third digit, and the ventral tip of the third phalanx
(cartilaginous) of the fourth digit. In Macrotus the muscle takes fibrous origin
from the distal part of the medial epicondyle, and fleshy origin on the surface
of the M. pronator teres and M. flexor carpi ulnaris; the insertion is on the
ventral base of the second phalanx of the thumb, and on the ventral base of
the third phalanx of the third digit.
Action.—In Eumops the muscle is a ventral flexor of the thumb and of
the phalanges of the fifth digit. In Myotis the action is ventral flexion of the
thumb and of the phalanges of digits three and four. The tendon to the
thumb is the largest, and that to the fourth digit is the smallest of the three
tendons of insertion. In Macrotus the muscle flexes the thumb and the
phalanges of digit three. In Myotis and Macrotus this muscle can also cause
posterior flexion of digits three and four and digit three respectively.
Flexion of the thumb while the bat is in flight lowers the distal end of the
propatagium, thereby increasing the camber of the proximal part of the wing.
By acting to flex the phalanges of any of the last three digits during the
94 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
downstroke this muscle resists the force of the air pressure that tends to
produce dorsal flexion of both the metacarpal and phalangeal segments of the
digit. In Eumops the tendon to the fifth digit helps maintain the optimal
camber of the plagiopatagium during the downstroke by acting as a ventral
flexor of the phalanges. Because of the great aerodynamic importance of this
action, I can not understand why the tendon to the fifth digit is not retained
in Myotis and Macrotus.
Extensor Group of Manus
M. interosseus dorsale
Oricin.—From posterodorsal base of second metacarpal.
INSERTION.—On anteroventral base of first phalanx of third digit.
ReMarks.—This muscle is present only in Eumops. The belly of the muscle
lies opposite the proximal third of the third metacarpal. The large, flat tendon
extends along the first half of the metacarpal on the anterodorsal surface,
then spirals ventrad to a sesamoid bone that lies on the anteroventral surface
of the third metacarpophalangeal joint. The sesamoid is attached by a short
ligament to the anteroventral base of the first phalanx; the distal part of the
tendon and the sesamoid bone are bound to the joint by a thick sheet of fascia.
Action.—Only in Eumops does flexion and extension of the first phalanges
of the third and fourth digits occur in the anteroposterior plane. Therefore,
this muscle extends the first phalanx of the third digit and partly spreads the
distal part of the wing.
Flexor Group of Manus
I did not determine the innervations of the intrinsic muscles of the hand,
and identified them on the basis of topographic relationships. Because these
relationships in bats differ from those in other mammals, comparisons are
difficult. In addition, the terminology of the muscles of the hand has not been
uniformly applied and the homologies are not completely understood (Rinker,
1954:84). In connection with my work, I found it especially difficult to iden-
tify the muscles of the fifth digit. The names M. abductor digiti quinti and
M. opponens digiti quinti are only tentatively applied; the two muscles may not
be homologous with the muscles bearing the same names in other mammals.
M. abductor pollicis brevis
Oricin.—From posteroventral surface of trapezium and ligament between
ventral base of second metacarpal and trapezium.
INsERTION.—Into the anteroventral part of pad on ventral surface of first
metacarpophalangeal joint (thumb pad).
ReMARKS.—The relationships of this muscle are approximately the same in
all three genera. The muscle is short and robust in Eumops, being relatively
largest in this genus. The muscle is thin, delicate and smallest in Macrotus
and originates largely on the fascia on the ventral surface of the trapezium.
ActT1ion.—This muscle abducts and flexes the first digit.
M. flexor pollicis brevis
Oricin.—From anteroventral surface of base of second metacarpal, from
adjacent ligament that extends from trapezium to second metacarpal, and
from tendon of M. flexor digitorum profundus at level of first metacarpal.
FUNCTIONAL MORPHOLOGY OF THREE BATS 95
INsERTION.—Into posteroventral part of thumb pad.
REMARKS.—This muscle is roughly the same in Eumops and Myotis, but
is relatively larger in the former. In Macrotus the muscle is thin and delicate
and has no part that takes origin from the tendon of the M. flexor digitorum
profundus.
Action.—This muscle is a ventral flexor of the first metacarpal.
M. abductor pollicis
Oricin.—From along first 8 mm. of posteroventral surface of second
metacarpal.
INSERTION.—By small tendon on dorsal base of second phalanx of thumb,
and on lateral border of tendon of M. extensor pollicis brevis where this
tendon passes over dorsum of first phalanx of thumb.
Remarks.—In Eumops this muscle is short and robust. It is relatively
smaller in Myotis; the origin is more restricted and the insertion is on the
lateral surface of the first metacarpophalangeal joint and on the tendon of
the M. extensor pollicis brevis where it crosses the joint. The muscle is absent
in Macrotus.
Action.—In Eumops and Myotis this muscle adducts and rotates the thumb.
M. adductor digiti secundi
Oricin.—From first 9 mm. of ventral surface of second metacarpal, by
fleshy attachment.
INsERTION.—By tendon, on posteroventral surface of trapezium.
ReMarks.—In Myotis this muscle is relatively smaller than in Eumops,
but the attachments are similar. In Macrotus the muscle is absent.
Action.—This muscle acts as a posterior flexor of the second digit and
thus serves to fold the distal part of the wing.
M. abductor digiti quinti
Ornicin.—From posterior border of scaphoid.
INsERTION.—On ventral surface of fifth metacarpophalangeal joint.
Remarks.—In Eumops this muscle takes origin by a broad, thick tendon,
which extends distad over the entire belly of the muscle as a glistening fascial
sheet. The tendon of insertion is also large and flat, and attaches to the
strong fibrous sheath that covers the fifth metacarpophalangeal joint. In
Myotis the origin is from the pointed tubercle that projects posteriorly from
the proximal end of the pisiform; the tendon divides distally and inserts on
either side of the fifth metacarpophalangeal joint. In Macrotus this muscle
is represented by a strong tendon, which runs from the radial sesamoid to the
fifth metacarpophalangeal joint. A fusiform bundle of fibers attaches onto the
dorsal surface of the proximal part of the tendon. This bundle originates on
the pisiform.
Action.—This muscle reinforces the fifth metacarpal against the force of
the air pressure created during the downstroke, and braces the fifth carpo-
metacarpal articulation. Because the muscle is also a weak flexor of the first
phalanx of the fifth digit, it helps hold the proper camber in the distal part
of the digit and resists the upward force of the air pressure against the wing
membrane.
96 UNIVERSITY OF Kansas PuBis., Mus. Nat. Hist.
In the discussion of the action of the pectoralis group of muscles, the im-
portance of the fifth digit in controlling the angle of attack of the plagio-
patagium, and hence the amount of lift created by this part of the wing, was
stressed. In Eumops the M. abductor digiti quinti is specialized to act as a
powerful, partially elastic band that helps maintain the camber of the fifth
metacarpal. The large tendons of the muscle, and the complete covering of
the belly by thick fascia, strongly reduces the elasticity of the muscle. Thus,
probably with little muscular effort, the muscle can act like a strong bow-
string, and can resist forces that tend to straighten the dorsally bowed meta-
carpal. The effectiveness of this action is increased by contraction of the M.
abductor pollicis longus, which pulls the scaphoid craniad and proximad.
This pull increases the distance between the origin and insertion of the M.
abductor digiti quinti. In addition, due to the structure of the carpus, ex-
tension of the fifth digit increases the distance between the scaphoid and the
base of the fifth metacarpal, thus tending to stretch the muscle. In Eumops
and Macrotus, because of its origin on the scaphoid, this muscle spans the
ventral surface of the fifth carpometacarpal articulation and keeps the joint
from allowing slight dorsal extension of the fifth metacarpal during the down-
stroke of the wings. In Macrotus the muscle is nearly non-elastic owing to
the tendon that extends throughout the length of the muscle, and its action is
probably similar to that in Eumops. In Myotis, however, the muscle is smaller
and seemingly weaker than in the other genera. Perhaps powerful bracing
of the fifth metacarpal is not needed in a small bat.
M. adductor digiti quinti
Oricin.—Along first 4 mm. of posteroventral surface of second metacarpal.
INsERTION.—Along proximal fifth of anteroventral surface of fifth meta-
carpal.
REMARKS.—This muscle is present only in Eumops, is broad and thin, and
is one of the largest muscles of the manus.
Action.—This muscle is a flexor of the second digit. Because the M. flexor
carpi ulnaris will not allow extension of the fifth digit beyond a right angle to
the radius when the wing is fully extended, and flexes the fifth digit fully when
the forearm is flexed, the M. adductor digiti quinti must act mainly to fold
the distal part of the wing by flexing the second digit. This muscle is
probably of importance mainly in folding the distal part of the wing when the
bat is at rest, and in holding the wing in a fully flexed position during
terrestrial locomotion. The retention of this muscle in Eumops, and its com-
plete loss in the other two genera, is probably correlated with the importance
of terrestrial locomotion in Eumops.
M. opponens digiti quinti
Oricin.—From distal end of pisiform.
INsERTION.—On ventral surface of fifth metacarpophalangeal joint imme-
diately lateral to insertion of M. abductor digiti quinti.
ReMARKS.—This muscle is similar in all three genera. It has a tendinous
origin and insertion, and is smaller than the overlying M. abductor digiti
quinti.
FUNCTIONAL MorPHOLOGY OF THREE BATS 97
Action.—This muscle acts with the M. abductor digiti quinti to brace the
fifth metacarpal, to flex partly the first phalanx of the fifth digit, and to steady
the fifth carpometacarpal articulation.
Mm. interossei
There are four of these muscles on the ventral surface of the manus; their
arrangement is as follows:
(1) Oricryx.—From posteroventral base of second metacarpal and distal
part of tendon of M. adductor digiti secundi. INsertion.—On pos-
terior surface of third metacarpophalangeal joint.
(2) Oricrn.—From proximal end of pisiform. INSERTION.—On posterior
surface of proximal end of first phalanx of third digit.
(3) Ortcry.—Along posteroventral base of third metacarpal and _ antero-
ventral base of fourth metacarpal. INsERTION.—On anteroventral
surface of fourth metacarpophalangeal joint.
(4) Oricry.—On sesamoid bone that lies on posteroventral base of fourth
metacarpal. INsERTION—On posteroventral surface of fourth meta-
carpophalangeal joint.
Remarxs.—In Eumops the anterior interosseus muscle of the fourth digit
is especially large. The posterior interosseus muscle of the third metacarpal
is small in this genus, and was not found in Myotis or Macrotus. In these
two genera the interosseus muscles of the fourth metacarpal originate on the
unciform.
Action.—These muscles function principally to brace the third and fourth
metacarpals against the air pressure during the downstroke, but the muscles
act also as weak flexors of the phalanges of the third and fourth digits. In
Eumops, because the first phalanx of the fourth digit flexes posteriorly instead
of ventrally as in the other two genera, the anterior interosseus muscle of this
digit extends the first phalanx.
Muscles of Pelvic Girdle and Limb
M. psoas minor (lumbar nerves )
(This muscle should be listed separately as belonging to the myotomic
musculature; it is placed here with the pelvic girdle musculature for the sake
of convenience. )
Oricin.—From a strip roughly 1 mm. wide along ventrolateral surfaces of
bodies of lumbar vertebrae two to four.
INSERTION.—On tip of pubic spine.
ReMarks.—In Eumops this muscle inserts by a large cylindrical tendon
that extends craniad as a heavy fascial sheet to the ventral surface of the
last lumbar vertebra. The middle of the belly of the muscle is dorsoventrally
flattened. The muscle is fleshy for roughly two thirds of its length. In
Myotis the origin is from the ventrolateral surfaces of the first three lumbar
vertebrae; the insertion is on the pubic spine by a flat tendon that comprises
half the length of the muscle. In Macrotus the origin is from the ventro-
lateral surfaces of the bodies of the last thoracic and first three lumbar
vertebrae, and by a few muscle fibers on the M. psoas major. The muscle
4—4357
98 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
is relatively largest in Macrotus, being thick and cylindrical in this genus.
The fleshy part of the muscle extends almost to the insertion.
Action.—Contraction of this muscle tends to pull the ventral part of the
pelvis forward, thereby arching the lumbar section of the vertebral column.
This action may be useful in helping to double up the body when the bat
is grooming itself or when an insect is being pinned against the uropatagium
while the jaws adjust their grip on the prey. In all three genera this muscle
is so robust and inserts by so heavy a tendon, however, that it seems it must
perform some function, in addition to those mentioned above, that demands
powerful action. This function may be to help brace the dorsally arched
vertebral column while the bat is flying, and, at least in Macrotus, when the
bat alights. During flight the bracing would be against the shock transmitted
by the hind limbs and tail to the vertebral column when the air stream sud-
denly strikes the ventral surface of the uropatagium in rapid maneuvers. In
Macrotus the shock of the bat’s alighting is passed from the hind limbs
through the pelvis to the vertebral column, and tends to straighten the dorsally
arched column. By acting as a strong elastic brace, extending from the
anteroventral part of the pelvis to the last thoracic vertebra, this muscle may
absorb part of this shock. The large size of this muscle in Macrotus may be
associated primarily with this bat’s unique method of alighting.
Iliacus Group
M. iliacus (femoral nerve )
Oricin.—From lateral surfaces of bodies of lumbar vertebrae two to five,
from lateral half of ventral surface of expanded anterior end of ilium, and
from all but posterior 3 mm. of lateral edge of ilium.
INsERTION.—On low knob at distal end of lesser tuberosity of femur.
RemMarks.—In all three genera this muscle has a thick fibrous insertion.
In Myotis the origin is from the lateral surfaces of the bodies of lumbar
vertebrae three and four, the lateral edge of the iliac crest, and the lateral
border of the ilium. In Macrotus the origin is from the triangular lateral
surface of the iliac crest, and the insertion is along roughly 1 mm. of the
middle of the medial ridge of the femur. The muscle is short and robust in
Macrotus and has no vertebral origin. This muscle is most evident super-
ficially in Macrotus, for in this genus the muscle is pulled upward and away
from the surrounding muscles by the dorsally directed femur.
Action.—This muscle flexes and rotates the femur. It is most effective as
a rotator in Macrotus, for the insertion is on the medial ridge of the femur,
which is especially high in this genus. In all three genera the rotation this
muscle effects is comparable to supination in terrestrial mammals, but because
of the unusual posture of the pelvic limb of bats the muscle pulls the femur
forward and swings the lower leg ventrad and craniad. Due to the dorsal
flexion of the bodies of these bats the anterior end of the pelvis is tilted
upward. As a result, this muscle not only flexes the femur but draws it
dorsad. In Eumops and Myotis this muscle is important in crawling. Con-
traction of the muscle helps accomplish the forward component of the stride
and tends to bring the lower leg to a right angle with the substrate. In
Macrotus this muscle is probably of major importance in connection with
FUNCTIONAL MORPHOLOGY OF THREE BATS 99
movements made while the bat is hanging from the ceiling of a cave or grotto.
Often when roosting this bat shifts its foothold slightly in preparation for
hanging by one foot, or to change from one foot to the other for hanging.
During this shifting of position the body is generally pulled toward the ceiling
by the flexion of the hind limbs. This flexion is also made preparatory to
launching into flight. Occasionally Macrotus “walks” slowly along the ceiling
of a cave by swinging the body and gaining a new foothold with one foot
while hanging on with the other. During this type of progression the pelvic
limbs are more or less flexed. Much of the power for these flexion movements
is supplied by the Mm. iliacus and gluteus medius.
The M. iliacus is equally important in connection with aerial locomotion.
In all three genera this muscle, together with other members of the iliacus
group, helps attain anteroposterior stability of the hind limb during flight by
acting against the antagonistic extensors of the femur. Rotational stability of
the pelvic limbs may be gained in part by the opposing rotary actions of this
muscle and M. gluteus medius acting against the extensors of the femur. In
Eumops and Myotis, when sudden changes of direction are demanded while
the animals are foraging, the uropatagium may be pulled downward to change
its angle of attack and hence affect the lift it develops or to increase its ef-
fectiveness as a braking surface. The M. iliacus helps in the lowering of the
posterior margin of the uropatagium by rotating the femur and swinging the
shank downward and forward. Because of the different position of the hind
limb in Macrotus this muscle is seemingly not involved in spreading or lowering
the uropatagium.
M. psoas major (femoral nerve )
Oricin.—From ventrolateral surfaces of bodies of last four lumbar verte-
brae (lumbars three to six), from medial half of ventral surface of iliac crest,
and by a short slip from ventrolateral surface of pubis just posterior to pubic
spine.
INSERTION.—On entire anterior surface of lesser trochanter of femur, by
fleshy attachment.
Remarxks.—In Eumops the origin of the muscle is immediately deep to that
of the anterior part of the M. iliacus. The area of origin of the M. psoas major
is approximately 0.5 mm. wide at the level of lumbar vertebrae three, but
broadens posteriorly to cover most of the lateral and ventrolateral surfaces of
the bodies of lumbar vertebrae five and six. In Myotis the origin is from the
last three Jumbar vertebrae (lumbars three to five). In Macrotus the muscle
takes origin from the lateral and ventral surfaces of the bodies of the last four
lumbar vertebrae (lumbars three to six), the ventrolateral surface of the iliac
crest, the anterior half of the lateral rim of the ilium, and by a short slip from
the flat dorsal surface of the pubic spine. This slip, which occurs in Eumops
and Macrotus, is assumed to be part of the M. psoas major, This is not a
certain identification, however, for the innervation of the slip was not deter-
mined.
Action.—This muscle flexes and rotates the femur. When Eumops and
Myotis crawl, the entire iliacus group of muscles probably works together to
move the leg forward at the start of the stride. The net effect of their common
contraction is to pull the femur forward and upward, and to rotate the femur
100 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
in such a way as to bring the shank to a right angle with the substrate. In
Macrotus by flexing the femur, this muscle helps in many movements made
while the bat is roosting. The iliacus group probably functions as a unit during
flight to help steady the hind limb. ,
M. pectineus (femoral and obturator nerves)
Oricin.—From base and lateral surface of pubic spine.
INsERTION.—Along 9 mm. of anteroventral surface of femur, starting roughly
3 mm. distal to lesser trochanter.
ReMaArRKs.—In Myotis the origin is from the pubic spine, and the insertion
is along 2 mm. of the anteroventral surface of the femur starting immediately
distal to the insertion of the iliacus. The origin is from the posterior two
thirds of the pubic spine in Macrotus, and the insertion is on the anterior
surface of the ventral ridge of the femur. This muscle is relatively largest in
Eumops, and the insertion extends farther distad than in the other genera.
The muscle can therefore act on the femur most powerfully in Eumops.
Action.—Adduction and flexion of the femur.
Gluteal Group
M. tensor fasciae latae ( gluteal nerve)
Oricin.—From posterodorsal border of iliac crest and neural spine of first
sacral vertebra.
INSERTION.—On anterior border of M. gluteus maximus and center of lateral
surface of femur.
RemMarks.—In Myotis the muscle arises from the posterodorsal edge of the
iliac crest and the neural spines of sacral vertebrae one to three. The in-
sertion is along the distal part of the anterior edge of the M. gluteus maximus
and on the proximal end of the lateral ridge of the femur. In Macrotus the
M. tensor fasciae latae is continuous with the M. gluteus maximus. That part
of the sheet of muscle which originates from the posterodorsal edge of the
iliac crest and the middorsal fascia over the first sacral vertebra probably
represents the M. tensor fasciae latae. The insertion is along the anterior edge
of the M. gluteus maximus.
Action.—This muscle abducts and flexes the femur. The functional signi-
ficance of these actions is discussed in the remarks on the M. gluteus maximus.
M. gluteus maximus (gluteal nerve )
OricIn.—From neural spines and middorsal fascia of last three sacral verte-
brae (sacral vertebrae two to four) and neural spine of first caudal vertebra.
INSERTION.—Along approximately 3 mm. of middle of lateral surface of
femur.
ReMARKS.—Relative to the total mass of the muscles of the thigh, this
muscle is larger in Eumops than in the other genera. In Myotis the origin is
from the middorsal fascia and neural spines of the last two sacral vertebrae
(sacral vertebrae three and four) and the neural spine of the first caudal
vertebra. The insertion is along roughly the second fifth of the lateral surface
ot the femur, starting on the tubercle at the distal end of the lateral ridge of
the femur. In Macrotus the origin is from the fused neural crests of the last
FUNCTIONAL MORPHOLOGY OF THREE BATS 101
four sacral vertebrae (sacral vertebrae two to five) and the neural spine of the
first caudal vertebra. The insertion is on the posterior surface of the M. rectus
femoris and along roughly the middle third of the anterolateral surface of
the femur.
Action.—This muscle, together with the M. tensor fasciae latae, abducts
the femur and flexes it when the hind limb is extended. Seemingly, the pri-
mary function of these muscles is to help control the stability of the pelvic
limbs during flight by acting against the antagonistic adductors of the femur.
The hind limbs serve as the base for the posterior part and trailing edge
of the plagiopatagium. Dorsoventral movements of the hind limbs while the
bat is in flight change the camber and angle of attack of the plagiopatagium,
as well as lowering or raising the uropatagium. It is of great importance,
therefore, that during flight the hind limbs be stabilized at the proper angle
to maintain the aerodynamic efficiency of the flight membranes. There are
a number of forces tending to move the hind limbs during flight. The force
of the air stream against the uropatagium tends to move the limbs. Par-
ticularly at the top of the upstroke and bottom of the downstroke of the wings,
the elastic wing membranes tend to disrupt the dorsoventral stability of the
hind limbs. At all times while the wings are extended, the wing membranes
pull laterally with considerable force on the pelvic limbs, and when the bat is
in flight the air pressure against the wing membrane stretches them still further,
increasing the lateral pull. Thus, while a bat is in flight, the hind limbs must
partly resist forces tending to cause their adduction, abduction, and extension.
The uropatagium is probably of some help in bracing the pelvic limbs
against lateral pull, but the major burden of maintaining their rigidity is on
the muscles of the pelvic girdle. For this reason, the functions of these muscles
in bats can not be explained simply in terms of terrestrial locomotion, but
rather should be considered in relation to both terrestrial and aerial locomo-
tion. The latter has perhaps had the greater effect on the evolution of the
hind limb musculature. Certainly, in bats that roost by hanging and that
seldom practice any sort of terrestrial locomotion, the morphology of the pelvic
limb must be in large part the result of adaptations for flight.
In Eumops and Myotis, then, the M. gluteus maximus acts to maintain the
dorsoventral stability of the hind limb by resisting the downward pull caused
by the wing membranes at the bottom of the downstroke. In Macrotus, due
to the different position of the pelvic limbs, this muscle resists lateral pull on
the limbs.
M. gluteus medius (gluteal nerve )
Oricin.—From posterodorsal edge of iliac crest and iliac fossa.
INSERTION.—On greater trochanter of femur.
ReMarks.—The origin extends onto the lateral surface of the neural spine
of the first sacral vertebra in Myotis. In all three genera this is a thick,
barrel-shaped muscle with a fleshy origin and insertion.
Action.—This muscle flexes, abducts and rotates the femur. In terrestrial
locomotion this muscle helps in the forward part of the stride. Probably more
important, however, is the action of bracing the hind limb during flight by
acting with the iliacus group against the powerful, antagonistic extensors and
adductors of the femur.
102 UNIVERSITY OF KANSAS PuBxs., Mus. Nat. Hist.
Quadriceps Femoris Group
M. quadriceps femoris (femoral nerve )
In the bats under study this muscle is composed of two parts, the M. rectus
femoris and a muscle that probably represents, in Eumops at least, the fused
Mm. vastus lateralis, vastus medialis and vastus intermedius. In the other
two genera the part of the M. quadriceps femoris that originates on the femur
is probably homologous with the M. vastus lateralis in other mammals. For the
sake of convenience, the femoral division of the quadriceps is here called the
M. vastus lateralis, although all of it may not be homologous with this muscle
in other mammals.
Oricin.—M. rectus femoris: from lateral edge of ilium immediately anterior
to acetabulum. M. vastus lateralis: from greater trochanter, from proximal
two thirds of anterolateral surface and entire anterior surface of femur.
INSERTION.—On proximal end of patella.
RemMarks.—In Eumops the M. rectus femoris has a fibrous origin via a small
sesamoid bone. The M. vastus lateralis attaches fleshily to the posterior surface
of the M. rectus femoris throughout the length of the latter. The two muscles
merge roughly half way along the femur and form a broad, flat tendon of
insertion. In Eumops the M. vastus lateralis is approximately twice as large
as the M. rectus femoris. In Myotis this muscle takes origin from a small
tubercle immediately anterior to the acetabulum. The M. vastus lateralis
originates on the anterior surface of the greater trochanter and the proximal
third of the anterolateral surface of the femur. In Macrotus the M. rectus
femoris has an unusually strong origin via a sesamoid bone on the anterodorsal
rim of the acetabulum, and is larger than the M. vastus lateralis. This muscle
originates from the greater trochanter and by fleshy attachment along the
proximal fourth of the lateral surface of the femur. In Macrotus the tendons
of insertion of these two muscles remain separate, but are bound in a common
fascial sheath.
Action.—This muscle extends the shank at the end of the rearward com-
ponent of the stride in Eumops and Myotis but probably is even more im-
portant in steadying the shank during flight. The muscles that flex the shank
and supply most of the resistance to the lateral pull exerted by the wing mem-
branes are the gracilis, the semimembranosus and the semitendinosus. These
are among the largest muscles in the pelvic girdle, and they are directly an-
tagonistic to the M. quadriceps femoris. This muscle must act against these
flexors throughout most of the time that the bat is in flight. The lateral pull
caused by the wing membranes is greatest during the downstroke of the wing,
but is greatly reduced in the upstroke largely because the wings are then partly
folded. This intermittent pull is probably not resisted by intermittent con-
tractions of the flexors of the shank, for this would demand considerable energy
at a time when a premium is placed on energy conservation. It is likely that
simply the tonus or weak contractions of the powerful flexors of the shank and
the M. quadriceps femoris is sufficient to steady the shank during level flight.
Powerful bracing action, calling for strong contractions of many of the muscles
of the pelvic girdle, may be needed only while the bat is making rapid ma-
neuvers to catch insects or avoid obstacles. In the course of these maneuvers
the wings often beat more rapidly than usual and are moved into positions
FUNCTIONAL MORPHOLOGY OF THREE BATS 103
causing more air pressure to be directed against the membranes of the wing.
The retention of a large, powerful M. quadriceps femoris in Macrotus,
a bat that seldom uses terrestrial locomotion, suggests that this muscle has an
important function connected with flight. Because of the 180 degree rotation
of the hind leg in Macrotus, contraction of this muscle has a different function
in relation to flight in this genus from that in the other genera, although the
action (extension of shank) is the same. In Macrotus this muscle helps control
the dorsoventral stability of the shank and resists the downward pull of the
plagiopatagium at the bottom of the downstroke of the wings. In addition,
the placement of the origin of the M. rectus femoris and the length of its
belly and tendon help direct the femur dorsad. Aerodynamically, this func-
tion is important. The entire vertebral column of Macrotus is arched sharply
dorsad with the result that the long axis of the pelvis is nearly dorsoventral.
Therefore, to create the proper angle of attack and camber of the plagio-
patagium, the hind legs must be held in a spider-leg-like posture with the
femur directed dorsad. The large size of the M. rectus femoris and the shift
of its origin toward the dorsal rim of the acetabulum in Macrotus is probably
associated with the peculiar posture of the pelvic limb in this genus.
Tibial Extensor Group
M. extensor digitorum longus (deep peroneal nerve )
Oricin.—F rom lateral condyle of femur and posterior border of M. extensor
hallucis longus.
INsERTION.—On dorsal bases of distal phalanges of digits two to five.
ReMarKs.—In all three genera the tendon is undivided as far distally as the
dorsum of the tarsus; there the tendon divides into the tendons that continue
lo the insertions on the digits. In Eumops the fleshy part of this muscle lies
along the entire length of the shank. The muscle originates by a short tendon
on the lateral condyle of the femur in Myotis and the belly extends along
roughly the proximal two thirds of the shank. The insertion is on the dorsal
bases of the distal phalanges of digits one to five. In Macrotus the origin
is from the lateral condyle of the femur and along the lateral border of the
proximal third of the tibia; the insertion is on digits one to five, as in Myotis.
In Macrotus the belly of the muscle is relatively thinner than in the other
genera, and lies approximately opposite the proximal half of the shank.
Action.—Extension of the digits of the foot.
M. extensor hallucis longus (deep peroneal nerve)
Oricin.—From lateral condyle of femur and anterior border of M. extensor
digitorum longus.
INSERTION.—On medial surface of base of first metatarsal.
Remarxs.—This muscle is relatively much larger in Eumops than in the
other two genera. In Eumops it is the anteriormost muscle of the shank, and is
fleshy to the level of the distal part of the tarsus. In Myotis the origin is from
all but the proximalmost 3 mm. of the posterior surface of the tibia, and from
the lateral surface of the distal third of the fibula. The insertional tendon
passes along the medial surface of the tarsus, is bound by fascia to the medial
surface of the base of the first metatarsal, and inserts on the lateral surface
of the base of the first phalanx of the first digit. This is an extremely delicate
104 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
muscle having a thin insertional tendon in Macrotus. The origin is by a nar-
row band of muscle fibers from the lateral surface of the distal half of the
tibia; the insertion is on the medial base of the first metatarsal.
Action.—This muscle swings the foot craniad and dorsad. The large size
of this muscle in Eumops is probably associated with this animal’s roosting
habits and well-developed ability to crawl. In the forward component of the
stride the foot swings forward in order to increase the length and effectiveness
of the stride; this movement is controlled largely by this muscle. Its degenerate
nature in Macrotus is probably correlated with this bat’s habit of roosting by
hanging and the unimportance of terrestrial locomotion in this genus.
Mm. extensores breves (deep peroneal nerve )
Oricin.—There are seven of these muscles. The medial slip takes origin
from the anterodorsal surface of the distal end of the fibula, and the lateral
slip from the dorsal surface of the proximal part of the calcaneus. The other
five slips originate on the tuberosity which projects dorsad from the distal
part of the calcaneus.
INSERTION.—The medial slip inserts on the dorsomedial surface of the base
of the first phalanx of the first digit, and the lateral slip on the dorsolateral
surface of the base of the first phalanx of the fifth digit. The other five slips
insert on the distal phalanges of digits one to five.
ReMarks.—There are no important differences in these muscles among the
three genera.
Action.—Extension of the digits.
Peroneal Group
M. peroneus longus (peroneal nerve )
Oricin.—From posterior part of lateral condyle of tibia and proximal third
of anterolateral surface of fibula.
INsERTION.—On ventrolateral base of fourth metatarsal.
RemMarks.—In Eumops the insertional tendon crosses to the lateral edge
of the tarsus, passes over the dorsal surface of the cuboid, and through the
space between the distal end of the calcaneus and the proximal end of the
fifth metatarsel to the ventral surface of the foot. The belly of the muscle
is broad and flat, and extends along roughly the proximal three quarters of
the shank. The origin in Myotis is from the head and lateral surface of the
proximal two thirds of the fibula. The insertional tendon follows approximately
the same course as that in Eumops, but inserts on the ventral base of the third
metatarsal. In Macrotus the origin is from the anterior surface of the base
and the thin distal remnant of the fibula. The insertional tendon follows a
course similar to that in the other two genera, but continues across the ventral
bases of metatarsals three and four to which it is bound by fascia, and inserts
on the base of the second metatarsal. In Macrotus the tendon of this muscle
is the largest of the extensor tendons of the shank.
ActTIonN.—This muscle rotates the foot, tending to pull the lateral part of
the foot dorsad and laterad. In terrestrial locomotion in Eumops and Myotis
the result of the common contraction of this muscle and the M. gastrocnemius
FUNCTIONAL MORPHOLOGY OF THREE BATS 105
is to pull the foot laterad and bring it into line with the shank, thus giving
a final push at the end of the propulsion stroke of the stride. In Macrotus
the rotation of the foot controlled by this muscle may be important in alighting,
for in this maneuver the feet seem to be so rotated that the plantar surfaces
tend to face one another. This muscle also helps to flex the foot and releases
its grip on the substrate as Macrotus launches into flight.
M. peroneus brevis (peroneal nerve )
OriciIn.—From approximately the middle half of anterolateral surface of
fibula.
INSERTION.—On dorsolateral surface of base of fifth digit.
REMARKS.—The insertion in Myotis is on the dorsal surface of the shaft
of the fifth metatarsal. The origin in Macrotus is from the anterolateral surface
of the shaft of the fibula.
Action.—Flexion and rotation of the foot. In general, this muscle has
the same action as the M. peroneus longus.
Adductor Group
M. gracilis (obturator nerve)
OriciIn.—From posterior 3 mm. of insertional tendon of M. psoas minor
and ventrolateral edge of pelvis from tip of pubic spine to posteroventral angle
ot ischium.
INSERTION.—By a common tendon with M. semitendinosus on postero-
medial surface of tibia roughly one third of the way along shank.
ReMaARKS.—In Eumops this muscle and the M. semimembranosus are almost
exactly the same weight and are the two largest muscles in the pelvic girdle.
Among the bats under study, the M. gracilis is relatively largest in Eumops.
In this genus the broad, flat belly extends nearly the length of the thigh -and
gives rise to a heavy tendon shared by the M. semitendinosus. In Myotis
the origin is along approximately the anterior two thirds of the ventrolateral
edge of the pelvis including the pelvic spine; the insertion is on the posterior
surface of the tibia one sixth of the way along its length. The belly of the
muscle in Myotis lies along the proximal half of the femur. In Macrotus
the muscle arises from along the entire ventrolateral rim of the pelvis, including
the pubic spine; the insertion is by a separate tendon on the posteromedial
surface of the tibia one tenth of the way along the shank. The flat belly
extends less than half way along the femur. The tendons of the M. gracilis
and the Mm. semimembranosus and semitendinosus are separate and are
not fused, but they are bound together by fascia and appear superficially to
be a single tendon.
Action.—This muscle is a flexor of the shank and adductor of the hind
limb. In Eumops and Myotis it is important in connection with both terrestrial
and aerial locomotion. In Macrotus the M. gracilis is important in flight and
by flexing the shank it aids in many movements made while the bat is roosting.
In Eumops and Myotis the M. gracilis helps support the weight of the
body during terrestrial locomotion, but its importance as an adductor may
be limited by its great power as a flexor of the shank. Only when flexion of
106 UNIVERSITY OF Kansas Pus.s., Mus. Nat. Hist.
the shank is resisted by the antagonistic Mm. rectus femoris and vastus
lateralis or by the grip of the claws on the substrate may the M. gracilis act
as an adductor.
The large size of the M. gracilis in Eumops and Myotis and the fusion
of the insertional tendon with that of the heavy M. semitendinosus seem to be
associated more with the importance of the M. gracilis during flight than with
its function in terrestrial locomotion. The fusion of the tendons of the Mm.
gracilis and semitendinosus indicates that these muscles perform their most
important function by working together. The action caused by their simul-
taneous contraction is flexion of the shank. This action, in which these muscles
must usually be aided by the M. semimembranosus, helps steady the hind
limb by bracing the shank against the lateral pull exerted by the wing mem-
branes. Photographs of bats in flight show that the flexors of the shank are
under tension during the downstroke of the wing, and it is probably at this
time that the lateral pull by the wing membranes is strongest. In Macrotus,
with its delicate flight membranes and slow flight, the lateral pull is probably
not great. In Eumops, in contrast, the flight membranes are thick and leathery
and strongly braced with elastic cartilage, and the flight is rapid. Accordingly,
the lateral pull on the shank must be strong during the downstroke. In
Eumops the pronounced specializations of the M. gracilis (the extension of its
origin onto the insertional tendon of the M. psoas minor, the broad _ belly
extending nearly the length of the shank, the complete fusion of its insertional
tendon with that of the M. semitendinosus, and the distal position of the
insertion) all serve the end of developing a strong system of flexors of the
shank. These modifications have probably attended the development of
speedy flight in this genus and are not primarily associated with its strong
terrestrial locomotion.
In Macrotus, because of the posture of the pelvic limbs, the M. gracilis
does not brace the shank against lateral pull but serves mainly to resist the
dorsal pull by the wing membranes on the shank at the top of the upstroke
and beginning of the downstroke. In general, this muscle acts against the
Mm. rectus femoris and vastus lateralis to maintain the dorsoventral stability
of the hind limb.
M. adductor longus ( obturator nerve )
Oricin.—From lateral surface of ventral half of ascending ramus and pos-
teroventral angle of ischium and from lateral surface of pubis posterior to
pubic spine.
INsERTION—Along roughly proximal third of posteromedial surface of
femur starting immediately distal to lesser trochanter.
ReMARKS.—In Eumops this is a large, robust muscle having fleshy attach-
ments. The anterior part of the origin covers the entire lateral surface of
the pubis from its ventral border to the ventral rim of the obturator fenestra.
In Myotis the origin is from the lateral surface of the posteroventral angle
of the ischium and the insertion is along the middle third of the posteromedial
surface of the femur. In Macrotus the origin is along the lateral surface
of the ventral border of the pelvis from the posteroventral angle of the ischium
FUNCTIONAL MORPHOLOGY OF THREE BaTs 107
to the base of the pubic spine; the insertion is on the proximal half of the
medial ridge of the femur.
Acrion.—This muscle adducts the femur. The muscle is large in Eumops
and Myotis and its insertion extends well beyond the lesser trochanter of the
femur, thus enabling it to act powerfully on the femur. The muscle is relatively
small in Macrotus and the insertion is situated farther proximad than in the
other two genera. These differences are probably due to the contrasts between
the functions of this muscle in the bats under discussion. This muscle supports
the weight of the posterior part of the body during terrestrial locomotion in
Eumops and Myotis, but in Macrotus it is used primarily for flight. Also,
with the other adductors of the femur, this muscle aids in the dorsoventral
stabilization of the pelvic limbs during flight in Eumops and Myotis. In
Macrotus, because of the different position of the hind limbs, this function
is largely taken over by other muscles. The femoral adductors in this genus
seem to be more important in connection with erratic flight, in which the
hind legs may be adducted—pulled downward to roughly their usual position
in the other two genera—in order to increase the area of the uropatagium.
Photographs of Macrotus hovering or turning sharply, show the legs adducted
and the uropatagium spread. This movement is probably mostly under the
control of the adductors of the femur.
M. adductor brevis (obturator nerve )
Oricin.—From along 3.5 mm. of lateral surface of ilium adjacent to
posteroventral rim of obturator fenestra.
INsERTION.—Along 4 mm. of proximal part of posterolateral surface of
femur starting on ridge of greater trochanter.
Remarxks.—In Myotis the muscle arises from the lateral surface of the
ascending ramus of the ischium adjacent to the posterodorsal rim of the
obturator fenestra; the insertion is along 1.5 mm. of the posteromedial surface
of the femur starting just distal to the trochanteric fossa. In Macrotus the
muscle originates on the lateral surface of the posteroventral angle of the
ischium and inserts along the distal half of the medial ridge of the femur.
Action.—In Eumops this muscle adducts and rotates the femur; as a
result the shank is pulled downward and tends to become vertical to the
substrate. This muscle acts solely as an extensor of the femur in ects In
Macrotus the action adducts the femur.
This muscle (and most other muscles) probably does not act ance but
usually in co-operation with certain other adductors of the femur (adductor
longus, pectineus and obturator externus) to steady the pelvic limb against
the opposing abductors during flight. The net effect of the common action
of the abovementioned adductors is not only adduction but also rotation of
the femur; this rotator is counter to the rotation caused by the action of the
main abductors (M. tensor fasciae latae, M. gluteus maximus and M. gluteus
medius). During flight the antagonistic pull fo these two groups of muscles
probably stabilizes the pelvic limbs.
In Eumops and Myotis this muscle functions in connection with terrestrial
locomotion; in Eumops the action is to help support the posterior part of
108 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
the body and hold the shank vertical to the substrate, and in Myotis the muscle
aids in the propulsion stroke of the hind limb.
M. adductor magnus (obturator nerve )
Oricin.—Along 4 mm. of dorsal border of obturator fenestra adjacent to
ischial tuberosity.
INsERTION.—On posterior surface of greater trochanter of femur.
REMARKS.—The origin of this muscle is approximately the same in all three
genera. In Myotis and Macrotus the insertion is along roughly 1 mm. of the
posteromedial surface of the femur immediately distal to the level of the
trochanters.
ActTion.—This muscle extends and rotates the femur. In Eumops the ro-
tation tends to swing the shank downward and forward and orient it verti-
cally to the substrate. In Myotis and Macrotus the rotation is in the opposite
direction, and tends to pull the shank upward and backward.
M. obturator externus (obturator nerve )
Oricin.—Along 3 mm. of rim of obturator fenestra adjacent to posteroven-
tral angle of ischium.
INSERTION.—On posteromedial surface of greater trochanter.
ReMaARKS.—In Eumops the insertion is by a short, flat tendon. In Myotis
the origin is along the ventral half of the rim of the obturator fenestra; the fleshy
insertion is in the trochanteric fossa. The attachments are similar in Myotis
and Macrotus.
Action.—Extension and adduction of the femur.
Ischiotrochanteric Group
M. gemellus (sciatic nerve )
Oricin.—Along dorsal border of superior ramus of ischium from adjacent
to posterior rim of acetabulum to superior tuberosity.
INSERTION.—On posterior surface of greater trochanter of femur.
RemMarks.—This muscle was not found in Myotis. In Macrotus it is a
thin band of fibers extending from along 1 mm. of the dorsal rim of the su-
perior ramus of the ischium immediately posterior to the acetabulum to the
posterior cartilaginous rim of the acetabulum. This muscle seems to be a
vestige of one or both of the gemellus muscles.
Action.—In Eumops this muscle is a weak extensor and rotator of the
femur. In Macrotus the muscle is probably functionless.
Hamstring Group
M. caudofemoralis (tibial nerve )
Oricin.—From transverse process of last sacral vertebra and lateral surface
of body of first caudal vertebra.
INsERTION.—Along proximal half of posterior surface of femur.
- Remarxks.—In all three genera the insertion starts immediately distal to the
level of the trochanters of the femur. In Eumops the posterior part of the
origin is fibrous; otherwise the attachments are fleshy. In Myotis the origin
is from the transverse process of the last sacral vertebra; the insertion is
FUNCTIONAL MORPHOLOGY OF THREE BATS 109
along the proximal third of the posterior surface of the femur. The muscle
takes origin in Macrotus from the posterior part of the transverse process of
the last sacral vertebra and the transverse ridge of the first caudal vertebra.
The insertion is along the proximal quarter of the posterolateral surface of
the femur.
Action.—This muscle acts mainly to extend the femur. By acting against
the flexors of the femur the muscle helps steady the hind limbs during flight.
In terrestrial locomotion the muscle acts with the other members of the
hamstring group of muscles to produce the rearward component of the stride.
In Macrotus, in contrast to the other genera, the origin is on the posterolateral
surface of the femur, and the muscle abducts as well as extends the femur.
The action steadies the pelvic limb against the lateral pull of the wing mem-
branes.
M. semitendinosus (tibial nerve)
Oricin.—From tip and lateral surface of rim of ‘dorsal ischial tuberosity.
INSERTION.—On posterior surface of tibia one third of way along shank.
REMARKS.—This muscle and the M. gracilis insert by a common tendon
in Eumops and Myotis. In the latter genus the origin of the M. semitendinosus
is from the entire lateral surface of the dorsal ischial tuberosity; the insertion
is on the posterior surface of the tibia one sixth of the way along the shank.
In Macrotus this muscle takes origin by two heads: (1) from the entire lateral
surface of the dorsal ischial tuberosity, by fleshy attachment; (2) from the
middle of the dorsal border of the ischium, by fibrous attachment. The in-
sertion is on the posterior surface of the tibia one eighth of the way along its
length. The insertional tendons of the two heads are separate but are bound
in a common fascial sheath. In Eumops and Myotis the M. semimembranosus
is much larger than the M. semitendinosus. In Macrotus this situation is re-
versed; the first head of the M. semitendinosus is at least twice as large as
either the second head or the M. semimembranosus.
Action.—This muscle extends the femur and flexes the shank. The
functional significance of this action during flight in Eumops and Myotis is
discussed in the account of the M. gracilis. In these genera the M. semiten-
dinosus also provides much of the power for the propulsion part of the stride
of the hind limb. In Macrotus this muscle helps steady the hind limb against
the dorsal pull of the wing membranes by acting as a ventral flexor of the
shank. The dorsal ischial tuberosity is turned more sharply mediad in
Macrotus than in the other genera, thus putting the origin of the semitendinosus
medial to the head of the femur. Because its origin is posterior and medial
to the head of the femur this muscle is an effective abductor of the hind limb
in the genus, and helps the Mm. gluteus maximus and caudofemoralis resist
the lateral pull of the wing membranes. The flexor action of the M. semitendi-
nosus is also important when Macrotus is making shifts in roosting posture
and slight changes in roosting location.
M. semimembranosus (tibial nerve)
Oricin.—From lateral surface of superior tuberosity and caudal border of
ischium.
INsERTION.—On posteromedial surface of tibia roughly one quarter of way
along shank.
110 UNIVERSITY OF KAnsAS PuBis., Mus. Nat. Hist.
Remarxs.—In Eumops this muscle and the M. gracilis are the two largest
muscles of the pelvic girdle. The M. semimembranosus inserts by a broad,
flat tendon that comprises approximately two fifths of the length of the
muscle. In Myotis the origin is on the lateral surface of the caudal border
of the ischium. The insertion is on the posteromedial surface of the tibia
roughly one sixth of the way along the shank, and the insertional tendon
comprises half the length of the muscle. This muscle is relatively smaller in
Macrotus than in the other genera. The origin in Macrotus is from the dorsal
half of the caudal border of the ischium; the insertion is on the posterior
surface of the tibia roughly one eighth of the way along the shank. The
insertion of this muscle is proximal to that of the M. gracilis in all three
genera, the separation of the tendons being greatest in Eumops and least in
Macrotus.
Action.—This muscle extends the femur and flexes the shank. In Eumops
and Myotis this muscle acts with the Mm. semitendinosus and caudofemoralis
to supply most of the power for the propulsion part of the stride of the hind
limb in quadrupedal locomotion. The origin of the M. semimembranosus is
ventral to the acetabulum due to the tilting upward of the anterior end of
the pelvis in these bats. Extension of the femur, therefore, results in pulling
the femur caudad and ventrad, this action tending not only to help in the
main propulsion movement of the stride of the hind limb but also to assist
the adductors of the femur in supporting the weight of the posterior part
of the body. This muscle is also a strong flexor of the shank, but when the
bat is walking the hold the claws maintain on the substrate probably limits
the flexor action of the muscles and causes it to function as an extensor of the
femur. In Macrotus, by flexing the hind limb, this muscle would help in
movements made while the bat is roosting.
This muscle is important during flight. In Eumops and Myotis it steadies
the hind limbs against the lateral pull of the wing membranes, and in
Macrotus, serves mainly to resist the dorsal pull exerted by the wing mem-
branes.
M. biceps femoris (tibial nerve)
OriciIn.—From lateral surface of tip of dorsal ischial tuberosity.
INsERTION.—On lateral condyle of tibia.
REMARKS.—In Eumops this is a thin, straplike muscle, and has a tendinous
origin and insertion. In Myotis the muscle is absent. In Macrotus it is
vestigial, being represented by a delicate strand of muscle approximately 0.25
mm. in breadth. The origin is from the fascia overlying the dorsal ischial
tuberosity and the insertion is on the lateral surface of the knee.
AcTIon.—This muscle flexes the shank and extends the femur in Eumops.
In Macrotus the muscle is probably functionless due to its small size.
Flexor Group of Leg
M. gastrocnemius (tibial nerve)
Oricin.—Medial head: from posteroventral surface of femur immediately
proximal to medial condyle. Lateral head: from posterior surface of head
of fibula and lateral condyle of femur.
INsERTION.—On proximal end of calcaneus.
FUNCTIONAL MORPHOLOGY OF THREE BATS j OE
RemMarks.—The connections of this muscle are similar in all three genera.
In Eumops the belly is robust and extends the length of the shank. The
muscle is relatively more slender in Myotis and the belly gives way to the in-
sertional tendon near the middle of the shank. In Macrotus the muscle is
extremely slender; and roughly three quarters of its length is tendinous.
Action.—Extension of the foot.
M. plantaris (tibial nerve)
Oricin.—Along posterior surface of proximal two thirds of tibia and antero-
medial surface of proximal half of fibula.
INSERTION.—On ventral surfaces of distal phalanges of digits one to five.
RemMarks.—In Eumops the large tendon of this muscle broadens into an
aponeurosis in the plantar region and there is joined by the tendon of the M.
flexor digitorum fibularis (Fig. 17). From this junction a tendon extends onto
each digit. In Myotis the origin is from the posterior surface of the proximal
two thirds of the tibia. The aponeuroses of the Mm. plantaris and flexor
digitorum fibularis are fused at their distal ends in this genus, thus, they are
not so completely fused as in Eumops. The origin of the M. plantaris in
Macrotus is along the proximal half of the posterolateral surface of the tibia,
from the tendon of the M. popliteus, and by a few fibers from the vestigial
fibula. The tendon of the M. plantaris, which extends to the hallux, separates
from the main tendon at the level of the proximal part of the tarsus and does
not connect with the plantar aponeurosis. The degree of fusion of the two
aponeuroses is similar in Macrotus and Myotis.
The unspecialized condition in mammals is for the plantaris tendon to divide
into a superficial and a deep aponeurosis in the plantar region; the ventral
surface of the deep part serves as origin for the M. flexor digitorum brevis.
Deep to the two plantar aponeuroses and separate from them, lies the apo-
neurosis of the M. flexor digitorum fibularis, from which arise the Mm. lum-
bricales. The bat genera considered here differ from this general plan in the
following ways: the superficial plantar aponeurosis is absent; the M. flexor
digitorum brevis is completely separate from the tendon of the M. plantaris;
the deep plantar aponeurosis is fused distally to that of the M. flexor digitorum
fibularis; the Mm. lumbricales originate on the ventral surface of the plantar
aponeurosis. Seemingly, in the evolution of bats, the M. flexor digitorum brevis
gained an origin separate from the deep plantar aponeurosis, and the deep
plantar aponeurosis fused with the aponeurosis of the M. flexor digitorum
fibularis and shared its insertional tendons. Probably due to the fusion of these
two aponeuroses the Mm. lumbricales have been crowded from the aponeurosis
of the M. flexor digitorum fibularis onto the ventral surface of the deep plantar
aponeurosis.
Action.—Flexion of the digits of the foot. The fusion of the tendons of
the M. plantaris and the M. flexor digitorum fibularis indicates that these
muscles normally work together. Judging by the size of these muscles and
their tendons, together they form the strongest functional unit in the shank.
The muscles enable the bat’s claws to grip the substrate.
M. popliteus (tibial nerve)
Oricin.—On medial surface of head of fibula.
INsERTION.—Along proximal half of posterolateral surface of tibia.
112 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
ReMarKs.—In Eumops the muscle is thick and fleshy proximally, becoming
thin and fibrous distally. It is represented in Myotis by a thin strand of muscle
extending from the posterior surface of the lateral condyle of the tibia to the
posterior surface of the tibia immediately distal to the head. In Macrotus the
origin is from the posterior and medial surfaces of the vestigial head of the
fibula; the insertion is along the proximal eighth of the posterior surface of the
tibia. The muscle is short and thick in this genus and fills most of the con-
cavity in the posterior surface of the tibia immediately distal to the head.
Action.—This muscle binds together the heads of the tibia and fibula and
braces the proximal parts of these bones. The muscle is vestigial and function-
less in Myotis. In Macrotus the muscle pulls the small head of the fibula
caudad and mediad and helps brace the knee joint.
M. tibialis posterior (tibial nerve)
Oricin.—From along distal two thirds of medial surface of fibula, and from
medial surface of interosseus membrane and anterior surface of M. flexor
digitorum fibularis.
INsERTION.—On ventral surface of tarsus.
RemMarks.—In Eumops the tendon of this muscle spreads out on the ventral
surface of the navicular, is bound to it by fascia, and sends flat tendinous
bands to the distal part of the ventral surface of the calcaneus, and the ventral
surfaces of the cuboid and internal cuneiform. In Myotis the muscle lies
opposite the distal third of the shank, and takes origin from the posterior
surface of the tibia, the anteromedial surface of the M. flexor digitorum
fibularis, and the medial surface of the M. peroneus longus; the insertion
closely resembles that in Eumops. The muscle originates in Macrotus along
the distal three quarters of the posterolateral surface of the tibia and the
adjacent medial surface of the M. flexor digitorum fibularis. The insertion
is on the ventral surface of the navicular.
Action.—Extension of the foot.
M. flexor digitorum fibularis (tibial nerve )
Oricin.—Along all but distalmost 3 mm. of posteromedial surface of fibula.
INSERTION.—On ventral surfaces of distal phalanges of digits one to five.
ReMArRKS.—The differences between the insertions of this muscle in the
three genera of bats under study are described in the account of the M.
plantaris. The origin in Myotis is on the lateral condyle of the tibia and along
the entire posterior surface of the fibula. In Macrotus the muscle originates
on the lateral surface of the vestigial head of the fibula and on the posterior
surface of the shaft of the fibula.
Action.—Flexion of the digits of the foot.
Flexor Group of Pes
M. flexor digitorum brevis (tibial nerve)
Oricin.—From dorsal surface of proximal end of calcaneus.
INSERTION.—On lateral surfaces of second phalanges of digits two to four.
Remarks.—The tendon of each slip divides at the level of the proximal
end of the first phalanx and sends a tendon on either side of the insertional
FUNCTIONAL MORPHOLOGY OF THREE BATS Ss
tendon used by the M. plantaris and the M. flexor digitorum fibularis. In all
three genera the latter is a thin muscle having slender and delicate tendons.
In Macrotus there is an extra slip from which a tendon extends to the fifth
digit. The unusual topographic relationships of this muscle are discussed
under the M. plantaris.
Action.—Flexion of digits two to four in Eumops and Myotis and digits
two to five in Macrotus.
M. abductor hallucis brevis (tibial nerve )
Oricin.—From distal edge of medial tarsal bone.
INSERTION.—On medial sesamoid bone of first metatarsophalangeal joint.
ReMARKS.—The medial tarsal bone is much larger and this muscle has a
more extensive origin in Eumops than in the other genera. Otherwise this
entirely fleshy muscle has similar attachments in all three genera.
Action.—Abduction of the hallux. This muscle and the M. abductor digiti
quinti spread out the foot by abducting the medial and lateral digits. Photo-
graphs of Eumops running show that the medial and lateral digits are abducted
while the foot is on the ground; this probably gives the foot a firmer purchase
on the substrate.
M. abductor ossis metatarsi quinti (tibial nerve)
Oricin.—F rom middle of ventrolateral surface of calcaneus.
INSERTION.—On lateral edge of plantar aponeurosis.
RemMarks.—This muscle usually lies deep in the tendon of the M. plantaris
and inserts on the fifth metatarsal. In the bats considered here, however, the
muscle passes mediad, ventral to the tendon of the M. flexor digitorum fibularis,
to insert on the side of the plantar aponeurosis. My identification of this muscle
is tentative.
Action.—This muscle pulls the plantar aponeurosis laterad and proximad.
In all three genera when the foot is relaxed it is not in line with the plane of
movement of the shank, but is turned inward. The tendons of the Mm. plan-
taris and flexor digitorum fibularis must, therefore, turn mediad at the level of
the base of the tarsus. In Eumops, in which the inturning of the foot is most
pronounced, the angle of the turn is approximately 45 degrees. When these
muscles contract, the tendons tend to straighten, pulling the foot back toward
an alignment with the plane of action of the shank and forcing the tendons
toward the medial edge of the tarsus. Contraction of the M. abductor ossis
metatarsi quinti reduces the latter force by pulling the tendons of the Mm.
plantaris and flexor digitorum fibularis toward the lateral side of the foot, thus
effecting some straightening of these tendons. In addition, by pulling the
plantar aponeurosis proximad this muscle helps flex the digits.
M. abductor digiti quinti (tibial nerve)
Oricin.—From ventral surface of distal part of calcaneus and from ventral
surface of lateral half of expanded base of fifth metatarsal.
InNsERTION.—On lateral sesamoid bone of fifth metatarsophalangeal joint.
Remarks.—This muscle has fleshy attachments and similar topographic
relationships in all three genera.
Action.—Abduction of the fifth digit.
114 UNIversity OF Kansas Pusts., Mus. Nat. Hist.
M. adductor hallucis (tibial nerve)
Oricix.—From ventral surfaces of bases of second and third metatarsals
and raphe between this muscle and M. adductor digiti quinti.
INsERTION.—On lateral surface of base of first phalanx of first digit and
lateral sesamoid bone of first metatarsophalangeal joint.
Remarxs.—This muscle shows only minor variation between the three
genera studied.
Action.—Adduction of the hallux.
M. adductor digiti quinti (tibial nerve)
Oricin.—From ventral surfaces of bases of metatarsals two and three and
from raphe between this muscle and M. adductor hallucis.
InsERTION.—On medial surface of base of first phalanx of fifth digit and
medial sesamoid of fifth metatarsophalangeal joint.
Remarxs.—This muscle is essentially the same in all three genera.
Action.—Adduction of the fifth digit.
Mm. lumbricales (tibial nerve)
Oricix.—From the ventral surface of the plantar aponeurosis.
InsERTION.—On dorsal surfaces of second phalanges of digits one to five.
Remarxs.—There are nine of these muscles. They send a tendon spiraling
dorsad around each side of the bases of the first phalanges of digits two to
five, and around the medial surface of the first digit. In Myotis these muscles
originate between the bases of the insertional tendons that extend distad from
the plantar aponeurosis. These muscles are relatively more slender in Macrotus
than in the other genera, and have similar attachments to those in Eumops.
Acrion.—Flexion of the digits of the foot.
Mm. interossei (tibial nerve)
Oricin.—By ten slips, two on each digit, one from lateral and one from
medial surface of bases of metatarsals one to five.
InsERTION.—On lateral and medial base of first phalanx of each digit and
lateral and medial sesamoid bones of each metatarsophalangeal joint.
Remarxs.—These muscles are relatively largest in Eumops. Their relation-
ships are similar in all three genera.
Action.—Flexion of the digits.
CONCLUSIONS
Adaptations for Flight
The remarks below, based on the bats under consideration,
probably apply at least to the microchiropteran families Molos-
sidae, Vespertilionidae and Phyllostomidae.
Many types of flying animals have tended to develop body shapes
that are aerodynamically favorable, bodies that neither create ex-
cessive amounts of drag nor counteract the lift developed by the
flight surfaces. The short, dorsally arched and moderately flat
FUNCTIONAL MORPHOLOGY OF THREE BATS 115
bodies of bats achieve this end surprisingly well. The occipital
region of the head lies close to the inter-scapular region in bats
and the head and body together are somewhat teardrop-shaped.
This body form does not create undue amounts of drag while
the bat is in flight. Because of its rounded dorsal surface and
more nearly flat ventral surface, the body forms, together with the
uropatagium, an airfoil from which some lift is obtained during
flight. Because the bodies of most bats are more nearly flat than
those of most birds, the bodies of bats are probably more effective
as airfoils; that is to say, they develop relatively greater amounts
of lift.
A tendency toward inflexibility of the vertebral column is evi-
dent in both bats and birds, being more strongly developed in the
latter. A strong base to which the wings and flight muscles can
attach is important in both of these groups. Some rigidity of the
vertebral column is necessary in bats because powerful flight
muscles that require strong, steady surfaces for attachment originate
on the thoracic vertebrae and on the ribs. Furthermore, dor-
soventral and lateral movements of the vertebral column while a
bat is in flight affect the drag and lift created by the animal’s body,
the camber of the plagiopatagium, the tautness of this membrane,
and to some extent the position of the uropatagium. During flight
a bat maintains a fairly high energy output, and it is of advantage
to the animal that the rigidity of the vertebral column is primarily
the result of the structure of the vertebrae and not of muscular
effort.
For the sake of aerodynamic stability of the animals and in-
creased manageability of their appendages, there has been a trend
in both bats and birds toward concentration of weight near the
center of gravity. Various parts of the skeletal and muscular sys-
tems of the bats here considered have become specialized so as to
effect a remarkable division of labor between the muscles, and at
the same time to concentrate the responsibility for the most de-
manding jobs on the large muscles near the center of gravity. A
locking device between articulating limb bones also commonly
serves these ends in bats.
Several interesting examples are worthy of mention. The lock-
ing of the supraglenoid tuberosity of the scapula into the depres-
sion in the proximal end of the humerus tends to stop the extension
of the humerus by relaying the force of the extension to the scapula.
This transfers the burden of stopping the extension to the group
116 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist.
of powerful muscles that bind the scapula to the body and at the
same time transfers the burden away from the relatively small
M. teres major, M. latissimus dorsi, and the M. spinodeltoideus.
During flight, when the M. spinodeltoideus is helping to elevate
the wing during each upstroke, this muscle is probably relieved of
the job of stopping the extension of the humerus and thus is given
more time for rest between contractions. This extra recovery
period probably is important when the wings are beating rapidly
for fairly long periods of time. The locking of the greater tuberosity
of the humerus against the scapula has been mentioned; this allows
a longer recovery period for the pectoral muscles by making the
posterior division of the M. serratus anterior responsible for stop-
ping the upstroke, and probably also for starting the downstroke.
Locking arrangements between bones occur also between the
spinous process of the medial epicondyle of the humerus and the
proximomedial part of the radius, between the distal epiphysis
of the radius and the lunar, and between the trapezoid and the
base of the second metacarpal. These locking devices all tend to
stop mechanically the extension of some segment of the wing and
transfer the force of that extension proximad to the heavy muscles
attaching the scapula to the axial skeleton. This transfer obviates
the need for distally situated musculature to stop the extension
movement and also concentrates the weight of the animal near the
center of gravity. Certain specializations of some of the forearm
muscles also serve to achieve this weight concentration by making
extension and flexion of the manus occur automatically with the
corresponding movements of the forearm. Because their origins
are slightly proximal to the lateral epicondyle of the humerus,
the thin, largely tendinous Mm. extensor carpi radialis longus
and brevis are stretched when the humerus is extended, thereby
extending the digits. Much of the power necessary for the ex-
tension of the manus is thus transferred proximad to the triceps
muscles. In Eumops and Myotis the origin of the M. flexor carpi
ulnaris is on the long spinous process of the medial epicondyle
of the humerus; flexion of the forearm by the M. biceps brachii
stretches the M. flexor carpi ulnaris and automatically causes
posterior flexion of the manus. In the course of the wing-beat
cycle the manus is extended at the top of the upstroke and held
rigid against the force of the airstream throughout the downstroke,
and then is flexed at the start of the upstroke; most of the power
necessary for the mentioned sequence of action probably is supplied
FUNCTIONAL MorPHOLOGY OF THREE BATS na by
in the Molossidae and the Vespertilionidae by the Mm. triceps
brachii and biceps brachii rather than by the muscles situated
in the forearm. The limiting of the planes of movement at certain
joints also serves to increase mechanical efficiency and concentrate
the weight. The humeroradial, radiocarpal and carpometacarpal
joints all limit movement in the anteroposterior plane. This limi-
tation of movement enables the control of rotational stability,
adduction and abduction, to be concentrated in the centrally lo-
cated muscles that act on the humerus or proximal end of the
radius.
Bats and birds have obviously met many of the problems of flight
in different ways. A striking example of different specializations
serving parallel functions is afforded by the muscles that control
the wings. The trend in birds has been to concentrate all of the
power for beating the wings in the pectoralis muscles, the pectoralis
major causing the downstroke, and the pectoralis minor, the up-
stroke. The pectoral girdle in birds serves as a rigid base for the
wings. The claviculae and coracoids are braced against the sternum
and the scapulae are long, thin, bladelike structures that lie nearly
immovably against the ribcage. Almost the reverse of this situation
occurs in bats, in which an intricate division of labor has evolved
between a number of muscles that control the wingbeat. Operation
of this system is based upon the retention of a large, broad scapula,
and a pectoral girdle that is nowhere attached immovably to the
axial skeleton and that retains considerable mobility. The down-
stroke is controlled mostly by the M. pectoralis, the posterior divi-
sion of the M. serratus anterior, and the M. subscapularis. The
coracoid head of the M. biceps brachii also helps slightly in the
downstroke in Myotis and Macrotus; in Eumops (and in all of the
other genera of molossid bats that I have examined) this muscle
is large and because of specializations of the coracoid process of the
scapula helps considerably in the downstroke. The division of
labor is possible because the scapula is moveable and has a large
surface area. The scapula has become faceted; therefore its surface
area is increased and it can provide origin for large flight muscles.
Furthermore, the scapula is free to rock on its anteroposterior axis;
thus, when the humerus is locked against the scapula at the top of
the upstroke the anterior division of the M. serratus anterior can
help in the wing-beat cycle by tipping the lateral edge of the scap-
ula ventrad. The upstroke is powered by dorsally situated muscles,
including the trapezius group, the deltoideus group, and the large
118 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
M. infraspinatus. Whereas the control of the wings is seemingly
as effective in bats as in birds, the major flight muscles in bats are
not all ventrally situated; hence, in terms of stability during flight,
the weight distribution in bats is not so favorable as that in birds.
To summarize: birds have a strongly braced and rigid pectoral
girdle and the pectoralis muscles provide the power for both the
upstroke and the downstroke of the wings; bats have a moveable
scapula, and a group of muscles share the burden of controlling the
wing-beat.
When the animals are in flight the actions of the large muscles
controlling the wings seem to control the respiratory cycle in both
bats and birds. During flight this is probably of major importance
in connection with meeting the increased metabolic demands of the
animals, and tends to put to use the force exerted on the ribcage by
some of the powerful flight muscles. Because certain muscles that
control the wings change the volume of the thoracic cavity in birds
.it is thought that the wing-beat and respiratory cycles in these
animals are synchronized (Krogh, 1941, Zimmer, 1935). Authors
have disagreed as to whether inspiration occurred during the up-
stroke and expiration during the downstroke or vice versa, but
experimental work clearly has shown that, at least in flights of short
duration in the domestic pigeon (Columba livia), inspiration is
synchronous with the upstroke and expiration synchronous with
the downstroke (Tomlinson and Kinnon, 1957). Judging by the
hinging of the sternum and the actions of certain major flight
muscles, an analogous situation probably obtains in bats. During
the upstroke of the wings in bats the M. pectoralis and the posterior
division of the M. serratus anterior are relaxed and the humerus is
elevated by the abductors of the brachium. As the humerus rises
the pectoralis muscle is stretched and tends to pull the post-manu-
brial part of the sternum dorsad and reduce the volume of the
thoracic cavity. Towards the top of the upstroke, when the humerus
locks against the scapula, the axillary border of the scapula is pulled
upward and the posterior division of the serratus anterior is
stretched, tending to pull the ribs dorsad and craniad, push the
posterior segment of the sternum ventrad, and increase the volume
of the thoracic cavity. When that muscle and the M. pectoralis are
both stretched their actions on the ribcage partly counteract each
other; but when the axillary edge of the scapula tips sharply upward
at the top of the upstroke the effect of the serratus muscle probably
overrides that of the pectoralis muscles and the ribcage expands.
FUNCTIONAL MORPHOLOGY OF THREE BATS 119
At the top of the upstroke, if the posterior division of the M. serratus
anterior contracts while the M. pectoralis is still relaxed (as may
occur in the course of the usual wing-beat cycle) the force tending
to expand the ribcage would increase further. After the downstroke
is begun and the pectoralis muscles contract, the volume of the
thoracic cavity is reduced by the upward force exerted against the
post-manubrial part of the sternum and the direct pressure of the
contracting muscles against the ribcage. Therefore, during the
upper part of the upstroke and the start of the downstroke inspira-
tion probably takes place, and expiration probably occurs during
most of the downstroke. Without experimental evidence the exact
time of inspiration in relation to the wing-beat cycle in bats can not
be stated with certainty. The anatomical evidence strongly sug-
gests, however, that in bats as in birds the rate of the wing-beat and
the respiratory rate are in synchrony, and that expiration is simul-
taneous with the downstroke.
The hind limbs of bats, unlike those of birds, enter into the
mechanical arrangement for flight by anchoring both the wing and
interfemoral membranes. In order to function effectively in this
capacity the hind limbs have a strikingly reptilian posture, in which
the femur extends laterad at roughly right angles to the body.
Owing to this posture, the range of movement of which the hind
limbs are capable allows them to assume a position during flight that
stretches the uropatagium and holds it in approximately the position
of a bird’s tail, whereas when bats crawl the shank is brought down-
ward and forward and is oriented almost vertically to the sub-
strate. The odd postures of the hind limbs of bats seem to be the
result of the importance of these appendages in both aerial and
terrestrial locomotion.
The Mechanics of Bat Flight
High-speed photographs show that the wing-beat cycle in bats
closely resembles that in birds in terms of the movements performed.
Many of those movements in bats were described by Eisentraut
(1936). The downstroke is the power stroke; the wings are fully
extended throughout the stroke and the direction of movement is
downward and forward. The upstroke is the recovery stroke; the
wings are partly folded during this phase, and the stroke is directed
upward and backward. The deflection of the wings away from a
direct up and down movement is due in part to the force of the air
pressure against the wing membranes. The proximal segment of
120 UNIVERSITY OF Kansas PuBLs., Mus. Nat. Hist.
the wing supplies most of the lift and the distal segment supplies
the thrust developed by the wing-beat. Flight in bats and birds,
then, is accomplished in nearly the same way; but the means by
which the movements are achieved differ greatly.
At the beginning of the upstroke in bats the adductors of the
humerus and the flexors and extensors of the radius relax and the
forearm flexes slightly as a result of the tonus of the Mm. biceps
brachii and triceps brachii, the automatic flexion of the manus by
the stretched M. flexor carpi ulnaris (this automatic flexion does
not occur in Macrotus), and the force of the air stream. Con-
traction of the abductor muscles of the brachium then raises and
flexes the humerus while the distal parts of the limb are relaxed.
The medial edge of the scapula is tipped ventrad by the trapezius
muscles, and the humerus is elevated by the deltoideus group and
the large M. infraspinatus muscle. The insertions of the deltoideus
muscles and the M. infraspinatus are so situated that as they raise
the wings they tend to rotate the leading edge upward and keep
the distal part of the wing at a fairly high angle of attack. Accord-
ingly, the wing surfaces produce some lift during this part of the
cycle, and the airstream helps to lift the wing. The convex dorsal
surfaces of the wings, and their partial flexion during the upstroke,
reduce the drag created during the upstroke (Fig. 2, Pl. 3). The
upstroke in the three bats studied by me is more rapid than the
downstroke, as noted by Eisentraut (op. cit.) in Myotis myotis,
Rhinolophus hipposideros and Plecotus auritus, and by Orr (1954:
206) in Antrozous pallidus. Although in terms of energy output
the upstroke is less efficient in bats than in birds, the upstroke in
bats probably requires only a fraction of the power demanded by
the downstroke.
At the top of the upstroke the greater tuberosity of the humerus
contacts the scapula and the force of the upstroke is transferred to
this element. Because the vertebral border of the scapula has
been tipped ventrad by the action of the trapezius muscles, the
broad, thick posterior division of the M. serratus anterior (that
inserts on the axillary border of the scapula) is stretched by the
stage in the cycle at which the humerus locks against the scapula.
The elastic effect of the tonus of the posterior division of the M.
serratus anterior may stop the upstroke; or this muscle may con-
tract as the humerus locks, thus stopping the stroke. In either
event, contraction of this muscle, with the consequent pulling
downward of the axillary border of the scapula and the resulting
.
FUNCTIONAL MORPHOLOGY OF THREE BATS UPA
adduction of the locked humerus, probably initiates the downstroke,
with the help of the tonus of the stretched M. pectoralis. When
the downstroke is started, the pectoralis muscles contract. Al-
though it supplies the largest share of the power necessary for
the downstroke, the M. pectoralis has considerable help in this
action. The posterior division of the M. serratus anterior relieves
the M. pectoralis of the burden of actively stopping the upstroke
and starting the downstroke and assists the M. pectoralis through
the upper part of the downstroke, while the humerus is still locked
against the scapula. The serratus muscle can not act on the
humerus when the greater tuberosity of the humerus loses contact
with the scapula, but throughout the downstroke the M. pectoralis
is helped by the large M. subscapularis (and in Eumops, and
probably in all molossid bats, by the action of the large coracoid
head of the M. biceps brachii). At the start of the downstroke
the M. subscapularis and the Mm. triceps brachii and biceps
brachii extend the humerus and the forearm. Extension of the
forearm stretches the largely tendinous M. extensor carpi radialis
longus and the M. extensor carpi radialis brevis, and the meta-
carpals are automatically extended. The digital extensors, with the
aid of the force of air pressure beneath the chiropatagium, com-
plete the extension of the chiropatagium. Thus, concurrent with
the start of the downstroke, the forelimb is fully extended and the
wing membranes are spread. Anteroposterior rigidity of the
forearm and manus during the downstroke is controlled largely
by the antagonistic actions of the flexors and extensors of the fore-
arm, the Mm. biceps brachii and triceps brachii; rotational stability
of the forelimb is maintained by the antagonistic actions of the
M. pectoralis and the M. biceps brachii.
The camber and angle of attack of the distal segment of the
wing is under the control of the rotators of the humerus, the
flexors of the phalanges of digits three to five, and the angle of
the dactylopatagium minus; but owing to the elasticity of the
membranes, the flexibility of the bones of the digits, and the
“give” that occurs at the interphalangeal joints, the camber is also
controlled by changing air pressures. The distal segment of the
wing supplies thrust during the downstroke, and to perform this
function the distal flight surfaces must be able to have a different
angle of attack during the downstroke than during the upstroke.
The leading edge of the distal segment of the wing is formed by
the second and third digits, and serves as a relatively rigid support
r27, UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
for the leading edge of the wing. The trailing edge of the wing
has no such continuous support, and the force of air pressure dur-
ing the downstroke tends to push the trailing edge and posterior
parts of the chiropatagium upward and overcome the slight ventral
flexion of the phalanges of the fourth digit. Because the distal
parts of the chiropatagium are less strongly braced and are sub-
jected to greater air pressures than the proximal parts, the “pitch”
of the chiropatagium (like the pitch of some airplane propellers)
increases distally, and the greatest part of the forward thrust seems
to be supplied by the section of the chiropatagium between the
third and the fourth digits.
In the proximal segment of the wing, the camber and angle of
attack are influenced by the positions of the propatagium, the fifth
digit, and the hind limbs. The humerus and radius form a relatively
inflexible brace for the anterior edge of the proximal segment of the
wing, whereas the trailing edge of this segment is unsupported, and
its rigidity depends upon its being tautly stretched between the fifth
digit and the hind limb. When the wing is extended the chiro-
patagium is pulled forward and held taut by the M. occipitopolli-
calis, and increases the area and camber of the proximal segment
of the wing. The angle that the propatagium makes with the
plagiopatagium seems to be fairly constant in a given species, but
can be varied somewhat by flexion or extension of the thumb. Be-
cause the proximal segment of the wing produces most of the lift
supplied by the flight membranes, that segment must retain roughly
the same angle of attack throughout the downstroke in order to
develop the maximum lift; this demands that the fifth digit be braced
so as to withstand the force of air pressure on the underside of the
wing during the downstroke phase of the wing-beat cycle. A num-
ber of remarkable specializations serve to make the fifth digit a
relatively rigid anchor for the distal edge of the plagiopatagium.
The fifth metacarpal is strongly built and is curved more strongly
ventrad than are the other metacarpals (in Eumops it is laterally
compressed). In addition, the fifth metacarpal is braced against
upward pressures by the pisiform bone and the M. abductor pollicis
longus and M. abductor digiti quinti. The hind limb forms the
proximal anchor for the posterior part of the plagiopatagium, and
is steadied during flight largely by the adductors and abductors
of the femur and the flexors of the shank.
The hind limbs not only serve as proximal anchors for the plagio-
patagium but govern the angle of attack of the uropatagium. The
FUNCTIONAL MORPHOLOGY OF THREE BATS 123
position of the uropatagium has an important effect on the equilib-
rium of the body during flight. A slight raising (abduction) of the
hind limbs with a consequent lifting of the uropatagium and tail
attends the partial flexion of the forelimb and the beginning of the
upstroke. Elevation of the uropatagium and tail continues through-
out the upstroke of the wing; at the start of the downstroke, when
the wings are fully extended, the hind limbs and uropatagium are
higher than they are at any other part of the cycle. This up-and-
down movement of the hind limbs compensates for the raising and
lowering of the anterior parts of the wing membranes and tends
to keep the angle of attack of these membranes fairly constant
throughout the entire cycle. The proximal part of the anterior
portion of the plagiopatagium is raised and lowered in relation to
the body as the wings are raised and lowered. If the hind limbs,
the places of anchorage of the proximal part of the posterior section
of the plagiopatagium, were to be held in the same position relative
to the body throughout the wing-beat cycle, the angle of attack
of the plagiopatagium in parts of the cycle would be such that this
membrane would not function efficiently as a lifting surface. With
the uropatagium moving up and down in synchrony with the strokes
of the wings, however, the angle of attack remains roughly the same
throughout the wing-beat cycle. The dorsal movement of the hind
limbs tends also to keep the body horizontally oriented during the
wing-beat cycle by compensating for the changing position of the
wings, which move from in front of the center of gravity of the
animal at the bottom of the downstroke to behind the center of
gravity at the top of the upstroke. The raising of the hind limbs
during the upstroke pulls the uropatagium upward so much that it
is at a low angle of attack and is developing but little lift. The
posterior part of the animal is thus supported to a lesser extent
than it is when the uropatagium angles downward, and the posterior
part of the body tends to drop. At the same time that the uro-
patagium is developing a minimum of lift, the wings are extended
at their posteriormost position, well behind the center of gravity
of the body, and the anterior end of the body tends to “topple”
frontward. As a result of the interplay of these two tendencies,
the body stays on a fairly even keel. In the downstroke the function
of the uropatagium is the same. As the wings are adducted in the
downstroke the hind limbs and uropatagium are lowered until at
the bottom of the stroke the uropatagium is at the highest angle
of attack it attains in the wing-beat cycle. Again, the two important
/
124 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
functions—maintenance of a fairly constant angle of attack of the
plagiopatagium and compensation for the changes of position of
the lifting surfaces relative to the center of gravity—are performed.
The hind limbs pull the trailing edge of the plagiopatagium ventrad
as the leading edge is moved ventrad by the forelimb, and the angle
of attack of the plagiopatagium remains nearly the same throughout
the stroke. When the wings are near the bottom of the downstroke
the uropatagium is at its greatest angle of attack and is developing
maximum lift, tending to lift the posterior part of the body. The
wings, at this point, are anterior to the center of gravity of the body
and are tending to tip the anterior part of the body upward. This
lifting of the anterior part of the body by the wings, then, is counter-
acted by the lifting of the posterior end of the body by the de-
pressed uropatagium; accordingly, the body stays nearly horizontally
oriented.
Orr (1954:206), studying Antrozous pallidus, noticed that the
uropatagium was lowered with the downstroke and raised during
the upstroke. He incorrectly, I think, concluded that the lift gained
by the lowered uropatagium during the downstroke compensated
for the loss of lift that occurs with the next upstroke.
In level flight the dorsal and ventral movements that the hind
limbs, uropatagium and tail perform during the wing-beat cycle
are clearly not the result of the legs being pulled up and down by
intermittent contractions of the femoral abductors and adductors.
The hind limbs are held in their flight-position by the tonus, or by
weak contractions of the muscles of these appendages, and instead
of moving the hind limbs with each wing stroke the muscles tend
to hold the limbs steady. The tail also is held in a fairly constant
position. As the wing is elevated and lowered, however, the amount
and direction of the pull exerted by the wing-membranes on the
limb varies, due to the variation in the dorsoventral position of the
wings, the force of air pressure against the membranes, and the
degree of extension of the wing. Accordingly, the force of the pull
on the hind limbs varies in the upstroke from a downward pull
during the lower part, to a weak lateral pull during the middle part,
to a dorsal pull during the upper part, and the hind limbs and
uropatagium are pulled progressively higher throughout the stroke
by the wing membranes. In the downstroke the pull on the hind
limbs changes from a strong dorsolateral pull during the upper
part of the stroke, to a strong lateral pull during the middle part,
to a strong ventrolateral pull during the lower part, and the hind
FUNCTIONAL MORPHOLOGY OF THREE Bats 125
limbs are forced progressively lower during the stroke. The dif-
ferences in the positions of the hind limbs, uropatagium and tail at
the different points of the wing-beat cycle, then, are the result of
the changes in the strength and direction of pull of the wing mem-
branes acting against the muscles of the hind limb that tend to hold
the limbs rigid. In deviations from level flight, of course, the
muscles of the hind limbs often effect changes in the position of
the uropatagium.
Comparisons of the Bats Studied
It is worth-while to re-examine some of the differences between
the three bats under consideration with the aim of learning which
modifications are more efficient adaptations to the basic chiropteran
mode of life. The kind of character here called “advanced” is one
that makes the bat a more effective flying animal. Modifications
that enable bats to occupy special environmental niches are re-
garded as specializations, whereas modifications that enable the
animals to fly more efficiently, no matter what particular type of
flight they use, are termed advanced characters.
The obvious differences between the sterna of the three genera
under study seem in part to be the result of the roosting habits of
these animals. Relative to the length of the thoracolumbar section
of the vertebral column, the sternum is longest in Eumops and
shortest in Macrotus. As an additional contrast, the sternum of
Eumops has almost no keel, and that of Macrotus bears a large
keel. The crevice roosting habit of Eumops makes an increase in
the depth (dorsoventral thickness) of the body disadvantageous,
yet the strong flight characteristic of this genus demands powerful
musculature. Thus, the long, unkeeled sternum serves to give a
large surface for the attachment of the pectoralis muscles without
increasing greatly the depth of the body. Because Macrotus roosts
exclusively by hanging, there probably is no selective pressure
exerted against the development of a deep body and it is under-
standable that the area needed for the attachment of the pectoral
muscles is provided by the short, keeled sternum. In Eumops, be-
cause of the shape of the sternum, the pectoralis muscles have a
long, thin origin. In body shape Myotis is roughly intermediate
between Eumops and Macrotus. Myotis has a short deep manu-
brium and the posterior part of the sternum has a low ridge. This
genus roosts both by hanging, in company with other individuals
of its kind, and by wedging itself into small holes or fissures. Per-
126 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
haps because of its small size and more generalized roosting habits,
a shallow body is not so important in Myotis as in Eumops. In any
case, the shape of the sternum seems not to be a character clearly
indicative of degree of advancement in bats, and is strongly in-
fluenced by the roosting habits of the animals.
The differences between the scapulae of the three bats may also
result in part from differences in roosting habits. Probably because
there has been less selective pressure against the development of
a deep thorax in bats that roost by hanging than in crevice-dwelling
bats, Macrotus has developed the M. pectoralis to an extent not
found in the other genera. As a result, there has been no need for
the perfection of an effective division of labor between several
flight muscles to limit the size of the M. pectoralis in Macrotus, and
the scapula has not been so specialized to serve as a place of origin
for some of the auxiliary flight muscles in Macrotus as it has in
the other genera. In the crevice-dwelling Eumops dorsoventral
thinness would seem to be advantageous and the scapula is highly
specialized to give origin to the muscles aiding the M. pectoralis
in the downstroke. Such characters as the elongation of the scapula,
sharp downward angle of its supraspinous part, large anterior flange
on the anterior rim of the supraspinous fossa, and medially pro-
jecting coracoid process indicate this adaptation to provide larger
area for insertion of muscles aiding the pectoralis muscles. Myotis
has some of these specializations of the scapula (for example large
anterior flange on rim of supraspinous fossa) but in general is not
so specialized in this regard as Eumops. Although specialization
of the scapula is characteristic of most groups of the suborder Mi-
crochiroptera, selective pressure influencing development of a spe-
cialized scapula varies according to roosting habit and kind of
flight of the bats. Nonetheless, because specializations of the scap-
ula in bats seem to indicate a more efficient utilization of the
mechanical scheme for flight based upon a movable scapula and
heavy musculature attaching this element to the axial skeleton,
these specializations are regarded as advanced characters.
Crevice-dwelling seems to favor the specialization of the humero-
scapular locking device. A firm lock is one detail of the chiropteran
shoulder joint seemingly highly efficient for flight. Accordingly,
enlargement of the greater tuberosity of the humerus and conse-
quent locking of the humeroscapular articulation at a lower point
in the upstroke is considered as an indication of advancement.
A few other characters indicative of advancement in bats can be
FUNCTIONAL MORPHOLOGY OF THREE Bats 127.
enumerated. Limitation of movement to the anteroposterior plane
at the elbow, wrist and carpometacarpal joints is necessary for the
limb to serve as an effective framework for the flight surfaces. Thus,
modifications that tend to brace these joints are advanced char-
acters. Centralization of weight is another important trend toward
aerodynamic efficiency. The locking devices that limit the ex-
tension of the segments of the forelimb, and the specializations of
the Mm. extensor carpi radialis longus and brevis and the M. flexor
carpi ulnaris that allow these muscles to operate as elastic cords that
automatically move the manus with movements of the forearm serve
this end, and are therefore advanced characters. The fifth digit,
together with the hind limbs, has the important function of con-
trolling the angle of attack of the plagiopatagium, which is the
main lifting surface of the flight membranes. Specializations that
strengthen the fifth metacarpal and modifications that enable other
bones or muscles to brace the fifth digit, then, represent advanced
characters.
Eumops has developed all of the advanced characters mentioned
above to the highest degree of any of the three genera under study.
Myotis exhibits many of these characters, but they are less perfectly
developed than in Eumops. Macrotus is the least advanced. Its
scapula, proximal end of the humerus and joints of the forelimbs
are far less advanced than those of the other two genera. The M.
flexor carpi ulnaris does not serve to flex the manus automatically
when the forearm is flexed as this muscle does in the other genera.
The fifth digit is slightly more strongly braced in Macrotus than in
Myotis, but is not nearly so well braced as in Eumops. On the basis
of the few post-cranial characters assumed to indicate advancement
within the suborder Microchiroptera, Eumops (Molossidae) is
judged to be the most advanced and Macrotus (Phyllostomidae)
is the least advanced; Myotis (Vespertilionidae) is intermediate.
I have examined additional members of each of the families repre-
sented by these bats, and judge that with respect to advancement
the families should be listed as above. This is in agreement with
Miller's (1907) currently accepted arrangement of these families.
Evolutionary Considerations
The fossil record of bats is scanty, and the stages represented by
the record seem to be well above the level at which the bats di-
verged from their ancestral stock. The earliest undoubted chirop-
teran fossils are from the middle Eocene of Europe and North
America, but even those bats were not extremely primitive; already
128 UnIversITY OF Kansas Pusts., Mus. Nat. Hist.
the basic chiropteran morphological pattern was established and
certain refinements are evident. In Palaeochiropteryx from the
middle Eocene of Europe, for example, the calcar is present, and
in the manus the pollex bears the only ungual phalanx. Bats must
have undergone their initial evolutionary development in the Paleo-
cene or earlier.
Without a background of fossil material a study of the morphology
of Recent bats can not certainly reveal their phylogeny. Neverthe-
less, the morphology of Recent bats suggests something of the early
stages of evolution of the Chiroptera.
Before considering the evolution of bats it should be stressed
that the modifications of the pectoral and pelvic girdles in these
animals have been influenced strongly by the dual use of both
the forelimbs and the hind limbs. Although aerial locomotion has
greatly affected the evolution of the appendages of bats, some type
of terrestrial locomotion in which the forelimbs are used is im-
portant in most bats. Consequently, natural selection may have
operated against anatomical modifications that tended to make the
forelimbs useless in terrestrial locomotion, even if such changes
would have made these limbs more efficient in flight; thus, the evo-
lution of the forelimbs has taken place within the limits imposed by
the use of these appendages in terrestrial locomotion. In the same
way, the evolution of the hind limbs, although influenced by the
use of these limbs in terrestrial locomotion, has been partly con-
trolled by the importance of these appendages as anchors for the
wing membranes during flight.
Judging completely on the basis of the morphology of Recent
bats, it seems that when bats were diverging from their insectivore
progenitors—when the basic adaptations for flight were being de-
veloped—bats usually sought diurnal refuge in narrow, crevicelike
retreats. Several lines of evidence support this idea.
The whole scheme of muscular and osteological specialization
that enables bats to perform the downstroke (powerstroke) of the
wings effectively may have resulted from a tendency, which prob-
ably originated due to the crevice-dwelling habits of early bats, to
avoid dorsoventral thickening of the body. In no known bat has
the M. pectoralis or the keel of the sternum developed to anything
approaching the degree seen in most birds. The division of labor
between the major flight muscles in bats has kept the size of the
pectoralis muscles at a minimum, while tending to create a broad
and moderately flat body that contrasts with a deep, fairly narrow
body of most birds. Although the bodies of bats may serve as more
FUNCTIONAL MorPHOLOGY OF THREE BATS 129
effective airfoils than those of birds, the weight distribution in bats
is not so favorable to equilibrium during flight as is that of birds.
In addition, because of the directions of the muscle pulls involved,
adaptations enabling bats to retain a flat body seem not to develop
strong muscular control of the wings so directly as is seen in birds.
Seemingly, the primary advantage of the chiropteran body-shape
is that it allows the bats to fit in small spaces. Clearly, to a crevice-
dwelling bat a deep thorax would be a decided disadvantage, for
it would greatly restrict the number of suitable roosting sites and
would make locomotion within the crevice difficult. The moderately
small pectoralis muscles, the relatively slightly keeled sternum, and
the specializations creating a division of labor between several
muscles that control the downstroke may result from bats having
perfected flight while they were restricted to roosting in crevices,
a niche that would have brought strong selective pressure to bear
against the avian method of solving the problem of the muscular
control of wings.
The posture of the hind limbs and forelimbs and the modifica-
tions of the pectoral and pelvic girdles in bats favor both the use of
the appendages to support flight surfaces (this function demanding
that the limbs be held out laterally from the body ) and the operation
of these limbs in connection with crawling within narrow spaces.
Locomotion within a crevice can be especially effectively accom-
plished by an animal the limbs of which are directed mostly laterad
instead of being oriented vertically to the substrate. In animals in
which the flight surfaces consisted of membranes stretched be-
tween the limbs, both the crevice-roosting habit and the develop-
ment of flight would have tended to force the development (or
retention) of the posture of limb typical of bats today. At the
critical early period in which flight in bats developed, the rate of
evolution conceivably was increased if both the roosting habits and
the foraging habits of the animals made the same general type of
limb posture advantageous.
A crevicelike retreat is one that was probably more nearly ubiq-
uitous during the early stages of the evolution of bats than any
other kind of shelter. Although caves are popularly considered
as the favored roosting places of bats, in many regions today where
bats are known to be abundant, both in total numbers and in di-
versity of species, there are no caves and most of the bats seek
refuge in some crevicelike retreat. The initial radiation of the
5—4357
130 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist.
chiroptera would probably have been favored by the habit of oc-
cupying crevices.
The Recent chiropteran fauna includes many species, and several
families, that roost exclusively by hanging pendant from some sup-
porting surface and that are specialized in various ways to exploit
successfully this roosting niche; characteristically these bats have
shorter bodies and deeper chests, relatively large pectoralis muscles
and less advanced scapulae and humeri than crevice-dwelling kinds.
In bats that roost in crevices and those that roost by hanging the
basic pattern of muscular control of the wings seems to be the same,
but selective pressure has seemingly not favored the perfection of
some of the major chiropteran adaptations to the extent in bats that
hang that it has in crevice-dwelling bats. The post-cranial skeletons
of certain representatives of the cave-roosting families Phyllosto-
midae and Rhinolophidae, and the mainly crevice-dwelling families
Vespertilionidae and Molossidae were examined. Without excep-
tion, the crevice-dwellers have more nearly flat bodies and more
advanced “chiropteran” adaptations for flight. Assuming that bats
underwent their early evolution when they were crevice-dwellers,
it seems that certain groups changed their roosting habits at a time
when the major bat specializations of the pectoral girdle had not
yet been perfected, and that because these animals were relieved
of the selective pressure against the development of deep thoraces
the pectoral girdle has not become so advanced (in terms of the
chiropteran type of adaptation), and that the division of labor be-
tween the flight muscles has not developed to the extent found in
present day crevice-dwellers. In any case, the differences between
the post-cranial morphology of such cave roosting groups as the
Phyllostomidae and the Rhinolophidae and the largely crevice-
dwelling Molossidae indicate that the crevice-dwelling habit favors
the development of the advanced “chiropteran” pectoral girdle.
Eumops is known to seek diurnal refuge in crevices and clearly
is an advanced bat. Its highly developed humeroscapular locking
device and the morphology of the scapula reflect an advanced
degree of development of the division of labor between the flight
muscles. These features and the long slender wing in Eumops may
be responses principally to a “need” for rapid and prolonged flight
and secondarily responses to a “need” for more efficient locomotion
within a narrow space; some of the adaptations that favor rapid
crawling within a crevice also favor the evolution of a long, high-
aspect-ratio wing. The second digit, the third and fourth meta-
FUNCTIONAL MORPHOLOGY OF THREE BATS ROL
carpals and the fifth digit are all approximately the length of the
forearm, and the phalanges of the third and fourth digits fold
accordion-fashion against their respective metacarpals. This ar-
rangement allows the wing to fold into a compact bundle that is
manageable when the bat is crawling rapidly in a crevice. Also,
this arrangement allows a long distal segment of the wing to be
developed, for the phalanges of digits three and four fold back
against the metacarpals and do not project awkwardly beyond the
proximal end of the radius when the wing is folded as in non-
molossid bats. The shallow body and fairly flat head of the mastiff
bat not only suit the bat for crevice dwelling but improve the airfoil
formed by the head, body and uropatagium. In the case of Eumops,
then, some of the animal’s most advanced characters, as well as
certain remarkable specializations, seem to have developed within
the limits imposed by the crevice-roosting habit. Conceivably
the development of these characters may actually have been
hastened by the selective pressure resulting from this habit.
It is evident that many of the major adaptations for flight differ
in bats and birds, and it seems that the basic chiropteran muscular
and osteological adaptations for flight are the result of bats having
been crevice-dwellers during the period of their early evolution.
SUMMARY
A comparative study was made of the functional morphology of
three North American bats, the western mastiff bat (Eumops pero-
tis of the family Molossidae), the cave myotis (Myotis velifer of
the family Vespertilionidae), and the leaf-nosed bat (Macrotus
californicus of the family Phyllostomidae). The myology and
osteology of the pectoral and pelvic girdles and their appendages
was described, and remarks were made on the actions of the
muscles. As a basis for functional considerations, a field study on
the foraging and roosting habits of these bats was made in the
desert and coastal regions of southern California in the summers
of 1953, 1954, and 1957. Additional information was gained from
high-speed photographs and observations of these bats in the
laboratory.
All three of the bats here considered are insectivorous, but their
roosting and foraging habits differ, as do their modes of flight and
terrestrial locomotion. Eumops seeks daytime refuge chiefly in
crevices or fissures in cliffs or large boulders, and roosting-sites are
usually high above the ground. This bat is fairly active in its roost
at various times of the day, and can crawl rapidly. The method
132 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. HIst.
by which this bat crawls is remarkably effective within the narrow
confines of a crevicelike retreat. Eumops is a speedy and enduring
flier, but is unable to take flight from a level surface and must
launch itself from at least six feet above the ground to attain
sustained flight. In summer this bat forages continuously for
roughly six and one half hours per night. Generally the nightly
exodus from the roost begins about one hour after sunset and the
bats return between one and two hours before sunrise. Mastiff
bats usually forage high above the ground; their foraging range
is at least five miles, and under certain conditions may be fifteen
miles or more.
Myotis velifer seems to prefer caves for daytime roosting, but
inhabits also buildings, and crevices and small holes in rocks. This
bat usually roosts in clusters by congregating on ceilings or in
crevices in ceilings of caves. The cave myotis can crawl fairly
rapidly, although not with the speed or agility of Eumops. The
flight of M. velifer is direct for a small bat, but when it clases
insects its flight is erratic and highly maneuverable. Often it forages
in riparian situations or over dry desert washes supporting scattered
large plants; then it usually flies six to 15 feet above the ground.
In the evening this bat generally emerges from its roost roughly
one half hour after sunset, and seems to have but one major forag-
ing period per night; each bat probably is on the wing no more
than one and a half hours per night.
The leaf-nosed bat rests in the daytime almost exclusively in
caves, and roosts by hanging from the ceiling with the body pendant
and only the feet in contact with the rock. Terrestrial locomotion
is confined to a unique kind of “walking” across a ceiling; Macrotus
is incapable of crawling. The flight is slow and extremely maneu-
verable, and the animal is able to hover for several seconds at a
time. In alighting, Macrotus performs a half roll and grasps the
ceiling with its feet. It usually forages within three feet of the
ground, and seemingly most of the prey is captured on the ground
or is picked from vegetation. There is one pre-midnight and one
early-morning foraging period, and each bat probably is on the
wing less than one hour and forty-five minutes per night.
Between the three bats studied here considerable variation occurs
in the aerodynamic characteristics of the flight surfaces, these
differences reflecting mostly the foraging habits of the animals.
Eumops has a long, narrow, high-aspect-ratio wing; the wing load-
ing is higher and the camber is lower than in the other bats. In
FUNCTIONAL MORPHOLOGY OF THREE BATS 1383
general, the wings of Eumops are well adapted for rapid, prolonged
flight. The wings of Myotis velifer are short, fairly broad and of
high camber, and are adapted to develop high lift at low speeds.
The wing loading is lower and the uropatagium is larger than in the
other two bats, these characteristics favoring maneuverability. The
wings of Macrotus californicus have approximately the same aero-
dynamic characteristics as those of Myotis velifer, but the uro-
patagium is smaller in the former. Macrotus probably owes its
maneuverable flight in large part to specialization of its sensory
equipment.
In bats the pectoral and pelvic girdles are highly modified for
support and for control of the flight membranes. In the bats here
considered the sternum of Macrotus is the only one having a keel,
but in all three bats the joint between the manubrium and the body
of the sternum allows the post-manubrial part of the sternum to
swing ventrad; this arrangement probably puts the breathing cycle
during flight partly under control of the major adductors of the
wing. The scapula is large and faceted and provides a large area
for muscle attachment. There are locking devices formed by the
articulating elements at the scapulohumeral, the humeroradial, the
radiocarpal, and the carpometacarpal joints; these joints mechani-
cally stop the extension of the various segments of the forelimb.
Because of specializations of the articular surfaces involved, flexion
and extension of the forearm and hand can occur only in the anter-
oposterior plane. Many of the osteological and myological spe-
cializations of the pectoral girdle serve to transfer responsibility
for control of the wings to muscles situated near the center of gravity
of the animal. Such adaptations tend to increase the manageability
of the forelimb and effect an advantageous weight distribution in
terms of aerodynamic stability. The hind limbs have a reptilian
posture in bats and are held more or less out to the sides of the
body; several of the largest muscles in the pelvic girdle seem to
function most importantly to steady the hind limbs and uropatagium
during flight.
Although there are parallel trends toward rigidity of the vertebral
column and centralization of weight in bats and birds, many of the
basic adaptations enabling these animals to control their flight
surfaces are different. In birds the pectoral girdle is braced rigidly
against the axial skeleton and the power for the wing-beat is sup-
plied almost entirely by the pectoralis muscles. Nearly the oppo-
site mechanical arrangement obtains in bats, for in this group the
134 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
job of controlling the wing-beat is divided between a number of
muscles, and this division of labor is possible mainly because of
a large and moveable scapula.
In chiropteran flight the proximal segment of the wing (plagio-
patagium and propatagium) supplies the lift and the distal segment
of the wing (chiropatagium) supplies the forward thrust during
the wing-beat cycle. The downstroke is directed forward and
downward and is mainly under the control of the M. pectoralis,
the anterior division of the M. serratus anterior, and the M. sub-
scapularis; the upstroke is an upward and backward movement
and is controlled principally by the trapezius and deltoideus groups
of muscles. The proximal segment of the wing is stretched between
the hind limb and the fifth digit. The hind limb is steadied mostly
by the adductors and abductors of the femur and the flexors of the
shank. As a result of a number of osteological and myological
specializations the fifth digit is braced against the force of air pres-
sure during the downstroke and maintains the plagiopatagium at a
fairly constant angle of attack during this phase of the wing-beat
cycle. The head, body and uropatagium form a crude airfoil that
develops lift during flight. The uropatagium moves up and down
with corresponding movements of the wings; during flight these
changes in the angle of attack of the uropatagium change the amount
of lift developed by this membrane and serve to keep the plagio-
patagium at a fairly constant angle of attack and compensate for
the changes of position of the wings relative to the center of gravity
of the animal.
The assemblage of muscular and osteological specializations for
flight characteristic of microchiropteran bats has resulted in these
animals having broad, moderately flat bodies. This body-form
suggests that in the time during which bats underwent their early
evolution they were crevice dwellers, and that selective pressure
operated against the development of the deep-chested body-form
typical of birds. Because the demands of flight and of locomotion
within a crevice favor the development of the kind of limb posture
occurring in most bats, it is conceivable that the rate of evolution
of these animals was increased by an early predilection for crevice-
dwelling.
FUNCTIONAL MORPHOLOGY OF THREE BATS 135
LITERATURE CITED
Cuvier, G.
1800-1805. Lecons d’anatomie comparée. Paris. 5 vols.
EIsENTRAUT, M.
1936. Beitrag zur Mechanik des Fledermausfluges. Zeitschrift f. wis-
sensch. Zoologie, 148: 159-188, 18 figs., 1 table.
FisHer, H. I.
1955. Avian anatomy, 1925-1950, and some suggested problems. Chap-
ter 4 in “Recent Advances in Avian Biology,” edited by A. Wolfson.
Univ. Illinois Press, x + 479 pp., illustrated.
GRINNELL, H. W.
1918. A synopsis of the bats of California. Univ. California Publ. Zool.,
17: 223-404, pls. 14-24, 24 figs.
Hatt, E. R.
1946. Mammals of Nevada. Univ. California Press, Berkeley and Los ©
Angeles, xi-+ 710 pp., frontispiece, colored, 11 pls., 485 figs., un-
numbered silhouettes.
HATFIELD, D. M.
1937. Notes on the behavior of the California leaf-nosed bat. Jour.
Mamm., 18: 96-97.
Hore J. E.
1937. Morphology of the pocket gopher, mammalian genus Thomomys.
Univ. California Publ. Zool., 42: 81-172, 26 figs.
Howe Lt, A. B.
1920a. Contribution to the life history of the California mastiff bat. Jour.
Mamm., 1: 11-117, 2 pls.
1920b. Some Californian experiences with bat roosts. Jour. Mamm., 1:
169-177, 1 pl.
1937. Morphogenesis of the shoulder architecture, part vi, therian Mam-
malia. Quart. Rev. Biol., 12: 440-463.
Huey, L. M.
1925. Food of the California leaf-nosed bat. Jour. Mamm., 6: 196-197.
Humenury, G. M.
1869. The myology of the limbs of Pteropus. Jour. Anat. and Physiol.,
3: 294-319, pl. 6, 7.
KNupsEN, V. O.
1931. The effect of humidity upon the absorption of sound in a room,
and a determination of the coefficients of absorption of sound in
air. Jour. Acoustical Soc. Amer., 3: 126-138, 7 figs.
19385. Atmospheric acoustics and the weather. Scientific Monthly, 40:
485-486.
KoLenatr, F. A.
1857. Beitrige zur Naturgeschichte des europdischen Chiroptern. Allge-
meine Deutsche Naturhistorische Zeitung, 3: 1-24, 41-68, figs. 1-10.
Krocu, A.
1941. The comparative physiology of respiratory mechanisms. Univ.
Pennsylvania Press, Philadelphia, vii + 172 pp., 84 illust.
136 UNIVERSITY OF KANSAS PusLs., Mus. Nat. Hist.
Krutzscu, P. H.
1955. Observations on the California mastiff bat. Jour. Mamm., 36: 407-
Al4,
LAWRENCE, B. .
1943. Miocene remains from Florida, with notes on the generic characters
of the humerus of bats. Jour. Mamm., 24: 356-369, 2 figs.
MACALISTER, A.
1872. The myology of the cheiroptera. Philos. Trans. Royal Soc. London,
162: 125-171, pls. 12-16,
Miter, G. S.
1907. The families and genera of bats. U. S. Nat. Mus. Bull., 57:
xvii + 282 pp., 14 pls., 49 figs.
Orr, R. T.
1954. Natural history of the pallid bat, Antrozous pallidus. Proc. Cali-
fornia Acad. Sci., 28: 165-246, 28 figs.
Poo eg, E. L.
1936. Relative wing ratios of bats and birds. Jour. Mamm., 17: 412-418.
RINKER, G. C.,
1954. The comparative myology of the mammalian genera Sigmodon,
Oryzomys, Neotoma, and Peromyscus (Cricetinae), with remarks
on their intergeneric relationships. Misc. Publ. Mus. Zool. Univ.
Michigan, 83: 1-124, 18 figs. 2 tables.
SAvILE, D. B. O.
1950. The flight mechanism of swifts and hummingbirds. Auk, 67: 499-
504.
1957. Adaptive evolution in the avian wing. Evolution, 11: 212-224, 9
figs., 1 table.
Sracer, K, E.
1939. Status of Myotis velifer in California, with notes on its life history.
Jour. Mamm., 20: 225-228.
1943. California leaf-nosed bat trapped by desert shrub. Jour. Mamm.,
24: 396.
ToMLInson, J. T., and McKinnon, R. S.
1957. Pigeon wing-beats synchronized with breathing. Condor, 59: 401,
fig:
TweENTE, J. W.
1955. Some aspects of habitat selection in cavern-dwelling bats. Jour.
Ecology, 36: 706-732, 14 figs., 1 table.
ZIMMER, K.
1935. Beitrige zur Mechanik der Atmung bei Vogeln in Stand und Flug.
Zool. Stuttgart, 33: 1-69, 6 pls., 88 figs.
PLATE Il
Fic. 1. View of terrain which offers suitable roosting places for Eumops
perotis. The domelike boulder of granodiorite on the lett has several crevices
in which E. perotis roosts in the daytime. Photo August 28, 1957, three miles
northeast Perris, California.
Fic. 2. Closeup of part of the boulder shown in Fig. 1. The crevice in the
lower left was usually occupied by a single Eumops perotis; the crevice
beneath the tongue-shaped slab of rock in the upper right generally harbored
three to six individuals.
PLATE 2
Fic. 1. View of an oxbow in the floodplain of the Colorado River where
Myotis velifer and Macrotus californicus forage. Such riparian situations
provide optimal foraging habitat for Myotis velifer in the Riverside Mountains
area. The strip of vegetation on the left is mostly tamarisk and screw bean;
cattails border the water and some arrowweed are in the foreground. Photo
August 22, 1957, 35 miles north and two miles east Blythe, Riverside County,
southeastern California.
Fic. 2. View looking south across a small desert wash to the Riverside
Mountains. The wash is a preferred foraging habitat of Macrotus californicus;
Myotis velifer also forages there. Photo August 21, 1957, 36 miles north
Blythe, Califormia.
Fic. 1. Macrotus californicus Fic. 2. Macrotus californicus in slow, nearly
hanging from ceiling of cave. hovering flight, with the uropatagium low-
Note that leg by which animal ered. The wings are in the middle of the
hangs is extended nearly upstroke. Note the high camber of the
straight behind the bat. Photo right wing due to flexion of phalanges of
June 29, 1954, 35 miles north the fifth digit. Photo June 29, se 35
Blythe, California. x 3s. miles north Blythe, California. >
Fic. 3. Macrotus californicus in level flight. wings are at the top of the
upstroke, but the digits have not yet been fully extended. Note that at this
point in the wing-beat cycle the hind limbs extend almost directly behind
bat and that camber of wings is reduced by extension of phalange s of fifth
digit. Photo June 29, 1954, 35 miles north Blythe, California. x %.
PLATE 4
Fic. 1. Head of Eumops perotis, female, from two miles east of El Cajon,
California, June 23, 1954, showing the position in which the ears are held
when the animal is using its eyes. 114.
Fics. 2 and 3. Same individual as shown in Fig. 1. 2) Front view, of bat
crawling. Note that the right forelimb, which is in the middle of the forward
component of the stride, is parallel to the substrate. 3) Side view of bat
crawling rapidly. The forelimb is near the end of its forward movement;
the hind limb is near the end of the propulsion part of the stride. « 34.
FUNCTIONAL MORPHOLOGY OF THREE BATS 137
SC
Fic. 1. Wings: Eumops (A), * %; Macrotus (B), *%; Myotis (C),
« %. Side view of the head of Eumops (D), * 14, showing the position
of the ear during flight.
138 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Nite (ide ape
occipito-pollicalis
- -pectoralis
/
serratus anterior~ , - -coraco-cutaneus
pect. abdom~
iy,
SY
\ZAZ
jy
ey G7 Uf > tensor plagiopatagii
= 2 eee ! YY Up
Sse NG
2D =
LE) x ——_ i S
Zi. a - - ~ -calcar
Fic. 2. Ventral view of Eumops, showing ie proportions of the body and
certain muscles, « 1
139
FUNCTIONAL MorpHo.ocy oF THREE Bats
“(W) suo
0% X “(@) snjosovy pure “AT X
W JO “(% ‘By 9es) soposnur omy osye pue Ap oq 94} Jo suonsodord Surmoys ‘Mora [e.QUOA
‘9 ‘Ory
140 UNIvERsITy OF Kansas Pusts., Mus. Nat. Hist.
- ---xiphoid process
Fic. 4. Sternum: Eumops (A), x 2; Myotis
(B), < 4; Macrotus (C), 3. For each sternum
the ventral view is on the left and the lateral
view, on the right.
141
FUNCTIONAL MORPHOLOGY OF THREE BATS
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r spine
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Dorsal view and anterior view, respectively, of the left scapula:
Eumops (A, B), * 2%; Myotis (C, D), * 5; Macrotus (E, F), x 4.
Fic. 5.
74357
142
UNIVERSITY OF KANSAS PUBLS., Mus. Nat. Hist.
r-pectoral ridge greater tuberosity— 5
r ~greater tuberosity
i
\
i)
|
E
r-pectoral ridge
1
I
'
1
L}
-trochlea
-med. epi.
!--capitulum
as H
L—lesser tuberosity
M
Fic. 6. Anterior and lateral view, respectively, of the proximal
end of the right humerus: Eumops (A, B), < 3; Myotis (C, D),
<5; Macrotus (E, F), * 4. End view of the proximal end of
the humerus: Eumops (G), * 8; Myotis (J), <5; Macrotus
(M), <4. Anterior view and lateral view, respectively, of the
distal end of the humerus: Eumops (H,1), * 3; Myotis (K, L),
<5; Macrotus (N, O), «4. Med. epi.—medial epicondyle;
lat. epi—lateral epicondyle.
FUNCTIONAL MORPHOLOGY OF THREE BATS
trapezium — 5
scaphoid--
i]
lunar-- 1
A
radius—- 4
_7~ trapezoid
. a
cuneiform -~
unciform- ~ 7
styloid process- -, magnum-~ //
ps. process— A pisiform— ~
~-sesamoid bone
7 -trapezium
/
, r-magnum
/
yr -lunar
Fic. 7. Ventral view of the carpus of Eumops (A), * 4. Pos-
terior view of the distal end of the left radius of Eumops (B),
<4. Dorsal view of the left carpus of Eumops (C), X 4. An-
terior views of the proximal ends of the left radii: Eumops (D),
<4; Myotis (E), * 7; Macrotus (F), 5. Inset shows cross
section of the fifth metacarpal of Eumops, * 4. Flex. fossa—the
depression into which the tendons of the Mm. biceps brachii and
brachialis insert. Ps. process—pseudostyloid process.
143
144 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
- - -pubic spine- - —{-/
dorsal ischial tuberosity- —
------- sympsysis
— -—--pubic spine
obturator fenestra- - —
dorsal rim of ischium- -
r -head
Fic. 8. Dorsal and lateral views, respectively, of the pelvis:
Eumops (A, B), * 2; Macrotus (C, D), * 4. Posterior view
of the proximal end of the left femur: Eumops (D), x 4:
Myotis (E), X 5; Macrotus (F), X 7.
FUNCTIONAL MORPHOLOGY OF THREE BATS
_--external cuneiform
_---middle cuneiform
— - --internal cuneiform
- - - -medial tarsal
-—=---- sesamoid bone
SS |} \ 424 --\- 7 oc astragalus
--\---- calcaneus
—-\- -tibia
- -\-fibula
---middle cuneiform
% is — -internal cuneiform
a -
-“--external cuneiform
a“
—-navicular
\\ > \ ~~ -calcaneus
~~ -astragalus
----- fibula
——--tibia
B
Fic. 9. Dorsal view of the left tarsus of Eu-
mops (A), < 5; and the right tarsus of Macrotus
(B), < 8. Note that in Eumops the tibia articu-
lates with the dorsal surface of the astragalus,
whereas in Macrotus the tibia articulates with
the proximal end of the astragalus. Inset shows
the posterior view of the proximal end of the
calear in Eumops.
146
UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
acromiotrapezius
Sosa acromiodeltoideus
\----—--- spinodeltoideus
NY : =
meh y\- ----- spinotrapezius
-- --- teres major
“Sogo triceps longus
- —— - --rhomboideus
S34 See triceps lateralis
------ brachialis
------ serratus anterior
----- biceps brachii
- — - - --latissimus dorsi
7 - — 7 -omocervicalis
a .
SSGs54 pectoralis
clavodeltoideus
epee subclavius
SS soe serratus anterior
=== ——- supraspinatus
- — —\ - - - — --teres minor
\ EN So levator scapulae
oN ae infraspinatus
SSeS sse subscapularis
— - - --- teres major
\------ triceps lateralis
\\ \ S eee triceps longus
aT ™
/ He I} i} Die :
IEYAZ = \ Vif
1
Y
Za
Z
jf
y
Fic. 10. A. Dorsal view of the shoulder region of Eumops. > 4%.
B. Dorsal view of the shoulder region of Eumops with some of the
superficial muscles removed. < 414.
FUNCTIONAL MORPHOLOGY OF THREE BATS 147
_7 — pectoral ridge
_7 ~biceps brachii (glenoid head)
- -biceps brachii (coracoid head)
- -subclavius
- - serratus anterior
= - subscapularis
rely pe es serratus anterior
Za F
7 teresmajorY ,
; , latissimus dorsi7
1
! w-triceps longus
'
I
1
i}
L -triceps lateralis
‘
t -triceps medialis
= S=>~< - - -biceps brachii(glenoid head)
a : 4 ee —lesser tuberosity
i; - —coracoid process
, \ [ fies — -flange of scapula
- -subscapularis
fuo---— serratus anterior
Fic. 11. A. Ventral view of the shoulder region of Eumops. X 4%.
B. Ventral view of the proximal part of the forelimb of Eumops. X 4%.
—----- triceps brachii
_77—brachialis
ps ~biceps brachii (glenoid head)
---- biceps brachii (coracoid head)
7-- -- extensor carpi radialis longus
_-7 777 extensor carpi radialis brevis
Pie supinator
-—— — extensor digitorum communis
\
SW - - - - extensor carpi ulnaris
--lateral epicondyle
SC oe LOR’ Br
----ulna
_-—7- extensor carpi radialis longus
--~ — pronator teres
a
—--palmaris longus
7
148 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
_--—-— flexor carpi ulnaris
_ 77 -extensor carpi ulnaris |
flexor carpi radialis
io ext. carpi radialis brevis
r-flex. digit. profund.
L—spinous process
Fic. 12. A. Lateral view of the proximal part of the forearm and distal
part of the upper arm of Eumops. X38. B. Medial view of the same
parts of the forelimb of Eumops. X 3
FUNCTIONAL MORPHOLOGY OF THREE BATS 149
~ —extensor pollicis brevis
7 7 —adductor pollicis
/ / Nets ste
, ¢ #7 77extensor indicis
‘
4 | -+—< =~ = — - extensor carpi radialis longus
=—iij 1S > —- extensor carpi radialis brevis
<p TAL j]
FE IN
— —-cuneiform
—s- - extensor carpi ulnaris
* \<- abd. poll. long.
~~ --interosseus dorsale
Fic. 13. Dorsal view of the wrist region of Eumops, showing the
tendons of some extensor muscles. 314.
150
r — -scaphoid
r — pisiform
'
ibe r abd. pollicis brevis
'
1
i]
i]
'
i]
'
'
‘
u
UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
-_——
Pies — adductor pollicis
ie oe ie ~ - occipito-pollicalis
tA 27 ~~ ~-adductor digiti secundi
Z a
-
—
SBA Oe _ 7 ~ interosseus dorsale
\
— \
——— == —S=
SY
. a \\'
oS ~ ale a 5
N Se interossel
—,
7} i
See So ~~ -abductor digiti quinti
SS \ ~ - adductor digiti quinti
. a —-flex. digit. profund.
~ ~apponens dig. quinti
i}
‘
|
!
!
1
|
I
'
|
'
U ff : :
| - flexor carpi ulnaris
'
i}
«—extensor carpi ulnaris
“ —flexor digitorum profundus
+ —palmaris longus
flexor carpi radialis
L - abductor pollicis longus
t —extensor carpi radialis longus
Fic. 14. Ventral view of the wrist region of Eumops, with part of
the fascial sheet of the insertion of the M. palmaris longus removed,
showing the tendons of some flexor muscles. x 344.
.
:
FUNCTIONAL MORPHOLOGY OF THREE BATS lsat
foc ccc — flexor digitorum fibularis
ro--cce peroneus brevis
,oorrccfle peroneus longus
4 ---—--extensor digitorum longus
7 --extensor hallucis longus
gracilis - -
— - - ~biceps femoris
semimembranosus
Fic. 15. A. Lateral view of the hind limb of Eumops. 2%.
B. Lateral view of the thigh of Eumops with the Mm. gluteus
maximus and tensor fasciae latae removed. 234
152 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist.
--adductor longus
---gracilis
\ - --semimembranosus
S
semitendinosus - - — &
gastrocnemius- --~
popliteus--~ _
Sy
plantaris— _
~
flexor digitorum fibularis- - -- -
tibialis posterior---.
'
gemellus --~-~ ;
adductor magnus--~~ -N
obturator externus--—~ ge \ Z
adductor brevis--~~ » ~ Ve ’ Y
psoas major---~ s\
Fic. 16. A. Medial view of the hind limb of Eumops. x 2%. |B and C.
Progressively deeper muscles of the thigh of Eumops from the medial
aspect. XX 2%.
FUNCTIONAL MORPHOLOGY OF THREE BATS 15s
iliacus — ~
>
psoas major— ~
psoas minor- ~~
< Ja
psoas major- —
{\x — - -extensor hallucis longus
~-extensor digitorum longus
\\. --peroneus brevis
\ Ss -peroneus longus
~---flex. dig. fib. lumbricales-~ x
Sor agg ee es gastroc. abd. hall. brev---+ +\
B \---depressor ossis styliformis med. tars-- — 4
\
flexor digitorum brevis-— —
plantaris - Gh.
tibialis posterior - -4
--abductor ossis metatarsi quinti
---plantaris adductor digiti quinti- - eS
Fic. 17. A. Lateral view of the pelvic region of Eumops. 5. B. Dorsal
view of the left foot of Eumops. C, D, and E. Progressively deeper muscles
of the left foot of Eumops from the ventral aspect. B-E are & 614.
Transmitted December 23, 1958. .-
‘=
27-4357
Vol. 8. 1.
Val. j/-9, 1.
Vol. 10.
15.
(Continued from inside of front cover)
The pigmy woodrat, Neotoma goldmani, its distribution and systematic posi-
tion. By Dennis re Rainey and Rollin H. Baker. Pp. 619-624, 2 figures in
text. June 10, 1955.
Index. Pp. 625-651.
2.
DSB ee
9.
10,
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.
Myology and serology of the Avian Family Fringillidae, a taxonomic study.
Py pe B. Stallcup. Pp. 157-211, 23 figures in text, 4 tables, November
An ecological study of the collared lizard (Crotaphytus collaris). By Henry
S. Fitch. Pp. 218-274, 10 figures in text. February 10, 1956.
A field study of the Kansas ant-eating frog, Gastrophryne olivacea. By Henry
S. Fitch. Pp. 275-306, 9 figures in text. February 10, 6.
Check-list of the birds of Kansas. By Harrison B. Tordoff. Pp. 307-359, 1
figure in text. March 10, 1956.
A population study of the prairie vole (Microtus ochrogaster ) in northeastern
tae By Edwin P. Martin. Pp. 361-416, 19 figures in text. April 2,
Temperature responses in free-living amphibians and reptiles of northeastern
Sere a By Henry S. Fitch. Pp. 417-476, 10 figures in text, 6 tables. June
Food of the crow, Corvus brachyrhynchos Brehm, in south-central Kansas. By
Dwight Platt. Pp. 477-498, 4 tables. June 8, 6.
Ecological observations on the woodrat, Neotoma floridana. By Henry S.
Fitch and Dennis G. Rainey. Pp. 499-533, 8 figures in text. June 12, 1956.
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.
2.
See Lae OS
Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 fig-
ures in text. December 10, 1955.
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.
A new long-eared myotis (Myotis evyotis) from northeastern Mexico. By Rol-
lin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.
Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming.
By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956.
The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6
figures in text, May 19, 1956.
Additional remains of the multituberculate genus Eucosmodon. By Robert W.
Wilson, Pp. 117-123, 10 figures in text. May 19, 1956.
Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures
in text. June 15, 1956.
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.
Extensions of known ranges of Mexican bats. By Sydney Anderson, Pp. 347-
351. August 15, 1956.
A new bat (Genus Leptonycteris) from Coahuila. By Howard J. Stains.
Pp. 358-856. January 21, 1957.
A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico.
By Robert J. Russell. Pp. 357-361. January 21, 1957.
Geographic variation in the pocket gopher, Thomomys bottae, in Colorado.
By ery M. Youngman. Pp. 363-384, 7 figures in text, 1 table. February
A new, bog lemming (genus Synaptomys) from Nebraska. By J. Knox
Jones, Jr. Pp. 385-888.
Pleistocene bats from San Josecito Cave, Nuevo Ledén, México. By J. Knox
Jones, Jr. Pp. 889-896, 1 figure in text. December 19, 1958.
New subspecies of the Rodent Baiomys from Central America. By Robert L.
Packard. Pp. 897-404. December 19, 1958.
More numbers will appear in volume 9.
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.
Comparative breeding behavior of Ammospiza caudacuta and A. maritima,
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.
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)
Vol. 11.
SS
iat
(Continued from inside of back cover)
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.
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.
The wood rats of Colorado: distribution and ecology. By Robert B. Finley,
Jr. Pp. 2138-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958.
More numbers will appear in volume 10.
The systematic status of the colubrid snake, Leptodeira discolor Giinther. By
William E. Duellman. Pp. 1-9, 4 figures. July 14, 1958.
Natural history of the six-lined racerunner. (Cnemidophorus sexlineatus). By
Henry S. Fitch. Pp. 11-62, 9 figures, 9 tables. September 19, 1958.
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.
A new snake of the genus Geophis from Chihuahua, México. By John
Legler. Pp. 327-384, 2 figures in text. January 28, 1959.
A new tortoise, genus Gopherus, from north-central México. By John Legler.
Pp. 335-343, 2 plates. April 24, 1959.
Fishes of Chautauqua, Cowley and Elk counties, Kansas. By Artie L, Metcalf.
Pp. 345-400, 2 plates, 2 figures in text. May 6, 1959.
Fishes of the Big Blue river basin, Kansas. By W. L. Minckley. Pp. 401-
442, 2 plates, 4 figures in text. May 8, 1959.
More numbers will appear in volume 11.
Vol. 12. Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry
A. Vaughan. Pp. 1-153; 4 plates, 17 figures. July 8, 1959.
—_~
Libis
AUG- ie 1959 ida
UNIVERSITY OF KANSAS PUBLICA sollR3 1 wy
MuUSEUM OF NATURAL HISTORY
Volume 12, No. 2, pp. 155-180, 10 figs.
Fil eens Se
The Ancestry of Modern Amphibia:
A Review of the Evidence
BY
THEODORE H. EATON, JR.
UNIVERSITY OF KANSAS
LAWRENCE
1959
UNIVERSITY OF KANSAS PUBLICATIONS, MusEUM OF NATURAL History
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson
Volume 12, No. 2, pp. 155-180
Published July 10, 1959
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED IN
THE STATE PRINTING PLANT
TOPEKA, KANSAS
1959
>
27-8362
MUS. COMP. 7001
LIBRARY
AUG - 61959:
HARVARD
UNIVERSITY
The Ancestry of Modern Amphibia
A Review of the Evidence
BY
THEODORE H. EATON, JR.
INTRODUCTION
In trying to determine the ancestral relationships of modern orders
of Amphibia it is not possible to select satisfactory structural an-
cestors among a wealth of fossils, since very few of the known fossils
could even be considered possible, and scarcely any are satisfactory,
for such a selection. The nearest approach thus far to a solution of
the problem in this manner has been made with reference to the
Anura. Watson’s paper (1940), with certain modifications made
necessary by Gregory (1950), provides the paleontological evidence
so far available on the origin of frogs. It shows that several fea-
tures of the skeleton of frogs, such as the enlargement of the inter-
pterygoid spaces and orbits, reduction of the more posterior dermal
bones of the skull, and downward spread of the neural arches
lateral to the notochord, were already apparent in the Pennsylvanian
Amphibamus (Fig. 1), with which Gregory synonymized Mioba-
trachus and Mazonerpeton. But between the Pennsylvanian and
the Triassic (the age of the earliest known frog, Protobatrachus )
there was a great lapse of time, and that which passed between
any conceivable Paleozoic ancestor of Urodela and the earliest
satisfactory representative of this order (in the Cretaceous) was
much longer still. The Apoda, so far as known, have no fossil
record.
Nevertheless it should be possible, first, to survey those char-
acters of modern Amphibia that might afford some comparison
with the early fossils, and second, to discover among the known
Paleozoic kinds those which are sufficiently unspecialized to permit
derivation of the modern patterns. Further circumstantial evidence
may be obtained by examining some features of Recent Amphibia
which could not readily be compared with anything in the fossils;
such are the embryonic development of the soft structures, including
cartilaginous stages of the skeleton, the development and various
specializations of the ear mechanism, adaptive characters associated
with aquatic and terrestrial life, and so on.
(157)
158 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
COMPARISON OF MODERN ORDERS WITH THE
LABYRINTHODONTS AND LEPOSPONDYLS
In both Anura and Urodela the skull is short, broad, relatively
flat, with reduced pterygoids that diverge laterally from the para-
sphenoids leaving large interpterygoid vacuities, and with large
orbits. (These statements do not apply to certain larval or peren-
Protcbatrachus
Fic. 1. Saurerpeton (X %, after Romer, 1930, fig. 6); Am-
phibamus, the palatal view 2h, from Watson, 1940, fig. 4
(as Miobatrachus), the dorsal view < 2%, from Gregory’s
revised figure of Amphibamus (1950, Fig. 1); Protobatrachus,
<1, from Watson, 1940, fig. 18, 19.
nibranchiate forms.) The skull in both orders has lost a number
of primitive dermal bones in the posterior part; these are: basi-
occipital, supraoccipital, postparietal, intertemporal, supratemporal,
and tabular. The exoccipitals form the two condyles but there are
no foramina for the 11th and 12th nerves, since these are not
separate in modern Amphibia. The opisthotic is missing in all
Tue ANCESTRY OF MODERN AMPHIBIA 159
except Proteidae (but see discussion of the ear). Although the
skull is normally autostylic, a movable basipterygoid articulation is
present among Hynobiid salamanders and in at least the meta-
morphic stages of primitive frogs, and therefore should be expected
in their ancestors. The vertebrae are, of course, complete; see dis-
cussion in later section. The quadratojugal, lost in salamanders,
is retained in frogs, and conversely the lacrimal, absent in frogs,
occurs in a few primitive salamanders. The situation in Apoda is
different, but postfrontal and jugal should be noted as bones re-
tained in this order while lost in the others.
Thus, in spite of minor differences, the above list shows that
there are numerous and detailed similarities between Anura and
Urodela with respect to the features in which they differ from the
Paleozoic orders. Pusey (1943) listed 26 characters which Ascaphus
shares with salamanders but not with more advanced frogs; a few
of these might be coincidental, but most of them are of some com-
plexity and must be taken to indicate relationship. The main
adaptive specializations of Anura, however, including loss of the
adult tail, extreme reduction in number of vertebrae, formation of
urostyle, elongation of the ilium and lengthening of the hind legs,
must have appeared at a later time than the separation of that order
from any possible common stem with Urodela, although they are
only partially developed in the Triassic Protobatrachus.
Turning to the Paleozoic Amphibia, there are two groups in
which some likelihood of a relationship with modern order exists.
In the Pennsylvanian Trimerorhachoidea (Labyrinthodontia, order
Temnospondyli) some members, such as Eugyrinus, Saurerpeton,
and notably Amphibamus (Fig. 1) had short, broad heads, an
expansion of palatal and orbital openings, posterior widening of
the parasphenoid associated with divergence of the pterygoids, a
movable basipterygoid articulation, and reduction in size (but not
loss) of the more posterior dermal bones of the skull. In recogni-
tion of Watson’s (1940) evidence that these animals make quite
suitable structural ancestors of frogs, Romer (1945) placed Am-
phibamus in an order, Eoanura, but Gregory (1950) indicated that
it might better be left with the temnospondyls. Association of the
urodele stem with this group does not seem to have been proposed
hitherto.
The other group of Paleozoic Amphibia that has been considered
probably ancestral to any modern type is the subclass Lepospondyli,
containing three orders, Aistopoda, Nectridia and Microsauria. In
160 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
these the vertebrae are complete (holospondylous ), the centra pre-
sumably formed by cylindrical ossification around the notochord,
and there is no evidence as to the contributions from embryonic
cartilage units. It is important to note at this point that precisely
the same statement can be made regarding the vertebrae of adults
of all three Recent orders, yet for all of them, as shown in a later
section, we have ample evidence of the part played by cartilage
elements in vertebral development. Therefore (a) we cannot say
that there were no such elements in embryonic stages of lepospon-
dyls, and (b) it would take more than the evidence from adult
vertebrae to relate a particular modern order (for example, Uro-
dela) to the Lepospondyli. Vague similarities to Urodela have been
noted by many authors in the Nectridia, Aistopoda and Microsauria,
but these are not detailed and refer mainly to the vertebrae. The
skulls do not show, either dorsally or in the palate, any striking
resemblance to those of generalized salamanders, and certainly most
known lepospondyls are too
specialized to serve as the
source of Urodela. It is true
that the elongate bodies, small
limbs, and apparent aquatic
habitus of some lepospondyls
accord well with our usual
picture of a salamander, but
such a form and way of life
have appeared in many early
Amphibia, including the laby-
rinthodonts. The family Ly-
sorophidae (Fig. 2), usually
placed among microsaurs, is
sufficiently close in skull struc-
Fic. 2. Lysorophus tricarinatus, lateral ;
and posterior views < 2%, modified after ture to the Apoda to be a
Sollas, 1920, Figs. 8 and 12, respectively; possible ancestor of these, but
palatal view after Broom, 1918, x 1%. it probablv has nothing to do
For explanation of abbreviations see [ ae ©
Fig. 3. with Urodela, by reason of
the numerous morphological
specializations that were associated with its snakelike habitus.
McDowell’s (1958) suggestion that it would be profitable to
look among the Seymouriamorpha for the ancestors of frogs seems
to be based upon a few details of apparent resemblance rather than
a comprehensive view of the major characters of the animals. In
most points which he mentions (limb girdles, form of ear, pterygoid
Tue ANCESTRY OF MODERN AMPHIBIA 161
articulation) the present writer does not see a closer similarity of
frogs to Seymouriamorpha than to Temnospondyli.
Still other opinions have been expressed. Herre (1935), for
instance, concludes “on anatomical, biological and paleontological
grounds” that the orders of Urodela, Anura, Apoda and Stego-
cephali were all independently evolved from fish, but beyond citing
the opinions of a number of other authors he does not present
tangible evidence for this extreme polyphyletic interpretation.
More notable are the views of several Scandinavian workers
(Saive-Sdderbergh, 1934; Jarvik, 1942; Holmgren, 1933, 1939, 1949a,
b), of whom Jarvik, in a thorough analysis of the ethmoid region,
would derive the Urodela from Porolepid Crossopterygii, and all
other tetrapods from the Rhipidistia; Sive-Sdderbergh and Holm-
gren, the latter using the structure of carpus and tarsus, see a
relationship of Urodela to Dipnoi, but accept the derivation of
labyrinthodonts and other tetrapods from Rhipidistia. All of this
work is most detailed and laborious, and has produced a great
quantity of data useful to morphologists, but the diphyletic theory
is not widely adopted; the evidence adduced for it seems to consist
largely of minutiae which, taken by themselves, are inconclusive, or
lend themselves to other interpretation. For instance Holmgren’s
numerous figures of embryonic limbs of salamanders show patterns
of cartilage elements that he would trace to the Dipnoan type of
fin, yet it is difficult to see that the weight of evidence requires this,
when the pattern does not differ in any fundamental manner from
those seen in other embryonic tetrapods, and the differences that
do appear may well be taken to have ontogenetic rather than phylo-
genetic meaning. Further, the Dipnoan specialization of dental
plates and autostylic jaw suspension, already accomplished early in
the Devonian, would seem to exclude Dipnoi from possible ancestry
of the Urodela, an order unknown prior to the Mesozoic, in which
the teeth are essentially similar to those of late Paleozoic Amphibia,
and the jaw suspension is not yet in all members autostylic.
THE EAR
In temnospondylous Amphibia the tympanum generally occupied
an otic notch, at a high level on the skull, bordered dorsomedially
by the tabular and ventrolaterally by the squamosal. In this posi-
tion the tympanum could receive airborne sounds whether the
animal were entirely on land or lying nearly submerged with only
the upper part of its head exposed. Among those Anura in which
the ear is not reduced the same is true, except that the tabular is
162 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
lost. In Temnospondyli (Fig. 3) the posterior wall of the otic
capsule was usually formed by the opisthotic, which extended up
and outward as a buttress from the exoccipital to the tabular, and
sometimes showed a paroccipital process for the insertion, pre-
sumably, of a slip or tendon of the anterior axial musculature. The
p.t.f.. pp. soc. exoc. par.proc.
lateral
Necturus
jugular 5
posterior
Fic. 3. Occipital region of skulls of Megalocephalus brevicornis (< *4o0, after
Watson, 1926, as Orthosaurus), Dvinosaurus (><, modified after Bystrow,
1938; the lower figure after Sushkin, 1936), and Necturus maculosus (X 8,
original, from K. U., No. 3471).
ABBREVIATIONS USED IN FIGURES
b’d.c.—basidorsal carti-
lage (neural arch)
b’oc.—basioccipital
ce.1-4—centrale1_4
ch.—ceratohyal
clav.—clavicle
clei.—cleithrum
cor.—coracoid
d.c.1-4—distal carpal-4
diap.—diapophysis
exoc.—exoccipital
ep.—episternum
hyost.—hyostapes
i—intermedium
Mk.—Meckel’s cartilage
n.—notochord
om.—omosternum
op.—operculum
opis.—opisthotic
par.—parietal
par. proc.—paroccipital
process
peri. cent.—perichordal
centrum
p p.—postparietal
prep.—prepollex
pro.—prootic
p’sp.—parasphenoid
pt.—pterygoid
p.t.f.—post-temporal
fossa
postzy g.—postzygapo-
physis
qj.— quadratojugal
qu.—quadrate
ra.—radiale
r.hy.—hyomandibular
ramus of VII
rib-b.—rib-bearer
r.md.—mandibular
ramus of VII
sc.—scapula
sc’cor.—scapulocoracoid
s’d.—supradorsal
cartilage
s’d.(postzyg.)—suprador-
sal (postzygapophysis)
soc.—supraoccipital
sp.c.—spinal cord
sq.—squamosal
s’sc.—suprascapula
s't.—supratemporal
sta.—stapes
ster.—sternum
tab.—tabular
uln.—ulnare
v.a.—vertebral artery
xiph.—xiphisternum
I,1V—digits I and IV
V, VII, X, XI1J—foramina
for cranial nerves of
these numbers (in
Fig. 4, VII is the fa-
cial nerve )
163
THE ANCESTRY OF MODERN AMPHIBIA
stapes, in addition to its foot in the fenestra ovalis and its tympanic
or extrastapedial process to the tympanum, bore a dorsal process
(or ligament) to the tabular, an “internal” process (or ligament)
to the quadrate or an adjacent part of the squamosal, and a ligament
to the ceratohyal. Some of these attachments might be reduced
or absent in special cases, but
—~—~_ dorsal process
otostapes r
they seem to have been the ones
originally present both phylo-
genetically and embryonically in AND 4
Amphibia. >
Among typical frogs (Fig. 4)
the base, or otostapes, is present
and bony, the extrastapedial
process (exiracolumella, or hyo-
stapes) is usually cartilaginous,
the dorsal process (processus
paroticus ) is of cartilage or liga-
ment, but the other two attach-
ments are absent in the adult.
Fic. 4. Diagram of middle ear struc-
The exoccipital extends laterally,
occupying the posterior face of
the otic capsule. Between it and
the otostapes is a small disc,
tures in Rana (upper figure, atter
Stadtmiiller, 1936, and lower left after
DeBeer, 1937), and Ambystoma (lower
right, after DeBeer, 1937); all > 4.
For explanation of abbreviations see
Fig. 3
usually ossified, the operculum,
which normally fits loosely in a portion of the fenestra! membrane,
and is developed from the otic capsule. The opercularis muscle
extends from this disc to the suprascapula, in many but by no means
all families of Anura.
Among Urodela (Fig. 4) the middle ear cavity and tympanum
are lacking, and the stapes (columella) consists of no more than
its footplate and the stylus, which is attached to the border of the
squamosal, thus corresponding to the “internal” process. In fami-
lies in which individuals metamorphose and become terrestrial
(Hynobiidae, Ambysiomidae, Salamandridae, Plethodontidae), an
operculum and opercularis muscle appear in the adult, just as in
frogs, except that in Plethodontidae, the most progressive family,
the operculum fuses with the footplate of the stapes. Among
neotenous or perennibranchiate urodeles there is no separate oper-
culum or opercularis. The evidence given by Reed (1915) for fusion
of the operculum with the columella in Necturus appears incon-
clusive, in spite of the great care with which his observations were
164 University oF Kansas Pusts., Mus. Nat. Hist.
made. On the other hand, Necturus and Proteus alone among
living salamanders have a distinct opisthotic on the posterior wall
of the otic capsule (Fig. 3), as do the Cretaceous Hylaeobatrachus
and the Eocene Palaeoproteus. Probably these Proteidae should
be regarded as primitive in this respect, although many other fea-
tures may be attributed to neoteny.
There is a contrast between Anura and most Urodela in the rela-
tive positions of the stapes and facial nerve, as shown in DeBeer’s
(1937) diagrams. In the latter (Ambystoma) the nerve is beneath,
and in the former (Rana) above, the stapes. Judging by figures of
Neoceratodus, Hypogeophis, and several types of reptiles and
mammals, the Urodela are exceptional. Necturus, however, has the
nerve passing above its stapes, and this may be primitive in the
same sense as the persistent opisthotic. There can be, of course,
no question of the nerve having worked its way through or over
the obstructing stapes in order to come below it in salamanders;
rather, the peripheral growth of neuron fibers in the embryo must
simply pursue a slightly different course among the partially differ-
entiated mesenchyme in the two contrasting patterns.
Although DeBeer (1937) shows in his figure of Hypogeophis
(one of the Apoda) an operculum, this is apparently a mistake.
The stapes has a large footplate, and its stylus articulates with the
quadrate, but no true operculum or opercularis has been described
in the Apoda. The facial nerve passes above the stapes. It does
not seem necessary to regard the conditions in this order as related
directly to those of either salamanders or frogs, but a reduction of
the stapes comparable to that in salamanders has occurred.
The presence in both frogs and terrestrial salamanders of a special
mechanism involving the opercularis muscle and an operculum
cut out in identical fashion from the wall of the otic capsule behind
the stapes seems to require some other explanation than that of a
chance convergence or parallelism. Although the stapes and otic
region are readily visible in a number of labyrinthodonts and lepo-
spondyls, no indication of an operculum seems to be reported among
them. But in the Triassic Protobatrachus (Fig. 1), which is un-
mistakably a frog in its skull, pelvis and some other features, Pive-
teau (1937) has shown, immediately behind the foot of the stapes,
a small bony tubercle, which he and Watson (1940) designated
opisthotic. Very clearly it served for insertion of a muscle, and it
is equally clear that the bone is a reduced opisthotic, carrying the
paroccipital process already mentioned as characteristic of it in
THE ANCESTRY OF MopERN AMPHIBIA 165
some temnospondyls. Since the remainder of the posterior wall of
the otic capsule consists of cartilage, meeting the exoccipital, it
may be that the opisthotic becomes the operculum in frogs. Proto-
batrachus was too far specialized in the Anuran direction, although
it still had a tail, and the forelegs and hind legs were nearly the
same size, to be considered a possible ancestor of the Urodeles.
But at one stage in the general reduction of the skull in the ancestry
of both groups, a condition similar to that in Protobatrachus may
have characterized the otic region, long before the Triassic.
In the argument thus far we have considered terrestrial, adult
amphibians, since it is only in these that either the normal middle
ear and tympanum, or the opercular apparatus, is present. But
among the urodeles several neotenic types occur (this term applies
also to the perennibranchs). For most of these there is nothing
about the otic region that would be inconsistent with derivation, by
neoteny, from known families in which adults are terrestrial; for
example, Cryptobranchus could have had a Hynobiid-like ancestor.
But this, as mentioned above, does not hold for the Proteidae, which
possess an opisthotic of relatively large size, distinctly separate
from the exoccipital and prootic. Either this bone is a neomorph,
which seems improbable, or there has not been in the ancestry of
this particular family an episode of reduction comparable to that
seen in the terrestrial families, where there is an operculum instead
of a normal opisthotic. Therefore the Proteidae probably are not
derived from the general stem of other salamanders, but diverged
sufficiently long ago that the bones of the otic region were reduced
on a different pattern. They need not be removed from the order,
but, in this respect, recognized as more primitive than any other
existing Urodela or Anura. A recent paper by Hecht (1957) dis-
cusses many features of Necturus and Proteus, and shows that they
are remote from each other; his evidence does not seem to prove,
however, that they were of independent origin or that they need
be placed in separate families.
VERTEBRAE AND RIBS
Development of the vertebrae and ribs of Recent Amphibia has
been studied by Gamble (1922), Naef (1929), Mookerjee (1930 a,
b), Gray (1930) and Emelianov (1936), among others. MacBride
(1932) and Remane (1938) provide good summaries. In this sec-
tion reference will be made to the embryonic vertebral cartilages
by the names used for them in these studies, although the concept
166 UNIveERSITY OF Kansas Pusts., Mus. Nat. Hist.
of “arcualia” is currently considered of little value in comparative
anatomy.
The centrum in Anura (Fig. 5) is formed in the perichordal
sheath (Rana, Bufo) or only in the dorsal portion thereof (Bombi-
nator, Xenopus). The neural arch develops from the basidorsal
cartilages that rest upon, and at first are entirely distinct from, the
perichordal sheath. Ribs, present as separate cartilages associated
with the 2nd, 3rd and 4th vertebrae in the larvae of Xenopus and
sd. (postzyg.)
Q- rib
oe ‘//}— pen. cent.
= ~ notochord
“hyvochord
epichordal centrum
Fic. 5. Development of Anuran vertebrae. Upper left, late tadpole of
Xenopus laevis; lower left, same just after metamorphosis; upper right,
diagram of general components of primitive Anuran vertebra. (After
MacBride, 1932, Figs. 35, 38, 47D, respectively.) Lower right, section
through anterior portion of urostyle, immediately posterior to sacral verte-
bra, in transforming Ascaphus truei (original, from specimen collected on
Olympic Peninsula, Washington). All < 20 approx. For explanation of
abbreviations see Fig. 3.
Bombinator, fuse with lateral processes (diapophyses ) of the neural
arches at metamorphosis, but in Leiopelma and Ascaphus the ribs
remain freely articulated in the adult. Basiventral arcualia have
been supposed to be represented by the hypochord, a median rod
of cartilage beneath the shrinking notochord in the postsacral
region, which at metamorphosis ossifies to produce the bulk of the
urostyle. Fig. 5, lower right, a transverse section taken immediately
posterior to the sacral ribs in a transforming specimen of Ascaphus,
THE ANCESTRY OF MODERN AMPHIBIA 167
shows that the “hypochord” is a mass of cartilage formed in the
perichordal sheath itself, and very obviously is derived from the
ventral part of postsacral perichordal centra; there are, then, no
basiventral arcualia, and the discrete hypochord shown in Mac-
Bride’s diagram (Fig. 5, upper right) of a frog vertebra does not
actually occur below the centrum, but only below the notochord in
the postsacral region.
In Urodela (Fig. 6) the pattern of vertebral and rib development
is more complex, and there has been much controversy over its
mesenchyme
(basiventral) bony process
basiventral
Fic. 6. Development of Urodele vertebrae. Upper figures, Triton: at left,
larva at 20 mm., at right, diagram of components of vertebra (from Mac-
Bride, 1932, figs. 17, 47C). Middle figures, Molge vulgaris larva: left, at
18 mm.; middle, at 20-22 mm.; right, at 25 mm. (from Emelianov, 1936,
figs. 33, 36, 38 respectively). Lower figures, Necturus maculosus larva:
left, at 21 mm.; right, at 20 mm. (from MacBride, 1932, figs. 41.5, 41.3
respectively, after Gamble, 1922). All & 20 approx. For explanation of
abbreviations see Fig. 3.
interpretation. Neural arches and perichordal centra form in the
same manner as in frogs, but with the addition in certain cases
(Triton) of a median supradorsal cartilage, which gives rise to the
zygapophyses of each neural arch. Difficulty comes, however, in
understanding the relationship of the ribs to the vertebrae. Each
168 UNIveERSITY OF Kansas Pusts., Mus. Nat. Hist.
rib, usually two-headed, articulates with a “transverse process” that
in its early development seems to be separate from both the vertebra
and the rib, and is therefore known, noncommittally, as “rib-bearer.”
This lies laterally from the centrum, neural arch, and vertebral
artery; upon fusing with the vertebra it therefore encloses the artery
in a foramen separate from the one between the capitulum and
tuberculum of the rib (the usual location of the vertebral artery).
At least four different interpretations of these structures have been
suggested:
(1) Naef (1929) considered the rib-bearer a derivative of the
basiventral, which, by spreading laterally and dorsally to meet the
neural arch, enclosed the vertebral artery. He then supposed that
by reduction of the rib-bearer in other tetrapods (frogs and am-
niotes ) the vertebrarterial foramen and costal foramen were brought
together in a single foramen transversarium. The implication is that
the Urodele condition is primitive, but it cannot now be supposed
that Urodela are ancestral to any other group, and the rib-bearer
is most probably a specialization limited to salamanders. This
does not, of course, invalidate the first part of his interpretation.
(2) Remane (1938), noting that rib insertions of early Amphibia
are essentially as in Amniota, argued that the rib-bearer is not from
the basiventral but is a neomorph which originates directly from
the neural arch and grows ventrally. This he inferred mainly from
Gamble’s (1922) observation on Necturus, but his assumption that
Necturus is more primitive than other salamanders (such as the
Salamandridae), where the pattern differs from this, is not neces-
sarily correct. Rather, the perennibranchs are distinguished mainly
by their neotenous features, and their development is likely to show
simplifications which are not necessarily primitive. The suggestion
of a “neomorph” ought not to be made except as a last resort, for
it is simply an acknowledgment that the author does not recognize
homology with any structure already known; sometimes further
information will make such recognition possible.
(3) Gray (1930), using Molge taeniatus, concluded that the
normal capitulum of the rib was lost, but that the tuberculum
bifurcated to make the two heads seen in Urodela, thus accounting
for the failure of the costal foramen to coincide with that of the
vertebral artery. This answer, too, seems to entail an unprovable
assumption which should not be made without explicit evidence.
(4) Finally, Emelianov (1936) regarded the rib-bearer as a rudi-
mentary ventral rib, on account of its relationship to the vertebral
artery, and considered the actual rib to be a neomorph in the dorsal
Tue ANCESTRY OF MODERN AMPHIBIA 169
position characteristic of tetrapod ribs in general. This argument
would fit the ontogenetic picture satisfactorily, provided that (a)
there were some evidence of ventral, rather than dorsal, ribs in
early Amphibia, and (b) we accept the invention of another neo-
morph in modern Amphibia as an unavoidable necessity. Emelia-
novs conclusion (p. 258) should be quoted here (translation):
“The ribs of Urodela are shown to be upper ribs, yet we find besides
these in Urodela rudimentary lower ribs fused with the vertebral
column. The ribs of Apoda are lower ribs. In Anura ribs fail to
develop fully, but as rare exceptions rudiments of upper ribs
appear.”
Of these various interpretations, that of Naef seems to involve
the minimum of novelty, namely, that the rib-bearer is the basi-
ventral, expanded and external to the vertebral artery. It is not
necessary to take this modification as the ancestral condition in
tetrapods, of course. The basiventral (= intercentrum) would
merely have expanded sufficiently to provide a diapophysis for the
tuberculum as well as the (primitive) facet for the capitulum.
No neomorph appears under this hypothesis, which has the distinct
advantage of simplicity.
Figures of early stages in vertebral development by the authors
mentioned show that the basidorsals chondrify first, as neural arches,
while a separate mass of mesenchyme lies externally and ventrally
from these. This mesenchyme may chondrify either in one piece
(on each side) or in two; in Molge the part adjacent to the centrum
is ossified in the 20-mm. larva, and subsequently unites with the
more dorsal and lateral cartilaginous part, while the rib, appearing
farther out, grows inward to meet this composite “rib-bearer.” In
Necturus the mesenchyme below the neural arch differentiates into
a cartilage below the vertebral artery (position proper to a basi-
ventral), a bridge between this and the neural arch, and a rib, the
latter two chondrifying later than the “basiventral” proper. In the
“axolotl” (presumably Ambystoma tigrinum) the rib-bearer grows
downward from its first center of chondrification at the side of the
neural arch (Emelianov, 1936).
Thus it appears that the simplest hypothesis to account for the
rib-bearer is that (a) it is the basiventral, (b) it is recognizable
just before chondrification as a mass of mesenchyme in contact
with both the notochordal sheath and the basidorsal cartilage,
(c) it may chondrify or ossify first in its ventral portion or in its
dorsal portion, the two then joining before it fuses with the rest
of the vertebra, (d) the enclosure of the vertebral artery is a con-
170 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
sequence of the extension of the basiventral beyond the position
occupied by it in primitive Amphibia, and (e) there is no indica-
tion that this took place in other orders than the Urodela.
It seems that the vertebrae in Urodela have at least the following
components: perichordal centra, separate basidorsal cartilages, and
basiventrals, which are somewhat specialized in their manner of
development. The vertebrae of Anura develop in the fashion just
described except that basiventrals are lacking. It would seem no
ARES
Trimerorhachis
Fic. 7. Vertebrae of Eusthenopteron (>< 1) and Ichthyostega
(< %, after Jarvik, 1952), Trimerorhachis (x 1%, after Case),
and Amphibamus (> 10, after Watson, 1940) in lateral and end
views; the two lower right-hand figures are from Watson (1940,
as Miobatrachus); the lower left is from a cast of the “Mioba-
trachus” specimen in Chicago Natural History Museum, No. 2000,
in the presacral region (original, « 10).
more difficult to accept the derivation of salamander vertebrae
from the temnospondylous type than it is in the case of frogs, if
other evidence points to such an ancestry.
Fig. 7, lower right, is Watson’s (1940) illustration of the anterior
trunk vertebrae of Amphibamus (Miobatrachus), in which the
intercentrum is shown as a single median piece. Fig. 7, lower left,
shows two of the more posterior trunk vertebrae seen as impressions
in a cast of the type of “Miobatrachus romeri;” evidently the inter-
centra were paired at about the level of the 16th vertebra, and
Tuer ANCESTRY OF MODERN AMPHIBIA 1
relatively large. Gregory’s (1950) figure of the type specimen of
“Mazonerpeton” (also equivalent to Amphibamus) shows the an-
terior trunk vertebrae in relation to the ribs essentially as they
appear to me in the cast of Miobatrachus, and rather differently
from Watson’s figure of the latter. Gregory is probably right in
considering the specimens to represent various degrees of imma-
turity. So far as present information goes, then, the vertebrae of
salamanders and frogs show no clear evidence of derivation from
those of any particular group among the early Amphibia, but their
features are not inconsistent with a simplification of the pattern of
Temnospondyli.
clei.
SC.
clav
SSC Loaves SS
Ce wae
Fic. 8. Pectoral girdles of Protobatrachus (after Piveteau,
1937), Notobatrachus (after Stipanicic and Reig, 1956),
Ascaphus (after Ritland, 1955 a) and Rana (original); all
<2. For explanation of abbreviations see Fig. 3.
PECTORAL GIRDLE
Hecht and Ruibal (Copeia, 1928:242) make a strong point of the
nature of the pectoral girdle in Notobatrachus, as described recently
by Stipanicic and Reig (1955, 1956) from the Jurassic of Patagonia,
and quite rightly recommend that the significance of the arciferal
and firmisternal types of girdle be restudied. That of Notobatrachus
is said to be firmisternal; in view of the arciferal condition in the
supposedly primitive Leiopelma, Ascaphus, Bombinator, etc., this
comes as a surprise. Is the firmisternal girdle, as seen in Rana, Bufo,
and others, actually the ancestral type, and has the arciferal been
derived from something like this?
12 UNIveERSITY OF KANSAS Pusts., Mus. Nat. Hist.
In the figures given by Stipanicic and Reig the ossified parts of
the girdle are figured in detail (Fig. 8) and Reig’s discussion of it
is thorough. The decision to call it firmisternal was taken with
some hesitancy, for no median elements are indicated, and the posi-
tion and shape of those seen is closely similar to the ossified parts
in Ascaphus and Leiopelma; there is no bony sternum or omoster-
num. It is safe to suppose that some cartilage lay in the midline
between the clavicles and coracoids, but there is no evidence as to
its extent, rigidity, or degree of overlapping if any. Apparently,
then, there is not sufficient reason to infer that this Jurassic frog
had a pectoral girdle comparable with the modern firmisternal type.
Piveteau (1955:261) remarks that the only living Anuran that
can be compared usefully with Protobatrachus (Triassic) with re-
gard to its pectoral girdle is Ascaphus. Again, the extent of car-
tilage in Protobatrachus (Fig. 8) can only be inferred, and there
are no median elements. The agreement with Ascaphus includes
the presence, in both, of a separate coracoid ossification situated
posterior to the ossified “scapulocoracoid” (actually scapula). This
ossification is evidently that shown in Notobatrachus as “coracoid.”
Direct comparison of the three genera with one another suggests
that if we use the term arciferal for any, we should use it for all.
In the remote predecessor of Anura, Amphibamus of the Penn-
sylvanian, the pectoral girdle was less substantial than in many of
its contemporaries, but it contained the primitive median inter-
clavicle in addition to the clavicle, cleithrum, and scapulocoracoid.
(The figure of Watson, 1940, and that by Gregory, 1950, are of
individuals of different ages, the latter being older.) It is clear
that the paired elements of such a girdle were held rigid by their
attachment to the interclavicle, via the clavicles. Subsequent elim-
ination of the interclavicle in the Anuran line of descent, and de-
crease of ossification, left a girdle like that of Protobatrachus, Noto-
batrachus, Ascaphus and Leiopelma. But in several advanced
families a more rigid median “sternum,” of one or two bony pieces
plus cartilage, is developed secondarily, possibly (as Cope, 1889:
247, suggested) in correlation with axillary amplexus.
Among Urodela no dermal bones occur in the pectoral girdle.
There is usually a scapulocoracoid ossified as a single piece, from
which a thin cartilaginous suprascapula extends dorsally and a
broad cartilaginous coracoid plate extends medially, overlapping
the one from the opposite side; a precoracoid lobe of this reaches
forward on either side, and a median, posterior “sternum” of carti-
Tue ANCESTRY OF MODERN AMPHIBIA 173
lage may make contact with the edges of the two coracoids. In
Siren and Amphiuma two centers of ossification are found for each
scapulocoracoid, and in Triton and Salamandra three. Probably
the more dorsal and lateral of these represents the primitive scapula
and the other one (or two) the primitive coracoid.
Comparing the girdle of a salamander with that of a frog, the
closest similarity can be seen between Ascaphus and a salamander
in which the scapula and coracoid ossify separately. Both have
the median “sternum” in contact with the coracoid plates. The
major difference, of course, is the lack of clavicle and cleithrum
in the salamander.
CARPUS AND TARSUS
In Ascaphus (Ritland, 1955a; cleared and stained specimens of
nearly grown males) distal carpals 1, 2, 3 and 4 are present and
separate, increasing in size in the order given (Fig. 9). A pre-
= a i.+uln.
rcen fs
77 &
Fic. 9. Skeleton of fore foot of Notobatrachus (after
Stipanicic and Reig, 1956, terminology revised) and
Ascaphus (after Ritland, 1955 a); all & 5. For explana-
tion of abbreviations see Fig. 3.
Notobatrachus Ascaphus
pollex rests against centrale 1; centralia 2 and 3 are fused; the
radiale fuses with centrale 4, and the intermedium fuses with the
ulnare; radius and ulna are fused with each other as in other frogs.
The digits (and metacarpals) are considered by Ritland to be 1-4,
in addition to the prepollex, rather than 2-5.
In the Jurassic Notobatrachus Stipanicic and Reig (1956) have
shown the carpus with surprising clarity (Fig. 9). If their nomen-
clature of the parts be revised, we obtain a fairly close resemblance
to Ascaphus, except that centralia 2 and 8 are not fused, distal
174 UNIVERSITY OF Kansas PuBts., Mus. Nat. Hist.
carpals 1 and 2 do not show (which would easily be understood
if they were of the size of those in Ascaphus, or not ossified), and
the intermedium remains separate from the ulnare.
In Salamandra (Francis, 1934; Nauck, 1938) distal carpals 1 and
2 are fused in both larva and adult, and 3 and 4 are separate; the
radiale, intermedium and ulnare are separate in the larva but the
latter two fuse in the adult; centrale 1 (labelled prepollical cartilage
by Francis) and centrale 2 are separate. Francis considers the
digits (and metacarpals) to be 1-4. Apparently the arrangement
here indicated for the larva is characteristic of other larval sala-
manders, except where further reduced, and reduction below the
number given for the adult is common in other terrestrial forms.
The radius and ulna are, of course, separate.
The ossification of carpals is more likely to be complete in adult
frogs than in salamanders, but some ossification of all parts named
is found in several of the latter. A common ancester of frogs and
salamanders could be expected to have the following elements
present and ossified in the adult: distal carpals 1-4 separate; 8
centralia; radiale, intermedium and ulnare separate. Comparison
with fossils older than Notobatrachus is fruitless on these points,
unless we go back to forms too distant to have any special value,
such as Eryops. This is because of inadequate preservation and
because the elements are not fully ossified in many immature speci-
mens.
For the purpose of this review there is no special value in a
comparison of the tarsi of frogs and salamanders, since the leaping
adaptation of the former leaves very little common pattern between
them. Even in Protobatrachus, where the legs were not yet con-
spicuously lengthened, the tibiale and fibulare (“astragalus” and
“calcaneum” respectively) were already considerably elongated.
The carpus and tarsus of Amphibamus are as yet undecipherable.
THE LARVA
Considering the postembryonic developmental! stages of modern
Amphibia, there can be no doubt that a gill-bearing, four-legged
larva of a salamander, in which lateral line pores and a gular fold
are present, represents much more closely the type of larva found
in labyrinthodonts than does the limbless, plant-nibbling tadpole
of the Anura. Salamander-like larvae of labyrinthodonts are well
known, especially those formerly supposed to comprise the order
Branchiosauria. Many, perhaps the majority of, labyrinthodonts
show some features associated with aquatic life even when full-
Tue ANCESTRY OF MODERN AMPHIBIA iS
grown, as do the lepospondyls. These features may include impres-
sions of sensory canals on the dermal bones of the skull, persistence
of visceral arches, reduction in size of appendages, and failure of
tarsal and carpal elements to ossify. In fact, it appears that very
few of the Paleozoic Amphibia were successful in establishing them-
selves as terrestrial animals even as adults.
Nevertheless, in the ancestry of Anura, and that of at least the
Hynobiid, Ambystomid, Salamandrid and Plethodontid salamanders,
there must certainly have been a terrestrial adult, transforming from
an aquatic larva. The leaping mechanism of Anura, shown in so
many features of their anatomy, is perhaps to be explained as a
device for sudden escape from land into the water, but it was not
yet perfected in the Triassic Protobatrachus or the Jurassic Noto-
batrachus.
The middle ear, its sound-transmitting mechanism, and the tym-
panum, well developed in most Anura, are readily derived from
those of early labyrinthodonts, and are presumably effective for
hearing airborne sounds whether on land or while floating in the
water. Reduction of these organs in Urodela may be correlated
with their customary restriction to subsurface habitats and inability
to maintain a floating position while in water.
Some light may be shed on the significance of the tadpole of
Anura by considering the early stages of the ribbed frogs, Liopelmi-
dae. Leiopelma and Ascaphus are so closely similar in the adult
that there is no doubt that they belong
in one family, primitive in some re-
spects (including articulated ribs; py-
riformis and caudalipuboischiotibialis
muscles) but not in others (absence
of tympanum and middle ear). In
both genera the eggs are large, 5 mm.
in Leiopelma, 4.5 mm. in Ascaphus,
and unpigmented; but at this point
the resemblance ends.
Stephenson (1955) showed that em-
bryos of L. hochstetteri develop
equally well on land (in damp places )
or in the water, and that embryos
prematurely released from egg cap-
sules develop successfully in the wa-
ter. The larvae possess both pairs of Fis. 10. Leiopelma_hochstet-
teri larva, lateral and ventral
legs (Fig. 10) and a broad gular fold (after Stephenson, 1955), X 4.
Leiopelma hochstetteri
176 UNIVERSITY OF Kansas Pusxs., Mus. Nat. Hist.
similar to that of larval salamanders. In L. hochstetteri the fold
grows back over the forelegs temporarily, but remains free from
the body and presently the legs reappear, whereas in L. archeyi
the forelegs are not covered at any time. No branchial chamber
or spiracle is formed. Of course direct development, without a
tadpole, occurs in several other groups of Anura, but in each case
terrestrial adaptations are obvious. This is not true of Leiopelma,
which Stephenson regards as more nearly comparable with Urodela
in its development than with other Anura, and he sees in it a “pri-
mary and amphibious” mode instead of a terrestrial specialization.
The Ascaphus tadpole bears no outward resemblance to the larva
of Leiopelma, but is a normal tadpole in form, although sluggish
in activity. Its greatly expanded labial folds bear numerous rows
of horny epidermal “teeth,” which, with the lips, serve to anchor
the tadpole to stones in the swift water of mountain brooks. Noble
(1927) noticed that particles of food were taken in through the
external nares, and that a current of water passed through these
openings and out by way of the median spiracle. It appears that
any action by the teeth and jaws in scraping algae from the rocks
(which were bare in the stream where I have collected Ascaphus)
would be quite incidental, and that the lips and teeth must be
primarily a clinging mechanism. Certain other mountain brook
tadpoles (for example, Borborocoetes) show similar devices, but
these are developed independently, as specializations from the
usual sort of tadpole.
May it not be that closure of the gill-chamber by the opercular
(= gular) fold, retardation of limb development, expansion of the
lips, growth of parallel rows of horny teeth, and other correlated
features that make a tadpole, were brought about as an adapta-
tion of the primitive Anuran larva to a swift-stream habitat, and
that this “basic patent” then later served to admit the tadpoles of
descendant types to an alga-scraping habit in quiet water as well?
The tadpole, as a unique larval type among vertebrates, bears the
hallmarks of an abrupt adaptive shift, such as might have occurred
within the limits of a single family, and it seems difficult to imagine
the enclosed branchial chamber, the tooth-rows, and lips of a
familiar tadpole as having evolved without some kind of suctorial
function along the way.
THE ANCESTRY OF MopERN AMPHIBIA 177
SUMMARY
The Anura probably originated among temnospondylous laby-
rinthodonts, through a line represented approximately by Eugyrinus,
Amphibamus, and the Triassic frog Protobatrachus, as shown by
Watson, Piveteau and others. The known Paleozoic lepospondyls
do not show clear indications of a relationship with Urodela, but
Lysorophus may well be on the ancestral stem of the Apoda.
Between Urodela and Anura there are numerous resemblances
which seem to indicate direct relationship through a common stock:
(1) a similar reduction of dermal bones of the skull and expansion
of palatal vacuities; (2) movable basipterygoid articulation in
primitive members of both orders; (3) an operculum formed in the
otic capsule, with opercularis muscle; (4) many details of cranial
development, cranial muscles, and thigh muscles, especially be-
tween Ascaphus and the Urodela, as shown by Pusey and Noble;
(5) essentially similar manner of vertebral development, quite
consistent with derivation of both orders from Temnospondyli;
(6) presence in the larva of Leiopelma of a salamanderlike gular
fold, four limbs, and no suggestion of modification from a tadpole
(Stephenson ).
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1938. Dvinosaurus als neotenische Form der Stegocephalen. Acta Zool.,
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Case, E. C.
1935. Description of a collection of associated skeletons of Trimero-
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1889. The Batrachia of North America. Bull. U. S. Nat. Mus., 34:1-525.
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1937. The development of the vertebrate skull. Pp. xxiii + 552. Oxford,
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DEVILLIERS, C. G. S.
1934. Studies of the cranial anatomy of Ascaphus truei Stejneger, the
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1936. Die Morphologie der Tetrapodenrippen. Zool. Jahrb. (Anat.),
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1934. The anatomy of the salamander. Pp. xxxi + 381. Oxford, Clar-
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GAMBLE, D. L.
1922. The morphology of the ribs and the transverse processes of
Necturus maculatus. Jour. Morph., 36:537-566.
Gray, P.
1930. On the attachments of the Urodele rib to the vertebra and their
homologies with the capitulum and tuberculum of the Amniote rib.
Proc. Zool. Soc. London, 1930(1931):907-911.
Grecory, J. T.
1950. Tetrapods from the Pennsylvanian nodules from Mazon Creek,
Illinois. Am. Jour. Sci., 248:833-873.
Hecurt, M. E.
1957. A case of parallel evolution in salamanders. Proc. Zool. Soc. Cal-
cutta, Mookerjee Mem.:283-292.
Houtmcren, N.
1933. On the origin of the tetrapod limb. Acta Zool., 14:185-295.
1939. Contributions to the question of the origin of the tetrapod limb.
Acta Zool., 20:89-124.
1949a. Contributions to the question of the origin of tetrapods. Acta
Zool., 30:459-484.
1949b. On the tetrapod limb problem again. Acta Zool., 30:485-508.
HeERRE, W.
1935. Die Schwanzlurche der mitteleocanen (oberlutetischen) Braun-
kohle des Geiseltales und die Phylogenie der Urodelen unter
Einschluss der fossilen Formen. Zoologica, 33:87, 1-85.
Jarvix, E.
1942. On the structure of the snout of Crossopterygians and lower
gnathostomes in general. Zool. Bidrag fran Upsala, 21:235-675.
1952. On the fish-like tail in the Ichthyostegid Stegocephalians. Med-
delelser om Grgnland, 114(12):1-90.
MookeERJEE, H. K.
1930a. On the development of the vertebral column of the Urodela. Phil.
Trans. Roy. Soc. London, B 218:415-446,
1930b. On the development of the vertebral column of the Anura. Philos.
Trans. Royal Soc. London, B 219:165-196.
MacBriwg, E. W.
1932. Recent work on the development of the vertebral column. Cam-
bridge, Biol. Rev., 7:108-148.
McDowELL, S. B.
1958. Are the frogs specialized seymouriamorphs? (Abstract) Anat.
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NaeF, A.
1929. Notizen zur Morphologie und Stammesgeschichte der Wirbeltiere.
15. Dreissig Thesen iiber Wirbelsiule und Rippen, insbesondere
bei den Tetrapoden. Zool. Jahrb., 50:581-600.
Tue ANCESTRY OF MODERN AMPHIBIA 179
Nos eE, G. K.
1922. The phylogeny of the Salientia; I. The osteology and the thigh
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1927. The value of life-history data in the study of the evolution of the
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1934. Some points of view concerning the evolution of the vertebrates
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180 UNIVERSITY OF Kansas PuBLs., Mus. Nat. Hist.
SusHKIN, P. P.
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1940. The origin of frogs. Trans. Roy. Soc. Edinburgh, 40(7):195-231.
Transmitted April 7, 1959.
27-8362
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UNIVERSITY OF KANSAS PUBLICATIONS
MvusEUM OF NATURAL HISTORY
Volume 12, No. 3, pp. 181-216, 49 figs.
February 19, 1960
The Baculum in Microtine Rodents
BY
SYDNEY ANDERSON
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. 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 pou
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 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 figures in text. June 29, 1951.
Phylogeny of the waxwings and allied birds. By M..Dale Arvey. Pp. 473-
580, 49 figures in text, 18 tables. October 10, 1951.
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. 1-308, 73 figures in text,
87 tables. August 25, 1952.
2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry
Ba SE Lewis L. Sandidge. Pp. 305-338, 5 figures in text. August
The silky pocket mice ( Perognathus flavus) of Mexico. By~-Rollin H. Baker.
Pp. 339-347, 1 figure in text. February 15, 1954,
North American jumping mice (Genus Zapus ). By Phillip H. Krutzsch. Pp.
849-472, 47 figures in text, 4 tables. April 21, 1954.
Mammals from Southeastern Alaska. By Rollin H. Baker and James S.
Findley... Pp. 473-477. April 21, 1954.
Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479-
487. April 21, 1954.
Subspeciation in the montane meadow mouse, Microtus montanus, in Wyo-
pine and serene: By Sydney Anderson. Pp. 489-506, 2 figures in text.
y
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. 518-582, 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, i ee castanops, in Coa-
huila, Mexico. By Rob bert 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. 618-618. June 10, ffs
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)
BO eS
UNIVERSITY OF KANSAS PUBLICATIONS
MusEUM OF NATURAL HISTORY
Volume 12, No. 3, pp. 181-216, 49 figs.
February 19, 1960
The Baculum in Microtine Rodents
BY
SYDNEY ANDERSON
UNIVERSITY OF KANSAS
LAWRENCE
1960
Unrversity oF Kansas PUBLICATIONS, MUSEUM OF NATURAL History
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson
Volume 12, No. 3, pp. 181-216, 49 figs.
Published February 19, 1960
UNIVERSITY OF KANSAS
Lawrence, Kansas
qasarmer mene ’
PRINTED IN
THE STATE PRINTING PLANT
TOPEKA, KANSAS
1960
The Baculum in Microtine Rodents
BY
SYDNEY ANDERSON
INTRODUCTION
Didier (1943, 1954) has described the bacula of several Old
World microtines, and other rodents. Argyropulo studied (1933a,
1933b) five species of Cricetinae and Microtus socialis. Ognev
(1950) illustrated numerous species of Eurasian microtines. Hamil-
ton (1946) figured and described the baculum of 11 species of
North American microtines. Hibbard and Rinker (1942, 1943)
figured the baculum of Synaptomys cooperi paludis and of Microtus
ochrogaster taylori. Dearden (1958) studied the baculum in two
Asiatic species of Lagurus, in six subspecies of Lagurus curtatus of
North America, and in six other species of microtines of other genera.
The baculum can be preserved easily with standard study skins,
and is potentially useful in interpreting relationships on any taxo-
nomic level, and especially in determining the relationships of species
within a genus, if used together with other structures.
The anatomical orientation of the baculum needs comment be-
cause some confusion exists in the literature, especially concerning
the use of the terms ventral and dorsal. The urethra lies on the
anatomically ventral side of the penis, and of the baculum. In the
center of the penis lies a single corpus cavernosum penis, shown in
cross section proximal to the baculum in Figure lc. Dorsally an
artery, thinner walled than the ventral urethra, ends in a somewhat
reticulate sinus surrounding primarily the middle part of the bacu-
lum within the bulbous glans penis. The corpus cavernosum penis
(the structure has no median septum, at least distally) terminates
with the baculum and is closely knit to it. The site of this bond is
evident in the tuberosities and sculpturing of the base of the bacu-
lum.
The part of the penis enclosing the baculum, when not erect, is
folded back as shown in Figures la and 1b. As a result the anatomi-
cally ventral surface faces upwards, or at least posterodorsally.
The use of the term ventral in this account refers to the anatomically
ventral side, that is to say to the side of the baculum facing the
urethra.
The baculum in microtines consists of an elongate stalk, having
a laterally, and to a lesser extent dorsoventrally, expanded base and
(183)
184 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
an attenuate distal shaft. Usually, three digitate processes of carti-
laginous material in which additional ossifications may occur arise
from the terminus of the shaft. The proportions and curvature of
the stalk vary as do the proportions of the terminal ossifications to
each other and to the stalk. In some species one or more of the
digital processes are frequently completely unossified.
ES
IN MILLIMETERS
LENGTH OF STALK OF
BACULUM
120 130 140 150 160 170
LENGTH OF ANIMAL IN MILLIMETERS
FicurE 1. The baculum in Microtus ochrogaster—orientation and variation
with age. a. Diagram of a sagittal section of the posterior half of a vole, natural
size. The penis, containing the baculum (in black), extends ventrally from a
point posterior to the pubic symphysis (stippled), along the body wall, and
bends posteriorly at the distal end. b. Distal end of penis (2) showing
baculum (in black), the urethra (solid lines) adjacent to the baculum, and
the corpus cavernosum (broken lines) proximal to the baculum. c. Oblique
view of the cross section of penis (> 4) shown in Figure 1 b. The thick-walled
urethra lies ventral to the curved corpus cavernosum. A thinner-walled blood-
vessel lies dorsal to the corpus cavernosum. The anatomically ventral side of
the baculum, in the normal non-erect penis shown, is seen to face dorsally.
d. Graph showing the relationship between size of baculum, size of animal,
and development of digital ossifications. Circles show presence of ossification
in stalk only; circles enclosing dots indicate presence of secondary ossification
in median process also; large dots indicate the addition of tertiary ossification
in one or both of the lateral digitate processes.
THE BACULUM IN MICROTINE RODENTS 185
Preserved specimens of Microtus arvalis, Microtus agrestis, Microtus orca-
densis, Microtus nivalis, Microtus guentheri, Microtus subterraneus, Cleth-
rionomys glareolus, and Ellobius lutescens were provided by Prof. Robert
Matthey of Lausanne, Switzerland. J. Knox Jones, Jr. carefully saved the
bacula with specimens of Microtus fortis and Clethrionomys rufocanus from
Korea. Dr. W. B. Quay, Department of Zoology, University of California,
supplied specimens of Synaptomys cooperi, Phenacomys intermedius, and
Microtus oregoni. Dr. Franklin Sturges and Mr. John W. Goertz, Museum
of Natural History, Oregon State College, Corvallis, have provided specimens
including bacula of Clethrionomys occidentalis, Microtus oregoni, and Microtus
townsendii. Dr. Randolph L. Peterson and Mr. Bristol Foster, Royal Ontario
Museum of Zoology, Toronto, Canada, provided specimens of Phenacomys inter-
medius. Dr. J. N. Layne, University of Florida, Gainsville, Florida, presented
me with a baculum of Microtus parvulus.
I am indebted to all of these persons for their aid, and to various collectors
for the Museum of Natural History, who preserved bacula with specimens.
Many of these specimens were obtained through the assistance of the Uni-
versity of Kansas Endowment Association and the National Science Foundation.
METHODS
Bacula were obtained from fresh specimens, specimens preserved in alcohol
or formalin, and dried study skins. The processing of bacula has been dis-
cussed by Hamilton (1946), Friley (1947), White (1951), and Dearden
(1958). The methods used to preserve bacula for my study differed some
from any of those reported. The terminal part of each penis including the
baculum imbedded in the glans penis was removed in its entirety and placed
in a vial. The catalogue number was kept with each specimen at all times.
A two per cent solution of potassium hydroxide was added. All specimens
were examined at least once a day. If tissues other than the glans penis were
present they were removed with forceps when softened usually at the end of
one day. Several drops of Alizarin red-S stain in a saturated alcoholic solution
were added to the 3 to 5 ccs. of KOH solution in each vial. Solutions were
replaced if they became turbid enough to obstruct observation of the clearing
penis. After one day the solution containing stain was removed and replaced
with two per cent KOH solution without stain. When the glans became
sufficiently cleared that the stained baculum could be seen easily, the solution
was replaced by glycerin in which clearing was completed. The time re-
quired for the entire process varied from one day to more than two weeks
depending on the size of the specimen and on its condition. Fresh specimens
clear more rapidly than dried specimens, and those that are dried more rapidly
than those that are preserved. A three or four per cent soluticn of hydroxide
will hasten the process, but more frequent observation is required to prevent
excessive maceration.
Specimens were then examined in a shallow dish containing glycerin under
a binocular microscope. The baculum can be viewed from any desired direc-
tion. The method described above leaves the baculum intact within the glans
penis; therefore its orientation can be determined relative to the thick walled
urethra and the thin walled dorsal artery that extends onto the dorsal side of
the baculum. The ventral curvature of the penis proximal to the baculum,
and the distal extension, characteristic of most species, of the dorsal border of
186 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
the glans (both shown in Figure 1) are other features aiding in correctly
orienting cleared specimens. The digitate processes are not so often injured,
lost, or displaced when the method described above is used as they are when
the penis is dissected. Specimens were stored in glycerin in glass shell vials
having polyethylene stoppers. A small card bearing the name, number, locality,
and other data was placed in each vial. A specimen thus enclosed can be
kept indefinitely, or removed and mounted in balsam as described by White
(1951:631) or in plastic as described by Dearden (1958:541) and thus stored
in the vial containing the skull of the specimen.
Drawings were made on millimeter ruled paper while the baculum was
viewed under a binocular microscope with a square ruled eyepiece.
Unless otherwise noted all specimens listed are in the University of Kansas
Museum of Natural History. Catalogue numbers are cited. Measurements
are accurate to within less than one-tenth of a millimeter. Proportions as stated
in the text are approximations, accurate to within one-twelfth (8.33 per cent).
The range of variation is unknown for some species. Mention is made if
maturity is known or suspected to differ in specimens being compared.
The development of the baculum has been studied by Callery
(1951) in Mesocricetus auratus and by Ruth (1934) in the lab-
oratory rat. In the rat (Rattus norvegicus) the bone is of endob-
lastemal origin being laid down by a condensation of undifferentiated
mesenchymal cells. At the distal end of the bone dense fibrous
tissue is then differentiated and at the proximal end hyaline cartilage.
Growth is by substitution at the proximal end and by subperiosteal
lamellation circumferentially. A marrow cavity is formed by re-
sorption. In the baculum of the hamster the primary center of ossi-
fication is in the stalk, and is present at the age of three days; the
secondary centers are in lateral processes and are present at 80 days
and enlarge subsequently. A tertiary center, in each median process,
may or may not develop later. Maximum development of the
baculum is reached late in the reproductive life of the hamster.
The early ossification of the baculum noted in the rat and the
hamster occurs in Microtus also. A specimen of Microtus montanus
fusus (76831, from 5 mi. N, 26 mi. W Saguache, 9600 ft., Saguache
County, Colorado) only 74 mm. in total length and weighing only
6.6 grams, had a slender ossified baculum having enlarged ends.
This vole was one-half of the average length and less than one-fifth
of the average weight of an adult, and of approximately the size at
which weaning takes place.
The development of the baculum in Microtus ochrogaster was studied in
$2 specimens of various ages. The specimens (between Nos. 74994 and 75074)
were collected between August 15 and September 4, 1957, at localities on the
Great Plains. These specimens were from breeding populations, as evidenced
by pregnancy of females and by large size of testes of males. The length and
width of the stalk of the baculum, the presence of digital ossifications, the
THE BACULUM IN MICROTINE RODENTS 187
total length of the animal, and the size of the testes were noted. Variability in
length of testes is greatest when voles are from 140 to 150 mm. in total length.
Sexual maturity is reached rather abruptly when the total length of most indi-
viduals is 140 to 150 millimeters. If the baculum likewise underwent more
rapid growth at the onset of sexual maturity, greater variability should be evi-
dent in the length of the baculum of voles 140 to 150 mm. in total length than
in bacula of voles of other sizes. This was the case (see Figure 1d). The
baculum does not, however, suddenly reach its maximum maturity.
The primary ossification is in the stalk. The secondary ossification
is in the median process except in Lagurus (Dearden, 1958:551) and
some individuals of Neofiber (see account on page 258). Tertiary
centers of ossification are in the lateral processes. The primary
ossification is present at an early age and subsequently increases in
size and solidity. The secondary and tertiary ossifications are pro-
gressively more common in older voles. The increase in degree of
ossification of all parts continues after sexual maturity is reached.
Individual variation and variation with age in the baculum of Micro-
tus pennsylvanicus have been illustrated by Hamilton (1946:380).
Figures 14, 15, and 17 illustrate variation with size, which is corre-
lated with age, and also illustrate individual variation. The three
bacula are from adult voles having testes that measured 15, 16 and
16 mm. in length, respectively. Each vole was trapped in late June.
The total lengths in millimeters of the three voles are 172, 167, and
181; weights are 55, 52.4, and 65.5 grams. I judge that the greater
size of the stalk and the better developed base shown in Figure 17
than in Figure 15 are illustrative of age variation; the difference in
the size of the lateral digitate processes is, in this case, attributable
to individual variation. Differences in the distal end of the baculum
in Figures 42 and 48, show individual variation also. Figures 35
and 36 represent two different subspecies; different individuals of
M. mexicanus mogollonensis, however, exhibit individual variation
of the same degree.
Hall and Cockrum (1953) list 44 species of microtines in North
America. At least twelve of these are insular or local forms perhaps
derived from some other species; for example Microtus coronarius,
an insular form derived from Microtus longicaudus; Microtus pro-
vectus, considered by Chamberlain (1954:587) and by Wheeler
(1956:176) as a subspecies of Microtus pennsylvanicus; and Micro-
tus ludovicianus, a close relative of Microtus ochrogaster.
All North American genera have been studied. Of the genus Microtus in
North America, all subgenera but Orthriomys and all species but the follow-
ing nine, have been studied: M. (Orthriomys) umbrosus, the insular M. (Steno-
cranius) abbreviatus, M. (Microtus) breweri, M. (Microtus) nesophilus, M.
188
UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
a
oC
Q
oO
Q
o
Q
Colt Face On adstie See
Ficures 2-13. Bacula of microtines. Unless indicated otherwise views are
(a) of the dorsum, (b) the right side, and (c) the proximal end with the dorsal
surface upward. Exact localities are given in accounts of species concerned.
Lemmus trimucronatus, 50678, Point Barrow, Alaska.
POs enus groenlandicus, 50539, Porcupine Lake, Brooks Range,
Alaska.
Dicrostonyx groenlandicus, 52524, Point Barrow, Alaska.
Synaptomys cooperi saturatus, WBQ 3-C-454, 3 mi. S Demotte, Indiana.
Synaptomys cooperi paludis, 13716, Meade County State Park, Kansas.
Phenacomys intermedius celatus, SA 2044, Quebec.
Phenacomys intermedius intermedius, WBQ 3-C-309, 5.4 mi. S Moran,
Teton Co., Wyoming.
Clethrionomys rufocanus, 60438, 1 mi. NW Oho-ri, Korea, (d) ventral
view.
Clethrionomys gapperi, 42108, 31 mi. N Pinedale, Wyoming.
Clethrionomys rutilus, 42865, 5 mi. NNE Gulkana, Alaska.
Clethrionomys occidentalis, FWS 30, Mary’s Peak, Benton Co., Oregon.
Clethrionomys glareolus, 67100, Zermatt, Valais, Switzerland.
THE BACULUM IN MICROTINE RODENTS 189
id \\\ 4 Uy, Wy
i uy
Ficures 14-25. Bacula of Microtus. Unless indicated otherwise views are
(a) of the dorsum, (b) the right side, and (c) the proximal end with dorsal
surface upward.
14,
15.
M. pennsylvanicus, 42439, 1 mi. S, 2 mi. E Eagle Nest, Colfax Co.,
New Mexico; abnormality | perhaps owing to injury; dorsal view.
M. pennsylvanicus, 42306, 5 mi. N, 26 mi. W Saguache, Colorado;
dorsal view.
M. pennsylvanicus, 43043, 20 mi. NE Anchorage, Alaska, ventral view.
M. pennsylvanicus, 42430, 1 mi. S, 2 mi. E Eagle Nest, New Mexico.
M. agrestis, 67102, Gryon, Switzerland.
ue sae eees amosus, 62241, % mi. E Soldier Summit, Wasatch Co.,
ta
. M. montanus nanus, 57470, 2 mi. N, 2 mi. W Pocatello, Idaho.
. M. montanus fusus, 42307, 5 mi. N, 26 mi. W Saguache, Colorado.
. M. arvalis, 67101, Vidy, Switzerland, possibly not mature.
5 We guentheri, 67104, Palestine.
M. orcadensis, 67106, Orkney Islands, orientation uncertain.
M. fortis, 63841, Chipo-ri, Korea, (d) ventral view.
190 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
0
log
Ficures 26-39. Bacula of microtines. Unless indicated otherwise views are
(a) of the dorsum, (b) the right side, and (c) the proximal end with the
dorsal surface upward.
26. M. (Pitymys) fatioi, 671038, Zermatt, Switzerland, immature.
27. M. (Pitymys) pinetorum, 76834, 2 mi. N Baldwin, Douglas Co., Kansas.
28. M. (Pitymys) pinetorum, 68545, 1 mi. NE Pleasant Grove, Kansas.
29. M. (Pitymys) quasiator, 30709, Teocelo, Veracruz, (d) ventral view.
30. M. (Pitymys) quasiator, 19878, 5 km. N Jalapa, Veracruz.
81. M. (Pedomys) ochrogaster, 75036, 1 mi. N, 2 mi. E Oberlin, Kansas.
82. M. (Stenocranius) miurus, 51152, Lake Schrader, Brooks Range, Alaska.
83. M. (Stenocranius) miurus, 51169, Lake Schrader, Brooks Range, Alaska.
34. M. (Stenocranius) gregalis, 8059, ‘“‘Eastern Europe.”
85. M. mexicanus mexicanus, 63094, Valle de Bravo, Estado de México, México.
836. M. mexicanus mogollonensis, 63298, Mt. Taylor, Valencia Co., New Mexico.
87. M. californicus, 76828, 1 mi. NE Berkeley, California; (d) ventral view.
88. M. (Arvicola) richardsoni, 42454, 31 mi. N Pinedale, Sublette Co., Wyoming.
39. M. richardsoni, 37903, 23% mi. S, 5 mi. W Lander, Wyoming; distal end.
THE BACULUM IN MICROTINE RODENTS 191
c 49
Ficures 40-49. Bacula of microtines. Unless indicated otherwise views are
(a) of the dorsum, (b) the right side, and (c) the proximal end with the dorsal
surface upward.
Microtus (Pitymys) parvulus, UF 1508, 1 mi. W Micanopy, Florida.
Microtus townsendii, 79186, Sec. 33, T. 11S, R. 5W, Benton Co., Oregon.
Microtus (Herpetomys) guatemalensis, 65895, 2 mi. S San Juan Ixcoy, Guatemala.
ae guatemalensis, 65921, 10 mi. E, 4 mi. S Totonicapan, Guatemala, dorsal view
of tip.
Microtus oeconomus, 43048, Kelsall Lake, British Columbia.
Microtus (Chilotus) oregoni, WBQ 3-C-248, 5 mi. N Orick, California.
. Lagurus (Lemmiscus) curtatus, 26053, 9 mi. S Robertson, Uinta Co., Wyoming.
Microtus (Chionomys) nivalis, 65127, Wetterstein, Germany, orientation uncertain,
Microtus (Chionomys) longicaudus, 50253, Crane Flat, Mariposa Co., California.
. Neofiber alleni, 27268, 2 mi. S Gainesville, Florida, orientation uncertain.
192 UNIVERSITY OF KAnsAs Pusts., Mus. Nat. Hist.
(Microtus) provectus (the last three are probably insular derivatives of M.
pennsylvanicus), M. (Microtus) fulviventer (perhaps derived from the same
stock as Microtus mexicanus), M. (Microtus) xanthognathus (perhaps related
to Microtus chrotorrhinus), M. (Microtus) coronarius, and M. (Pedomys)
ludovicianus.
SpEcIES OF WuicH BACULA WERE EXAMINED
Subfamily: Microtinae Number of
Tribe: Lemmi Specimens
Dicrostonyx groenlandicus*(Traill)... 2... .2. 022.52 s.0226+-0- 4
Lemmus trimucronatus (Richardson) .................0ee08: 6
Sunaptomys coopen: Baird |... ste. Meer. oe Oct aisle ae 5
Tribe: Microti
Genus: Clethrionomys Tilesius, 1850
Clethrionomys rutilus Pallas”, 4.4000. + -2 oes on oe eee
Clethrionomys gapperi (Vigors) 0.0.5.0. nee eee ewe ee
Clethrionomys occidentalis (Merriam) ..................-
Clethrionomys glareolus Schreber ................---000-:
Clethrionomys rufocanus Sundevall ...................++
Genus: Phenacomys Merriam, 1897
Phenacomys intermedius Merriam ...............22.--:-:
Genus: Ondatra Link, 1795
Ondatra-zibethicus (Acinnaeus) .) 4h Aos.. 4: SS ..2 Sette «ihe
Genus: Microtus Schrank, 1798
(Herpetomys) guatemalensis Merriam ..................-
(Arvicola) richardsoni: (DeKay)-% .0..<6 cals cues eee nee
(Chilotus) oregont (Bachman)’ ...05...<... «5. .f0u- oo
(Stenocranius) gregalis (Ballas) 2.0... .2 2: sae ue) ae oe:
(Stenocranius) miurus Osgood ............000000 neers eee
(Chionomys) longicaudus (Merriam) ...............-.++.
(Chionomys) nivalis: Martins)=.5:. .2 0). be 2 Gh 4. sen dseas
(Macrotis)sarealis (Pallas): 3) e893. ae Foe eo 32 Oe:
(Microtus) ‘orcadensisMillaist (.- i005) ve babes wale chet ees
(Microtus) guentheri Danford and Alston.................
(Mecrotus) fortis Buchner (364.5 oo.c6-ds.c fe ve eben eee
(Microtus) montanus: (Peale) 2252 6 i 3jein ots on iene tee Bae 1]
(Microtus) townsendii (Bachman) ...................0-
(Microtus) ceconomus’\(Pallas) . . . och. whens sles be leo: 10
(Microtus) mexicanus (Saussure) ............000-0020085 8
(Microtas) ‘californicuss (Peale) 4.5 o).6.5 62s Sepoehto ve eee 2
(Microtus) pennsylvanicus (Ord) ......0.....0.00c ec eens 13
(Microtus) agrestis “(lainnaeus) iis. 24. 0 es Bee eee i
(Pedomys) ochrogaster (Wagner) ..............22+se008- 41
(Pitumuys) spinetorum. GueConte) i. «05 ao acne eee eee 2
(Pitymys) naroulus. CHowelll)\s (003 «ahi dicokihe Be ee oe 1
(Pitumys). quasiater: (Goues))) cs wae « ceeded AAs ne eeu ke 5
(Ritymeys) cfatiOt NOH AZ A scant: ate.o soso ob seeder anos il
Genus: Neofiber True, 1884
Neofibertallentilrdee.: shy prec. ae Ae ces Rhee 2
Genus: Lagurus Gloger, 1841
Eagurusicurtatus’ (Cope)! 2846 oo ek ce ee es uf
‘Total mumbervexamined so 2 20. 10../aeee n e 184
WUD RB BN QGOrFWDNC Mm CO BRO,
THE BAacuLUM IN MICROTINE RODENTS 193
ACCOUNTS OF SPECIES
Dicrostonyx groenlandicus (Traill)
Figs. 8 and 4
Baculum: stalk elongate, greatest length (3.1 mm.) 2% to 2% times greatest
breadth, and 4% times greatest depth; digitate processes usually cartilaginous,
occasionally lateral processes partly ossified; basal tuberosities weakly to mod-
erately developed, medially confluent; posterior profile in dorsal view rounded
with rounded posterior apex or shallow notch; dorsal concavity in end-view
shallower and not so wide as ventral concavity; median constriction approxi-
mately 24 greatest depth; ventral part of base in end-view wider than dorsal
part; shaft straight or slightly curved; base of stalk placed dorsally relative to
axis of shaft; stalk spatulate, sometimes with distal enlargement; at mid-point
stalk wider than high; lateral profile in dorsal view sloping gradually without
abrupt curvature anterior to point of greatest width.
The baculum of Dicrostonyx torquatus figured by Ognev (1948:476) agrees
with that of D. groenlandicus in shape of stalk, and in lateral digitate processes
that are small relative to size of median process; but differs in more elongate,
terminally enlarged, bulbar shape of median process. None of my specimens
showed ossification in the lateral processes, observed by Hamilton (1946:381)
in Dicrostonyx rubricatus richardsoni [= D. groenlandicus richardsoni]. In all
of my specimens the cartilaginous median process was larger than that figured
by Hamilton, or by Dearden (1958:542).
Specimens examined: Four from; Point Barrow, Alaska, 52524 (Barrow
Village), 67264 (died in captivity); Brooks Range, Alaska, 50536 (Wahoo
eee 69°08’, 146°58’), 50539 (Porcupine Lake, 68°51'57”, 146°29’50”, 3140
Lemmus trimucronatus (Richardson)
Fig. 2
Baculum: Stalk heavy, broad, greatest length (2.8 mm.) in mature indi-
viduals (Fig. 2) as little as 1% times greatest breadth, greatest length no less
than 234 times greatest depth of base; three ossified processes, median one
from as long as to % longer than the lateral processes, and approximately %
wider and twice as deep as lateral processes; length of median process almost
8% times its breadth, approximately % length of stalk; basal fossae broadly
confluent; posterior profile in dorsal view evenly rounded; in end-view ventral
concavity deeper than dorsal concavity, constriction as little as % greatest depth
in mature specimens; shaft straight, bluntly rounded, or slightly decurved and
laterally inflated terminally; lateral profile in dorsal view a gradual slope from
widest point of stalk anteriorly onto shaft; in younger individuals stalk slenderer,
otherwise as described above.
Five specimens examined by me differ from one figured and described by
Hamilton (1946:379) in that stalk is better developed, larger relative to size
of processes, length of stalk in my specimen (Fig. 2) 2.8 as opposed to 2.1 mm.
in Hamilton’s specimen; median process shorter, 1.5 as opposed to 1.8 mm.,
proximal end rounded rather than concave, not partially enclosing tip of shaft;
proportion of and relative sizes of median and lateral processes approximately
same as in Hamilton’s Lemmus helvolus [= Lemmus trimucronatus helvolus].
194 UNIVERSITY OF KANSAS PuBts., Mus. Nat. Hist.
A specimen figured by Dearden (1958:542) has a basally trilobed median
process.
The baculum of the Asiatic Lemmus lemmus figured by Ognev (1948:413)
agrees with my specimens in the ossification of three processes, the relative
sizes of these processes to each other and to the stalk, the well-developed base
of the stalk and heavy bluntly rounded shaft; the baculum of Lemmus lemmus
differs in greater anterolateral extent of basal tuberosities, in proximal notch
seemingly separating these tuberosities, and in median process being slenderer.
Specimens examined; Five, of two subspecies; Lemmus trimucronatus alas-
censis, Point Barrow, Alaska, numbers 50591, 50678, 50731, 50758; Lemmus
trimucronatus subarcticus, Wahoo Lake, 69°08’, 146°58’, 2350 ft., Brooks
Range, Alaska, 50948.
Synaptomys cooperi Baird
Figs. 5 and 6
Baculum: Stalk elongate, greatest length (2.7 to 2.8 mm.) 2% to 2% times
greatest breadth, 4 to 5 times greatest depth; three processes ossified or lateral
processes unossified, ossifications relatively small (in 78380, median ossification
less than % as large as lateral ossifications although median cartilaginous process
is larger), length of median process 4% to % of length of stalk, cartilaginous
part of median process larger; posterior profile in dorsal view convex throughout
or bilobate; tuberosities moderately developed, deflected dorsal to axis of
shaft; in end-view medial construction 84 greatest depth of tuberosities; shaft
tapered from point of greatest width, slightly inflated terminally.
The specimen (KU 13716) figured by Hibbard and Rinker (1942:29) has
been restudied. It was first cleared and stained to soften the dry cartilage
binding the digital processes together and to differentiate bone and cartilage.
The lateral processes are small and cartilaginous (Fig. 6) and seem intact. The
differences between this specimen and others examined by Hamilton (1946:
881), Dearden (1958:542), and myself, namely the relatively larger median
ossification, the absence of ossification in lateral processes, and the distinctly
bilobate base and larger size, may represent geographic differences, or individual
‘variation. The proportions of length, width, and depth of the stalk, and the ap-
pearance in lateral view do not differ greatly from others examined by Hamilton,
by Dearden (1958:546), and by me.
Specimens examined: Five, representing four subspecies; S. cooperi gossii,
6 mi. N Midway, Holt Co., Nebraska 78379, 78380; S. cooperi relictus, 5 mi.
N, 2 mi. W Parks, Dundy Co., Nebraska, 72601 (immature); S. cooperi satur-
atus, 3 mi. S Demotte, Jasper Co., Indiana, 3-C-454, collection of W. B. Quay;
S. cooperi paludis, Meade County State Park, Kansas, 13716.
Clethrionomys rutilus Pallas
Fig; 11
Baculum: Stalk elongate, and proximally enlarged, greatest length (2.7 mm.)
2 times greatest breadth; less than 4 times greatest depth; three well-developed
ossified processes; length of stalk 214 times length of median process; median
process with basal (and ventral) protuberence and lateral lobes, arched in
dorsoventral plane; lateral processes as large as median process, flattened distally,
having ventromedial vane on distal half; basal tuberosities of stalk well devel-
oped, medially confluent; posterior profile in dorsal view trilobate or convex
THE BACULUM IN MICROTINE RODENTS 195
throughout with rounded posterior apex; dorsal concavity well developed,
ventral surface but slightly concave, medial constriction of base as little as %
greatest depth; shaft straight, slender, at mid-point of stalk but slightly wider
than high; basal tuberosities largely dorsal to axis of shaft in lateral view;
lateral profile in dorsal view with an abrupt curvature separating the gently
sloping sides of the shaft from the basal part at its greatest breadth.
The specimen of Clethrionomys rutilus figured by Ognev (1950:120) is
essentially like the North American specimens examined by me in the relative
sizes of the ossifications and the general shape of the stalk.
Specimens examined: Four, of one subspecies; C. r. dawsoni, west bank
Gakona River, 1700 ft., 5 mi. NNE Gulkana, Alaska, 42865, 42866; SW end
Dezadeash Lake, 2400 ft., Yukon Territory, 42910, 42921.
Clethrionomys gapperi (Vigors)
Fig. 10
Baculum: Stalk elongate, greatest length (2.8 mm.) 1% times greatest
breadth, and 3% times greatest depth; proximally enlarged, greatest depth
% greatest breadth; three well-developed ossified processes; length of stalk
21% times length of median process; median process arched in dorsoventral
plane, with basiventral protuberence or spine and lateral lobes; lateral processes
as large as median process, flattened distally, arched; basal tuberosities of
stalk well developed, medially confluent; posterior profile in dorsal view
trilobate or convex throughout with a rounded posterior apex; dorsal concavity
well developed, ventral surface but slightly concave, or in some cases slightly
convex; medial constriction of base 34 greatest depth; shaft straight, slender,
at mid-point of stalk twice as wide as high; basal tuberosities dorsally placed
relative to axis of shaft; lateral profile in dorsal view abruptly curved anterior to
point of greatest width; slender stalk distinct from angular enlarged base.
The most noticeable difference between the baculum of C. rutilus and C.
gapperi is size. The proportions of the four ossifications are approximately the
same. Ventral vanes on the lateral processes are not developed in C. gapperi.
C. gapperi and C. rutilus are more nearly alike in their bacula than any other
two species of Clethrionomys examined. Clethrionomys occidentalis, the other
New World species, is also much like C. gapperi and C. rutilus. The differ-
ences are of a magnitude comparable to those between the bacula in subspecies
of Microtus montanus (Figs. 19-21) for example, or in subspecies of Lagurus
curtatus (Dearden, 1958:542).
Specimens examined; Nine, of two subspecies; Clethrionomys gapperi atha-
bascae, British Columbia, 42922 (Indian Creek, Mile Post 234 of Alaskan High-
way), 64281 (West bank Racing River, 89 mi. W Muskwa), 64287 (North
bank Tetsa River, 56 mi. W, 11 mi. S Muskwa), 64290 (44 mi. W, 9 mi. S
Muskwa), 64310 (32 mi. W, 2 mi. S Muskwa); Clethrionomys gapperi galci,
31 mi. N Pinedale, Sublette Co., Wyoming, 42108; Grand Mesa, Delta Co.,
Colorado, 60014 and 60015 (5% mi. E, 12 mi. S Collbran), 60022 (8 mi. E, ¥%
mi. S Skyway).
Clethrionomys occidentalis (Merriam)
Fig. 12
Baculum: Stalk elongate, greatest length (2.8 mm.) 2% times greatest
breadth, 6 times greatest depth; three well-developed ossified processes; me-
dian process larger than lateral processes, % the length of stalk, curved, basally
196 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
broad, ventrally keeled, trilobate posteriorly; lateral ossifications large, flattened
distally, curved; posterior profile of stalk posteriorly slightly emarginate, thus
bilobate in outline; in end-view dorsal concavity deeper than ventral, con-
striction less than % greatest depth, tuberosities confluent, visible in dorsal view
at each side; shaft slender, especially in depth, straight; at mid-point of stalk
almost twice as wide as deep, slight terminal inflation.
The general proportions of the stalk and the relatively large, uniquely shaped
processes, are characteristic of most specimens of the genus Clethrionomys
examined.
Specimen examined: C. occidentalis californicus, one from Mary’s Peak,
Benton Co., Oregon, 30, F. W. Sturges’ collection.
Clethrionomys glareolus Schreber
Fig. 18
Baculum: Stalk elongate, greatest length (2.9 mm.) twice the greatest
breadth in the specimen examined, flattened proximally, greatest length almost
6 times greatest depth of base; three well-developed ossified processes; median
process arched in a dorsoventral plane, with basal notch and lateral lobes;
lateral processes as long as median process, bowed in dorsal view, flattened
distally, with ventromedial vane; basal tuberosities of stalk weakly developed,
medially confluent; posterior profile in dorsal view evenly rounded; in end-view
dorsal concavity shallow in comparison to most species but deeper than ventral
concavity, constriction % greatest depth; shaft straight, at mid-point slightly
wider than high, elongate, widest point of stalk less than % of total length from
proximal end, slight lateral inflation at tip; lateral profile in dorsal view sloping
at first abruptly and then gradually from widest point of stalk anteriorly onto
shaft.
The specimen of Clethrionomys glareolus figured by Ognev (1950:31) in
dorsal view as I interpret it, resembles my specimen in the rounded base; in
the elongate, distally inflated shaft; in the initially abrupt slope of the lateral
profile in dorsal view from the greatest width of stalk anteriorly; and in the
presence of three well ossified processes. Ognev’s specimen differs from mine
in the median process being more elongate relative to its width, and rounded
proximally, lacking lateral lobes and basal notch; in lateral processes being less
curved; in the greater terminal inflation of the shaft; and in the closer approxi-
mation of the terminal processes to the shaft. The baculum of Clethrionomys
glareolus as described and figured by Didier (1954:243-244) resembles my
specimen in general proportions, but is more pointed proximally and more
curved in dorsoventral plane. Didier states that the baculum is rather variable
in form in this species, in different regions, but that a large number of specimens
must be examined to assess the geographic nature of this variation.
Specimen examined: One from Zermatt, Valais, Switzerland, 67100.
Clethrionomys rufocanus Sundevall
Fig. 9
Baculum: Base of stalk broad but relatively flattened dorsoventrally, greatest
length (3.2 mm.) less than 1% greatest width, 4 times greatest depth; three
well-developed ossified processes; median process arched in dorsoventral plane,
having basal notch and lateral lobes; lateral processes as long as median process,
THe BACULUM IN MICROTINE RODENTS 197
flattened distally, with ventromedial vane; basal tuberosities of stalk weakly
developed, medially confluent; posterior profile in dorsal view convex with
rounded posterior apex; dorsal surface of base almost flat, ventral concavity
broad and shallow; constriction % greatest depth (not including an unusual
irregularity on the ventral surface of the base); shaft straight, at mid-point of
stalk distinctly wider than high, slender at distal end, widest point of stalk
almost 14 of total length from proximal end, tip of shaft rounded; lateral profile
in dorsal view gradually sloping from widest point anteriorly onto shaft.
The specimen of Clethrionomys rufocanus figured by Ognev (1950:97)
resembles my specimen in the presence of three well ossified processes. Ognev’s
specimen differs however in the lack of a proximal notch on the median process,
the lesser proportion of the stalk included in the basal enlargement, the more
posterior position of the point of greatest width, and the presence of a concavity
in the posterior profile of the stalk in dorsal view. These differences in the
stalk may be owing to a difference in age (my specimen perhaps being older).
Specimen examined: One from 1 mi. NW Oho-ri, 6 M., Korea, 60438.
Phenacomys intermedius Merriam
Figs. 7 and 8
Baculum: Stalk slender, greatest length (2.9 mm.) 2% to 2% times greatest
breadth, 4 times greatest depth; three well-developed ossified processes, median
one almost % length of stalk, curved, broad basally and slightly larger in all
dimensions than either lateral process; lateral processes flattened distally, curved;
base of stalk well developed, basal tuberosities medially confluent or separated
by medial emargination, posterolateral faces flattened or rough; emarginations
in the four adults examined; posterior profile in dorsal view bluntly pointed or
flattened except for emargination posterially, abruptly curved at point
of greatest width; shaft arising broadly from distal side of base of stalk; in
end-view hour-glass shaped, medial constriction pronounced, both dorsal and
ventral concavities deep; shaft having relatively straight but distally convergent
sides; at mid-point of stalk, 1 to 1% times as wide as deep; tip bluntly rounded,
or slightly inflated.
The specimens from Quebec differ from the one from Wyoming in smaller
size, relatively smaller lateral digital processes, larger more medial basal
emargination, and slender shafts. The baculum of Phenacomys intermedius
differs much from that of Phenacomys longicaudus, described by Hamilton
(1946:381) and by Dearden (1958:547). Dearden states that the three
bacula examined by him of Phenacomys longicaudus differ markedly from the
specimen described by Hamilton. It seems to me that in major features the
resemblance is greater between the specimens of Phenacomys longicaudus
examined by these two authors than between their specimens and specimens
of other microtines, including Phenacomys intermedius. Neither Hamilton nor
Dearden record the exact localities of capture, the collections in which the
specimens are deposited, or the catalogue numbers of specimens. Consequently
verification of identifications and observations is difficult.
Specimens examined: Five, of two subspecies; P. intermedius intermedius,
5.4 mi. S Moran, Teton Co., Wyoming, 3-C-309, collection of W. B. Quay;
P. intermedius celatus, four (including one immature specimen) from Authier-
2—T74
198 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist.
nord, Abitibi-ouest Co., Quebec, specimens in collection of Bristol Foster desig-
nated by numbers 2041-2044 of S. Anderson’s field catalogue. Smith and
Foster (1957:107) were of the view that Phenacomys ungava (including the
above specimens from Quebec) may be specifically distinct from Phenacomys
intermedius.
Ondatra zibethicus (Linnaeus)
Not figured
Baculum: In the single specimen examined, less mature than that figured
by Hamilton (1946:384), the digitate processes are cartilaginous, the basal
tuberosities are less well developed, and the shaft is slenderer throughout. The
cartilaginous processes are of the same proportions as ossified processes in the
figure mentioned. The shaft is also convex ventrally in lateral profile. The
view of the side here considered to be anatomically the ventral side (adjacent
to the urethra) is labelled dorsal view in Hamilton’s specimen.
Specimen examined: One, from Reserve, Brown Co., Kansas, 72405.
Microtus (Herpetomys) guatemalensis Merriam
Figs. 42 and 43
Baculum: Stalk moderately elongate, greatest length (3.5 mm.) 214 times
greatest breadth, spatulate, flattened throughout, greatest thickness 14 milli-
meter; three ossified processes; median process having three cornered base,
curved dorsally, wider than high, % to 14 greatest length of stalk; each lateral
process bent at middle, as long as median process, compressed laterally; base
of stalk curved dorsally, tuberosities marginal, hence narrow, lateral excavations
of tuberous margin not confluent medially; in end-view ventral concavity broad,
no dorsal concavity, medial constriction but slightly less than greatest thick-
ness (not depth); shaft wider than high throughout, at mid-point more than
3 times as wide as high; tip of shaft slightly inflated both laterally and dorso-
ventrally; lateral profile gradually sloping anteriorly from widest point of stalk.
Specimen number 65921 (Fig. 43) differs from number 65895 (Fig. 42)
described above. Terminus of shaft of number 65921 has lateral lobes from
which arise lateral cartilaginous processes; median terminal ossification irregu-
lar in shape, smaller, imbedded in terminally bilobate cartilage. In the spatulate
flattened stalk these two specimens are much alike. An immature specimen,
number 65908, is smaller (length of stalk 2.6 mm.) also flattened and spatulate,
has the terminal processes cartilaginous, the lateral processes bent medially,
and proportions as in the adult.
The baculum shows no noteworthy resemblance to that of any other species
of North American Microtus; on the other hand the differences between M.
guatemalensis and some other species are no greater than the differences be-
tween certain species included in the subgenus Microtus. The baculum neither
strengthens nor weakens the case for subgeneric rank for M. (Herpetomys)
guatemalensis.
Specimens examined: Three from Guatemala; 65895 (2 mi. S San Juan
Ixcoy 65908, (3% mi. SW San Juan Ixcoy), 65921 (10 mi. E, 4 mi. S Totoni-
capan).
THe BacuLuM IN MIcROTINE RODENTS 199
Microtus (Arvicola) richardsoni (DeKay)
Figs. 38 and 89
Baculum: Stalk broad, greatest length (3.7 to 4.3 mm.) 1% times greatest
breadth, relatively flattened, greatest depth 14 greatest breadth; single median
ossified process, in smaller of two specimens this ossification incomplete and
of unusual shape (Fig. 39); length of stalk 4 times length of median process;
concavities of basal tuberosities medially confluent, constriction less than %
greatest depth; widest point of shaft less than % length of shaft from posterior-
most point; shaft wider than high except at distal end that is inflated dorsally
and sometimes laterally; both ventral and dorsal concavities of base of stalk
broad and moderately deep; posterior profile in dorsal view evenly rounded or
having marginal notch.
In the absence of ossified lateral processes my two specimens differ from
bacula of Microtus (Arvicola) terrestris figured by Didier (1943:79, 1954:245,
247, 248) and by Ognev (1950:591). The median process relative to the
size of the shaft is smaller, and the shaft relative to its length is wider in M.
richardsoni than in M. terrestris. The stalk of M. (Arvicola) amphibius figured
by Didier is like that of M. richardsoni in its greater breadth and median notch
on posterior border.
The relationship of the New World water rat, M. richardsoni, to the Old
World water rats (genus Arvicola of some European authors) is uncertain.
Miller (1896:66) placed all of them in the subgenus Arvicola. Subsequent
authors, stressing differences in the teeth, have placed M. richardsoni in the
subgenus Aulacomys of Rhoads. Zimmerman (1955) has shown that teeth
in some Arvicola approach the more complex pattern of M. richardsoni. He
argues also that Arvicola is generically distinct from Microtus on the grounds
that the two groups have separate origins, Arvicola having descended from the
genus Mimomys and Microtus from some other group of microtines. This argu-
ment also was advanced by Hinton (1926:47-48). Pending further studies
of the possible polyphyletic origin of other subgenera of the genus Microtus, I
refer both M. richardsoni and M. terrestris to the subgenus Arvicola.
The evidence afforded by the bacula available is not conclusive as to re-
lations of Old World and New World water rats. No general agreement on
the number of species in this Palaearctic group has been reached, and bacula
of only three or four of the numerous Old World subspecies have been figured.
I have examined none.
Specimens examined: Two, from Wyoming; 42454 (31 mi. N Pinedale,
os ft., Sublette Co.), 37903 (23% mi. S, 5 mi. W Lander, 8600 ft., Fremont
o.).
Microtus (Chilotus) oregoni (Bachman)
Fig. 45
Baculum: Stalk broad, greatest length (2.2 mm.) 1% times greatest breadth,
3% times greatest depth; three well-developed ossified processes; median process
% length of stalk, rounded or tapered terminally, proximal end opposed to tip
of stalk and flattened obliquely; lateral processes 24 length of median process,
deeper than wide, curved; tuberosities of stalk well developed, confluent
medially, visible in dorsal view; in end-view dorsal concavity narrow, moder-
200 UNIVERSITY OF KANSAS Pusis., Mus. Nat. Hist.
ately deep, rounded, ventral concavity wide, deep, flattened; base wider
ventrally than dorsally; shaft tapering more or less uniformly, terminally inflated.
In the relative sizes, to each other and to the stalk, of the three digitate ossi-
fications M. oregoni resembles closely the Old World representative of the same
subgenus, M. (Chilotus) socialis, as figured by Argyropulo (1933b:181). In
M. oregoni the greatest width of the baculum is more proximal on the stalk
than in the M. socialis figured by Argyropulo but closely resembles the baculum
of the M. socialis figured by Didier (1954:242). In possessing a shallow
emargination in the base of the stalk and in possessing a median process that
is smaller than the lateral processes, M. socialis, as figured by Didier, differs
from M. oregoni. The baculum figured by Argyropulo (loc. cit.) of Sumeriomys
colchicus schidlovskii [= Microtus (Chilotus) socialis schidlovskii according to
Ognev, 1950:392] differs from other Chilotus that have been studied in having
an unusually elongate median process and a more distal placement of the widest
part of the stalk.
Specimens examined: Three, of the subspecies M. oregoni oregoni, from 5
mi. N Orick, Humboldt Co., California, 3-C-248, collection of W. B. Quay;
from Mary’s Peak, Benton Co., Oregon, 66, collection of F. W. Sturges; and
from Sec. 3, T. 11S, R. 5W, Benton Co., Oregon, 79183.
Microtus (Stenocranius) gregalis (Pallas)
Fig. 34
Baculum: Length of stalk (2.4 mm.) 1% times greatest breadth, 414 times
greatest depth; median ossified process well developed, more than 14 length of
stalk, higher than wide, slightly bowed, closely appressed to terminus of shaft;
basal tuberosities of stalk moderately developed, confluent medially, posterior
profile of medial apex rounded in dorsal view, lateral indentations present, hence
trilobate outline; in proximal end-view base wider ventrally, ventral concavity
broader than dorsal concavity but of equal depth, medial constriction 24 greatest
depth; shaft slender in distal part, inflated terminally, and wider than high at
mid-point of stalk; lateral profile a smooth slope of gradually decreasing curva-
ture from point of greatest width to near distal end.
The baculum of this species figured by Ognev (1950:461) differs in having
lateral ossified processes, and a more rounded base of the stalk. Resemblance
to the New World Stenocranius is discussed below.
Specimen examined: One from “Eastern Europe,” 8059.
Microtus (Stenocranius) miurus Osgood
Figs. 82 and 33
Baculum: Length of stalk (2.8 mm.) 1% times greatest breadth, 3% times
greatest depth; median process ossified, % to *4 length of stalk, laterally com-
pressed, sometimes arched in dorsoventral plane; lateral processes cartilaginous,
slender; basal tuberosities well developed, averaging less enlarged than shown
in Figure 32, but more angular in lateral outline than shown in Figure 33;
tuberosities confluent posteriorly; posterior profile smoothly rounded to trilo-
bate, curvature at point of greatest breadth usually acute; in proximal end-view
base wider dorsally, deep dorsal concavity, shallow ventral concavity, medial
constriction 34 of greatest depth; shaft slender anteriorly, at mid-point of stalk
THE BAcULUM IN MICROTINE RODENTS 201
twice as wide as high, at tip higher than wide, laterally inflated; lateral profile
in most specimens abruptly curved anterior to point of greatest breadth.
The single specimen of the Old World M. (Stenocranius) gregalis examined
resembles the New World M. (Stenocranius) miurus in the angular lateral
profile at the point of greatest breadth of the stalk, slender shaft in comparison
to broad base of stalk, and presence of a single well-developed laterally com-
pressed median process. The base of the stalk in the baculum of M. gregalis
is less well developed and smaller than in the baculum of M. miurus.
Specimens examined: Nine, all of the subspecies Microtus miurus muriei,
from the Brooks Range, Alaska; 51077 (Lake Schrader, 145°09’50”, 69°24’28”,
2900 ft., Romanzof Mts.); 51151, 51152, 51154, 51164, 51166, 51169 (last 6
from Wahoo Lake, 69°08’, 146°58’, 2350 ft.); 51210, 51213 (last 2 from Porcu-
pine Lake, 68°51'57”, 146°29’50”, 3140 ft.).
Microtus (Chionomys) nivalis Martins
Fig. 47
Baculum: Greatest length of stalk (2.7 mm.) 2% times greatest breadth,
4% times greatest depth; three digitate processes, lateral processes mostly
cartilaginous in single adult examined; median process well ossified, approxi-
mately 14 length of stalk, basally notched, not arched, laterally compressed
distally; base of stalk broad and flat, basal tuberosities well developed, separate;
posterior profile in dorsal view rounded, convex except for medial notch sep-
arating tuberosities; dorsal and ventral concavities deep, broad, equal; medial
constriction less than #% greatest depth; in dorsal view shaft tapering gradually
from widest point, terminally rounded; at mid-point of stalk almost twice as
wide as high.
In the elongate, largely cartilaginous lateral processes of the baculum, the
specimen described above resembles M. longicaudus. The size of the median
process in comparison to the size of the stalk is also the same. The lateral
processes have larger ossifications and the base of the stalk is more robust in
M. longicaudus than in M. nivalis.
The well ossified lateral processes and enlarged base of Didier’s (1954:240)
specimen suggest that it is of a more mature individual than the one described
above. These specimens of M. nivalis, as well as the specimens of M. longi-
caudus, exhibit dorso-ventral flattening of the mid-part of the base of the stalk.
The baculum of a specimen from Switzerland is weakly developed, of small
size (shaft 2.0 mm. in length), slender, thin, spatulate, and terminally inflated.
Digital processes were not observed, perhaps owing to excessive maceration in
preparation. The general appearance of the baculum is that of an immature
individual, although the animal was not small (165 mm. total length in
preservative ).
Specimens examined: Two Microtus nivalis nivalis; Zermatt, Valais, Switzer-
land, 67105; Wetterstein, Germany, 65127.
Microtus (Chionomys) longicaudus (Merriam)
Fig. 48
Baculum: Base of stalk well developed, greatest length (3 mm.) 1% times
greatest breadth, 3%4 times greatest depth; three ossified processes; base of
median process rounded; median process slightly curved in dorsoventral plane,
in length almost 14 greatest length of stalk; ossifications in lateral processes
202 UNIvERSITY OF KANSAS Pusts., Mus. Nat. Hist.
variable in size, frequently widely separated from shaft by cartilage, rarely as
large as median ossification; basal tuberosities usually well-developed, medially
confluent; profile of base in dorsal view trilobate or irregularly convex through-
out; constriction % greatest depth; shaft relatively straight or slightly bowed
ventrally or dorsally, shaft at mid-point of stalk wider than high; tip of shaft
laterally inflated; widest point of stalk approximately % length of stalk from
proximal end; lateral profile in dorsal view tapers gradually onto shaft anteriorly
from point of greatest width of stalk; shaft variable, from slender terminally
and nearly parallel sided (Fig. 48), to broad distally and tapered.
In many of the features that distinguish M. longicaudus (and the closely
related insular species M. coronarius) from other North American Microtus,
longicaudus resembles the Old World species of the subgenus Chionomys (that
is to say, M. nivalis, M. gud, and M. roberti). These features are medium
size, long tail, grayish color, montane habitat, relatively short molar tooth-row,
moderate sized and unconstricted incisive foramen, relatively decurved upper
incisors, elongate nasals, relatively broad interorbital region without well-
developed median ridge, and similar chromosomes (Matthey, 1955:178).
For these reasons I am here referring Microtus longicaudus to the subgenus
Chionomys; previously it has not been referred to that subgenus.
Specimens examined: Six, of three subspecies; Microtus longicaudus lit-
toralis, Sullivan Island, Alaska, 42972, 42969; M. l. mordax, % mi. N, 2 mi. W
Allenspark, 8400 ft., Boulder Co., Colorado, 50335, 76829; M. l. sierrae, Crane
Flat, Mariposa Co., California, 50252, 50253.
Microtus arvalis (Pallas)
Fig. 22
Baculum: In the single specimen examined, stalk small, greatest length
(2.3 mm.) 21% times greatest width, almost 6 times greatest depth, flattened
proximally; three well-developed digitate processes, the median one ossified,
the lateral processes cartilaginous; median ossification laterally compressed and
decurved at tip, bilobate at base; basal tuberosities of stalk weakly developed,
medially confluent; posterior profile in dorsal view evenly rounded; ventral
concavity deeper and narrower than dorsal concavity, but both comparatively
shallow; medial constriction 4 greatest depth; shaft straight, at mid-point
twice as wide as deep; lateral profile tapering from greatest width gradually
to parallel sides of distal third of stalk.
From the baculum of Microtus arvalis figured by Ognev (1950:173), and
from the baculum figured by Didier (1954:238) my specimen differs in the
absence of lateral ossifications in the digitate processes, smaller and slenderer
median ossification, and weaker base. These differences in part may be owing
to a difference in age, my specimen being the less mature.
Specimen examined: One from Vidy, Switzerland, 67101.
Microtus orcadensis Millais
Fig. 24
Baculum: In the one specimen examined, stalk broad, greatest length
(2.6 mm.) 1% times greatest breadth, 3% times greatest depth; three digitate
processes ossified; median process relatively broad, in length more than %
length of stalk, triangular in dorsal view, with small spurs posterolaterally,
THE BACULUM IN MICROTINE RODENTS 203
middorsal ridge posteriorly; lateral ossifications slightly curved, slenderer, less
than % depth and less than % transverse thickness of median process; basal
tuberosities well-developed, confluent medially; in end-view base wider dor-
sally than ventrally, dorsal concavity broader and more abruptly curved at
mid-point than ventral concavity; constriction % greatest depth; posterior profile
in dorsal view notched, setting off a posterior shelf; stalk including shaft wider
than deep throughout, at mid-point width twice depth; lateral profile abruptly
curved anterior to point of greatest width, sides of shaft tapering gradually
anteriorly to rounded uninflated tip.
The baculum of this insular species, placed in the “arvalis” group by Eller-
man (1941:595), resembles the baculum of both Microtus agrestis and Micro-
tus guentheri more than it resembles the baculum of Microtus arvalis. Simi-
larities in the chromosomes of M. arvalis and M. orcadensis were noted by
Matthey (1953:254, 279), who was of the opinion that M. orcadensis is an
insular derivative of the arvalis-group.
Specimen examined: One from the Orkney Islands, 67106.
Microtus guentheri Danford and Alston
Fig. 23
Baculum: In the one specimen examined, stalk broad, greatest length (2.9
mm.) 1% times greatest breadth, 3% times greatest depth; three digitate
processes ossified; median process slightly less than % length of stalk, broad,
dorsally curved; curved lateral ossifications shorter and more slender than
median ossification; basal tuberosities well developed, angular, confluent
across posterior border of projecting shelf; in end-view tuberosities projecting
ventrolaterally from central shelf; dorsal surface at medial constriction flat,
ventral surface broadly and deeply concave; posterior profile in dorsal view
trilobate, central lobe formed by posteriorly flattened shelf, surface of at-
tachment visible only on lateral lobes; at mid-point stalk almost twice as
wide as deep, depth of shaft greater than width proximal to inflated terminus.
Specimen examined: One from Palestine, 67104.
Microtus fortis Biichner
Fig. 25
Baculum: Stalk large, greatest length (3.8 mm.) 14 times greatest breadth,
4% times greatest depth; three digitate processes ossified; median ossification
almost % length of stalk; lateral ossifications slender, smaller than median
ossification; posterior profile of stalk in dorsal view trilobate, basal tuberosities
well developed, confluent medially; in end-view dorsal concavity broader and
deeper than ventral concavity; medial constriction pronuonced (less than %
greatest depth); lateral profile at widest point of stalk convex, becoming
abruptly concave as the flange of the basal tuberosities grades into the shaft,
then gradually converging to narrowest point % of length of stalk from the
terminus; stalk wider than deep in proximal %, circular in cross section in
terminal \4, slight terminal inflation.
A specimen figured by Ognev (1950:297) has the same general propor-
tions, slender lateral processes, and proximal placement of the point of greatest
breadth.
Specimens examined: Two from Chipo-ri, Korea, 604438, 63841.
204 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
Microtus montanus (Peale)
Figs. 19, 20 and 21
Baculum: Stalk broad, greatest length (varying with subspecies from 2.3
to 3.1 mm.) 1% to 1% times greatest breadth, 314 to 414 times greatest depth;
three ossified processes, median one largest, more than twice as wide and as
deep as shorter, slenderer, lateral processes; median process laterally com-
pressed distally except in one specimen in which moderately inflated distally,
proximally enlarged in some specimens (Fig. 21) and % to % length of stalk;
base broad, posterior profile in dorsal view evenly convex throughout, at widest
point of stalk abruptly incurved; basal tuberosities moderately to strongly de-
veloped, medially confluent; in end-view base wider ventrally than dorsally,
dorsal concavity slightly to much deeper than the nearly flattened ventral
concavity; medial constriction % to 4 of greatest depth; shaft relatively slender,
at mid-point of stalk slightly wider than high and % as wide as base of stalk,
terminally rounded or slightly inflated; lateral profile in dorsal view a gradual
curve from point of greatest width anteriorly onto shaft.
The different subspecies figured show the essential characteristics of the
species, differing primarily in size.
Specimens examined: Fourteen, of three subspecies; Microtus montanus
amosus, % mi. E Soldier Summit, Wasatch Co., Utah, 62241; M. montanus
fusus, La Manga Pass, Conejos Co., Colorado, 42164; 5 mi. N, 26 mi. W
Saguache, 9500 ft., Saguache Co., Colorado, 42307, 42315; 5 mi. N, 27 mi. W
Saguache, 9350 ft., Saguache Co., Colorado, 42308; 5 mi. N, 28 mi. W
Saguache, 9325 ft., Saguache Co., Colorado, 42309; 5 mi. S, 24 mi. W Antonito,
9600 ft., Conejos Co., Colorado, 42327, 42330; Prater Canyon, Mesa Verde Na-
tional Park, Montezuma Co., Colorado, 69456, 69457, 69463; Microtus mon-
tanus nanus, 2 mi. N, 2 mi. W Pocatello, Bannock Co., Idaho, 57470, 57472;
% mi. N, 2 mi. W Allenspark, 8400 ft., Boulder Co., Colorado, 50330.
Microtus townsendii (Bachman)
Fig. 41
Baculum: Stalk broad, greatest length (3.0 mm.) 1% times greatest breadth,
4% times greatest depth; three ossified processes, median one largest, deeper
and more than twice as wide as curved, shorter, compressed lateral processes
and more than % as long as stalk; base broad, in dorsal view posterior profile
trilobate, basal tuberosities visible; basal tuberosities well developed, medially
confluent; in end-view base wider ventrally than dorsally, dorsal concavity
deeper than ventral concavity; medial constriction % of greatest depth; shaft
broad, at mid-point more than twice as wide as high and 14 as wide as base of
stalk, terminally rounded.
Specimens examined: Three, all M. t. townsendii; Fort Lewis, Pierce Co.,
Washington, 57998, subadult; Sec. 33, T. 11S, R. 5W, Benton Co., Oregon,
79186; Sec. 5, T. 12S, R. 4W, Benton Co., Oregon, 79188.
Microtus oeconomus (Pallas)
Fig. 44
Baculum: Stalk broad and flattened, greatest length (3.5 mm.) 1% to 2
times greatest width, 4 to 5% times greatest depth; three ossified processes,
median one largest, lateral processes slender, relatively small; length of median
process % length of stalk; median process decurved, dorsoventrally flattened in
THE BACULUM IN MICROTINE RODENTS 205
some specimens, widened at base; attachment of processes to shaft displaced
ventrally; base of stalk widened, posterior profile in dorsal view usually trilobate,
in a few cases rounded, median lobe forming posterior shelf, lateral lobes
dorsally raised and forming margins of lateral tuberosities; in end-view thick-
ness frequently more or less uniform throughout central part, broad depression
dorsally, ventral concavity narrower and shallower (as figured); base, and
occasionally shaft, flattened, width at mid-point of stalk 2 to 3 times depth,
narrowest point posterior to terminal inflation of shaft in terminal 14 of shaft.
The baculum of M. oeconomus (Old World) figured by Ognev (1950:257)
resembles but exceeds that of M. oeconomus (New World) in the relatively
large median process and slender lateral processes, but differs noticeably in the
presence of a deep median notch in the base of the stalk. A specimen from
Hungary is intermediate between Ognev’s specimen and those from the New
World in both size of median process and size of lateral processes, and has an
unnotched base resembling that in Figure 44.
Specimens examined: Ten, of three subspecies; M. oeconomus gilmorei,
Umiat, Alaska, 51354, 51361, 51399, 51408; Lake Schrader, Brooks Range,
Alaska, 51422; M. o. macfarlani, 5 mi. NNE Gulkana, Alaska, 43039, 43041;
20 mi. NE Anchorage, Alaska, 43044; Kelsall Lake, British Columbia, 43048;
M. o. mehelyi, Kisbalatan, Hungary, 75159.
Microtus mexicanus (Saussure)
Figs. 85 and 36
Baculum: Stalk attenuate, greatest breadth relatively near proximal end;
greatest length (3.1 to 3.4 mm.) more or less twice greatest breadth, 4 to 5
times greatest depth; usually a single process ossified; lateral processes relatively
small, cartilaginous or (in three specimens, 63094, 69453, 68019) with small
ossifications; median process relatively small, sometimes appressed to tip of
shaft, in length less than % length of stalk; posterior profile in dorsal view
rounded, flattened posteriorly, or in some specimens trilobate with angular
edges; in end-view relative depths of dorsal and ventral concavities variable,
dorsal usually deeper than ventral; distal end of stalk frequently bowed
dorsally; shaft slender distally, sometimes slightly inflated terminally, or (in
one specimen, 63085) near tip small lateral projections that are perhaps fused
lateral ossifications; lateral profile in dorsal view a gradual slope anteriorly
from point of greatest width to slender tip.
Specimens examined: Thirteen, of four subspecies; Microtus mexicanus
mexicanus, Las Vigas, Veracruz, 30692; Nevada de Toluca, México, 63101;
Valle de Bravo, México, 63094; Microtus mexicanus mogollonensis, Mt. Taylor,
Valencia Co., New Mexico, 63298, 76830; Park Well, Mesa Verde National
Park, Montezuma Co., Colorado, 69448, 69453; Upper Nutria, McKinley Co.,
New Mexico, 69997, 70000; Microtus mexicanus phaeus, Sierra Patamba, 9000
ft., Michoacan, 63085; Microtus mexicanus subsimus, 2 mi. E Mesa de Tablas,
Coahuila, 58916; 13 mi. E San Antonio de las Alazanas, Coahuila, 68019, 68021.
Microtus californicus (Peale)
Fig. 37
Baculum: Stalk elongate, greatest length (3.0 mm.) 2%4 times greatest
breadth, 4% times greatest depth; median process ossified, % length of stalk,
basally broadened, flattened and shallowly grooved ventrally to fit tip of shaft,
to which the process is closely appressed; lateral processes cartilaginous; ends
206 UNIVERSITY OF KANsAs Pusts., Mus. Nat. Hist.
of stalk bowed upwardly; posterior profile of base of stalk rounded or slightly
trilobate if posterolateral concavities form in tuberosities; moderate develop-
ment of tuberosities, in end-view dorsal concavity slightly deeper and narrower
than ventral concavity, both comparatively shallow, median constriction %
greatest depth; shaft curved, more or less terete at mid-point of stalk, terminally
inflated dorsally; lateral profile in dorsal view gradually curved from point of
greatest width anteriorly onto shaft.
Specimens examined: Two, of two subspecies; Microtus californicus cali-
fornicus, 1 mi. NE Berkeley, in Contra Costa Co., California, 76828; Microtus
bel ee mohavensis, % mi. SE Victorville, San Bernardino Co., California,
Microtus pennsylvanicus (Ord)
Figs. 14, 15, 16 and 17
Baculum: Stalk heavy, broad, greatest length (2.2 to 3.0 mm.) 1% to
1% times greatest breadth, up to 3% times greatest depth; three ossified
processes, median one largest, usually not twice so deep as lateral ossifications;
median process usually distinctly widened basally, in length approximately %
length of stalk; base broad, frequently angular laterally and basally, sometimes
bilobate; basal tuberosities well developed, medially confluent; in end-view
more or less uniformly biconvex or ventral surface more flattened than dorsal
surface, medial constriction % to 74 greatest depth; shaft relatively heavy, at
mid-point stalk almost twice as wide as deep and 14 as wide as base of stalk;
shaft terminally rounded and sometimes slightly inflated; lateral profile in
dorsal view abruptly or gradually curved anterior to point of greatest width
and then gradually curved anteriorly.
Specimens examined averaged slightly smaller and were more variable than
those described by Hamilton (1946:382). The greater variation may be in part
geographic, as five subspecies are represented. Lateral processes are the last
to ossify. One specimen (75082) with well-ossified median process lacks any
lateral ossification. Four bacula of M. pennsylvanicus (locality not specified )
studied by Dearden (1958:547) agree in general with the description above.
One specimen shows a break, perhaps resulting from injury, in the shaft
(Fig. 14). One specimen has a posteromedian spine on the median digital
ossification (Fig. 16). Comparison with M. agrestis is included with the
description of M. agrestis.
Specimens examined: Thirteen, of six subspecies; Microtus pennsylvanicus
alcorni, 20 mi. NE Anchorage, Alaska, 43048; Microtus pennsylvanicus finitus,
Laird, Yuma Co., Colorado, 68544; Microtus pennsylvanicus modestus, 5 mi.
N, 26 mi. W Saguache, 9500 ft., Saguache Co., Colorado, 42306; 3 mi. N, 16
mi. W Saguache, 8500 ft., Saguache Co., Colorado, 42416, 42417, 42418; 1 mi.
S, 2 mi. E Eagle Nest, 8100 ft., Colfax Co., New Mexico, 42430, 42439;
Microtus pennsylvanicus pennsylvanicus, 2 mi. S, 3 mi. E Ft. Thompson, 1370
ft., Buffalo Co., South Dakota, 42379; Vermillion, Clay Co., South Dakota,
87070; Microtus pennsylvanicus pullatus, 12 mi. S, 5 mi. E Butte, Silver Bow
Co., Montana, 57501, 57503; Microtus pennsylvanicus uligocola, Muir Springs,
2 mi. N, 24 mi. W Ft. Morgan, Morgan Co., Colorado, 75082.
Microtus agrestis (Linnaeus)
Fig. 18
Baculum: Greatest length of stalk (2.9 mm.) twice greatest breadth, 4%
times greatest depth; stalk well developed, shaft not flattened dorsoventrally;
large median ossified process, minute lateral ossifications in single specimen
Tue BacuLuM IN MICROTINE RODENTS 207
examined; length of stalk 2% times length of median ossification which is higher
than wide, slightly decurved, sagittate in dorsal view, with three-cornered base;
basal tuberosities of stalk moderately well developed, medially joined; posterior
profile in dorsal view evenly rounded; ventral concavity broader than, but of
comparable depth to, dorsal concavity in end-view, base of stalk wider ventrally,
constriction % greatest depth; at mid-point of stalk shaft is but slightly wider
than high; pronounced terminal inflation of shaft; lateral profile in dorsal view
sloping abruptly from widest point of stalk anteriorly onto stalk which then
tapers more gradually to terminal inflation.
From the baculum of its New World counterpart, namely Microtus pennsyl-
vanicus, my specimen of Microtus agrestis and the specimen figured by Didier
(1954:239) differ in their minute lateral processes, relatively larger median
processes, and more elongate, less dorsoventrally flattened shafts.
The specimen of M. agrestis figured by Ognev (1950:320), in dorsal view
has lateral concavities producing a somewhat trilobate outline in the base of
the stalk, and the lateral processes are well developed; the median process is
larger and bulbous, wider distally than proximally. Without larger numbers
of bacula of M. agrestis I am unable to reconcile these differences. The
differences between M. agrestis and M. pennsylvanicus seem comparable to the
differences between some other species of Microtus.
Specimen examined: One, from Gryon, Switzerland, 67102.
Microtus (Pedomys) ochrogaster (Wagner)
Fig. 31
Baculum: Stalk broad, greatest length (3.2-4.0 mm.) 1% to 2 times greatest
breadth, 2% to 4 times greatest depth; median process ossified, relatively small,
less than 340 length of stalk; lateral processes arising from subterminal part of
stalk, cartilaginous or with small ossifications; posterior profile in dorsal view
broadly rounded or slightly angular, widest point of stalk 4% to % the length
of stalk from base; basal tuberosities well developed and medially confluent,
in end-view dorsally convex, or at least less deeply concave than ventrally;
shaft straight, base bent ventrally or more commonly dorsally; at mid-point of
stalk wider than high, often twice as wide as high; viewed from above, lateral
profile from point of greatest breadth to middle of shaft a gradual sigmoid
curve; slight terminal inflation of shaft.
Specimens examined: Forty-one, of three subspecies; Microtus ochrogaster
haydeni, Muir Springs, 2 mi. N, 2% mi. W Ft. Morgan, Morgan Co., Colorado,
74995, 74998, 74999, 75002; 1 mi. W Laird, Yuma Co., Colorado, 57304, 76833;
2 mi. N, 2 mi. W Haigler, Dundy Co., Nebraska, 75016; 2 mi. S Franklin,
Franklin Co., Nebraska, 75043, 75044; Atwood, Rawlins Co., Kansas, 75020,
75023, 75025, 75027, 75028; 1 mi. N, 2 mi. E Oberlin, Decatur Co., Kansas,
75030, 75032, 75034, 75035, 75036; 1% mi. N, 4 mi. E Norton, Norton Co.,
Kansas, 68327; 1 mi. SW Norton, Norton Co., Kansas, 75037; 2 mi. S, 1 mi.
W Norton, Norton Co., Kansas, 75038; M. ochrogaster ochrogaster, Rydal,
Republic Co., Kansas, 75047-75053, 75060, 75062, 75063, 75066, 75070, 75071,
75078; 1 mi. N, 1 mi. W Holton, Jackson Co., Kansas, 75077; 2 mi. W Court
House, Lawrence, Douglas Co., Kansas, 76832; Univ. Kansas Natural History
Reservation, Douglas Co., Kansas, 68536; M. ochrogaster taylori, Meade
County State Park, Kansas, 68539, 68542.
208 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
Microtus (Pitymys) pinetorum (LeConte)
Figs. 27 and 28
Baculum: Stalk broad, greatest length (2.5 to 2.7 mm.) 1% times greatest
breadth, 4 times greatest depth; median process ossified, size small, % length
of stalk, higher than wide, having small anterodorsal prominence in both
specimens examined; lateral processes cartilaginous, relatively small, displaced
posteriorly, attenuate; posterior margin in dorsal view broadly rounded, or
having blunt median apex, convex throughout; basal tuberosities moderately
well developed, medially confluent, barely visible in dorsal view when mature;
in end-view median constriction 4 greatest depth, ventral concavity deeper
than dorsal concavity, both comparatively shallow; stalk at mid-point 1% times
as wide as deep; shaft relatively slender, bowed dorsally at tip, relatively
straight otherwise; lateral profile in dorsal view a gradual concave slope from
point of greatest width anteriorly to distal part of shaft.
Specimens examined: Two, from Douglas Co., Kansas, 76834 (2 mi. N
Baldwin), 68545 (1 mi. NE Pleasant Grove).
Microtus (Pitymys) parvulus (Howell)
Fig. 40
Baculum: Stalk broad, greatest length (2.4 mm. in specimen examined) 1%
times greatest breadth, 4 times greatest depth; median process ossified, size
small, less than % length of stalk, wider than high, terminally flattened; lateral
processes cartilaginous, relatively small, attenuate; posterior margin in dorsal
view flattened, irregularly curved with concavities medially and laterally; basal
tuberosities well developed, medially confluent; visible in dorsal view; in end-
view median constriction %4 greatest depth, ventral concavity well-formed, no
dorsal concavity; stalk at mid-point twice as wide as deep; shaft relatively
slender, bowed dorsally toward tip; in dorsal view lateral profile a gradual
concave slope from point of greatest width anteriorly to distal part of shaft;
tip of shaft enlarged.
The baculum of M. parvulus resembles that of M. pinetorum more than it
resembles the baculum of any other microtine studied, differing primarily in
smaller size.
Specimen examined: One, from 1 mi. W Micanopy, Alachua Co., Florida,
Univ. Florida No. 1508.
Microtus (Pitymys) quasiater (Coues)
Figs. 29 and 30
Baculum: Stalk broad, greatest length (2.6-3.2 mm.) 14% to 1% times
greatest breadth, 314 to 3%4 times greatest depth; median process ossified, with
ventral depression, process 4% to 144 length of stalk, appressed to tip of shaft,
wider than high proximally, relatively broad terminally; lateral processes carti-
laginous, small, attenuate; posterior profile of stalk in dorsal view broadly
rounded, bilobate, or trilobate, median lobe formed by posterior projection of
dorsal shelf between enlarged lateral tuberosities that form outer lobes, postero-
lateral faces of these tuberosities visible in dorsal view of stalk; in end-view
dorsal surface slightly concave, ventral concavity broad and deep, median con-
Tue BacuLUM IN MICROTINE RODENTS 209
striction % greatest depth; shaft flattened except tip that is more terete, and
bowed dorsally; at mid-point, stalk twice as wide as high; shaft relatively slender
terminally, narrower than median ossification.
The baculum of M. quasiater is the largest and has the best developed
base and median process of the three American species of the subgenus Pitymys.
The three species closely resemble each other in basic form,
Specimens examined: Five, all from Veracruz; Teocelo, 4500 ft., 30709,
Se 4 km. N Tlapacoyan, 1700 ft., 24466; 5 km. N Jalapa, 4500 ft., 19869,
Microtus (Pitymys) fatioi (Mottaz)
Fig. 26
The baculum of a single specimen (KU 67103) of M. fatioi from Zermatt,
Valais, Switzerland, was examined. The baculum is immature, as evidenced
by its small size, slender stalk and absence of ossified processes, therefore no
characterization is included.
The baculum of another Old World species of the subgenus Pitymys, M.
pyrenaicus from France, figured and described by Didier (1954:242-243),
differs from all New World Pitymys examined in processing ossified lateral
processes.
The status of Pitymys, as a genus or as a subgenus, is uncertain. Hall and
Cockrum (1953:448) considered the North American Pitymys and Pedomys as
subgenera of Microtus. They did not state specifically the basis for this point
of view, but mention the fact that these two subgenera (Pitymys and Pedomys)
closely resemble each other cranially. These authors did not study nor com-
ment upon the status of the Old World Pitymys. It may be asked whether the
Old World and New World Pitymys have developed as fossorial Microtus
independently, or from an ancestor common to both groups and not common
to any other Microtus. Matthey (1955:202) found 62 chromosomes (2N) in
both the New World Pitymys pinetorum and the Old World Pitymys duodeci-
mcostatus. This suggests, but does not prove, common ancestry.
Neofiber alleni True
Fig. 49
Baculum: Stalk massive, greatest length (4.7 mm.) 1% times greatest
breadth, 4 times greatest depth; ossification in digitate processes variable; in
one (KU 27123) of two specimens examined lateral processes ossified and
median process unossified, as in two specimens examined by Hamilton (1946:
879) from “southern Florida”; in my other specimen (KU 27268) that is
possibly more mature, median process ossified although less deeply stained
than lateral ossifications or stalk; posterior profile in probable dorsal view
roughly rounded; in end-view probable dorsal concavity deep, ventral con-
cavity broad but shallow, and with center convex; median constriction %
greatest depth; shaft heavy, least depth 74 greatest depth of base; stalk, at
mid-point, slightly wider than deep and more than 14 width of base; lateral
profile in dorsal view sharply incurved distal to point of greatest breadth, shaft
therefore relatively distinct from basal part of stalk; slight subterminal con-
striction; tip less reduced in the two specimens examined than in two figured
by Hamilton. In preparation, the tissues that make it possible to distinguish
210 UNIVERSITY OF KANsAs Pusts., Mus. Nat. Hist.
with certainty the dorsal and ventral surfaces of the baculum were removed
in both specimens.
Specimens examined: Two, of the subspecies Neofiber alleni alleni, 2 mi. S
Gainesville, Alachua Co., Florida, 27268; 1 mi. E Courtenay, Merritt Island,
Brevard Co., Florida, 27123.
Lagurus curtatus (Cope)
Fig. 46
Baculum: Stalk slender, greatest length (2.5 mm.) 2 to 274 times greatest
breadth, 4 to 5 times greatest depth; three ossified processes; median one more
than 14 length of stalk, curved dorsally toward tip, proximally flattened and
having acute lateral angles in dorsal view, wider than deep except in distal
half; lateral processes smaller than median one, slenderer, shorter, of approxi-
mately same depth, also curved dorsally; base of stalk well developed, basal
tuberosities medially confluent, in part visible in dorsal view, in end-view
wider ventrally than dorsally, dorsal and ventral concavities of equal depth
and both wide, medial constriction % greatest depth; posterior profile in dorsal
view broadly bilobate; lateral profile with abrupt transition from basal tuberosi-
ties to gradually converging, slightly curved sides of shaft; shaft terminally
inflated.
Dearden (1958:543) described and figured the bacula of six subspecies of
Lagurus curtatus and two Asiatic species, Lagurus lagurus and Lagurus luteus.
He examined at least 34 specimens of L. curtatus and found geographic variation
in size, breadth of shaft distally, and proportions of digital ossifications to each
other and to the stalk. The description that I have given above pertains to
L. c. levidensis.
The baculum of the Asiatic Lagurus (Lagurus) lagurus figured by Ognev
(1950:554) agrees with that of Lagurus (Lemmiscus) curtatus, described
here, in the relatively elongate shaft and slender stalk, the proportions of the
processes, and the well-formed and moderately enlarged base of the stalk.
The bacula of three Lagurus lagurus examined by Dearden (1958:545) were
of older individuals than the specimen that Ognev figures and differ from it
and from bacula of Lagurus curtatus (all subspecies) in the unusual, almost
heart shaped, median process, and in larger size. Lagurus luteus examined by
Dearden (1958:545) differs from both Lagurus lagurus and Lagurus curtatus
in lacking lateral digital ossifications and in having shorter median digital
ossifications and wider base of stalk.
Specimens examined: Seven Lagurus curtatus levidensis from Wyoming;
9 mi. S Robertson, Uinta Co., 26045, 26053; 8 mi. S, 2% mi. E Robertson,
Uinta Co., 26049; Farson, Sweetwater Co., 37906; 16 mi. S, 11 mi. W Walt-
man, Natrona Co., 42457; 32 mi. S, 22 mi. E Rock Springs, 42465, 42466.
The following key to the bacula in some adult North American Microtinae
is intended to help point out some of the most important differences. It should
be noted that not all species can be keyed out on the basis of the baculum.
The most difficult group in this respect includes the species of Microtus that
have small or no ossified lateral processes, especially species of the subgenera
Pedomys and Pitymys, and the two species Microtus californicus and Microtus
mexicanus of the subgenus Microtus. Another complicating factor is the vari-
ability of bacula evident in some species even in the small samples available.
Tue BacuLuM IN MICROTINE RODENTS
211
It is to be expected that additional specimens will show variations not yet
observed.
10’.
elt.
. Size small, less than 3.4 mm. in total length. .... Microtus oregoni,
12"
. Greatest width of stalk at a point about 4 of length of stalk from
>
137.
14’,
15.
. Lateral ossifications equal in length to median ossification
Kry TO THE BACULA OF SOME NorTH AMERICAN MICROTINES
Length of lateral digital ossifications more than 44 breadth of
Stal ke eee A DY, Deis | piaree aie ie eal ope be Cums, sees 2
. Length of lateral digital ossifications less than 14 breadth of stalk
QENAISEN Ge tee sees ty Rah lta Bek Cie acho Desens Se 15
Size small (total length of baculum less than 5.5 mm.)........ 8
. Size large (total length of baculum more than 5.5 mm.)...... 14
. Width at mid-point of stalk more than 4% greatest breadth of
Ste cae NRE OE EA yatta ookaok cimaole 4
. Width at mid-point of stalk less than 4 greatest breadth of stalk, 8
. Stalk, viewed from proximal end hour-glass shaped, and width of
stalk less than % length of stalk...... Phenacomys intermedius,
. Stalk not both hour-glass shaped when viewed from proximal
end, and with width less than % length of stalk
Shaft thin basally, thickness less than 14 of greatest breadth.... 6
. Shaft thick basally, thickness 4 or more of greatest breadth.... 7
Stalk more or less straight, base not deflected. Microtus oeconomus,
. Stalk spatulate, and base deflected from axis of shaft............
Ne ete oebs Pr EP yo. oka ed 5, VRS Microtus guatemalensis,
Base enlarged, depth nearly % of breadth. . .Lemmus trimucronatus,
. Base moderately enlarged, depth near ¥ of breadth............
Be ee: Microtus pennsylvanicus, p. 206, or Microtus townsendii,
. Base hour-glass shaped as viewed from proximal end..........
ee at i Pee ci Oe AS 9S Phenacomys intermedius,
PRINGES SOF ete Ai ae ie A Ea een EE Eee na Stee 9
. Lateral processes separated from tip of shaft by more than the
thiekness-of the lateral process... -..s- 2. w.020--cre-e eee: 10
. Lateral processes separated from tip of shaft by less than the
thicknesssof ‘tue lateral “process #1. 5.) he ee ee eee ee 11
Lateral processes more than % the width of median process......
VS Pe ae, ce EN, SOONER in SCPE SOM Microtus longicaudus,
Lateral processes slender, less than % the width of median
NITOCESS ees ngs ne iE er pen | AS ed ass oe Microtus montanus,
BETA te CE PEE, DRE SOR I EY Clethrionomys,
Lateral ossifications shorter than median ossification.......... 12
Size medium, more than 3.4 mm. in total length............ 13
ase wut tcce. Microtus chrotorrhinus (Hamilton, 1946:382).
Greatest width of stalk at a point less than 4% of length of stalk
froma basew., Sseee eee ise AS eres ED. LS Lagurus curtatus,
. Size of baculum larger, base more than 3 mm. wide, processes
all well. devyelopeds..u 3) Seeae 26s ee ger Ondatra zibethicus,
Size of baculum smaller, base less than 3 mm. wide, processes
poorly developed in some animals............. Neofiber alleni, p.
At least one digital ossification present.................... 16
p.
p.
. 197
. 204
. 198
193
204
. 197
. 201
. 204
. 194
. 199
210
198
209
212 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
15’. Digital ossifications not Deesent Peers Dicrostonyx groenlandicus, p. 193
16. Breadth of stalk at least % length Ofistalicaeisce we eae 17
16’. Breadth of stalk less than % length of stalk.................. 19
17. Length of stalk greater than 3.6 mm. and less than 1% times its
greatest) breadth =. arta. el osnie phe Oates Microtus richardsoni, p. 199
17’. Length of stalk usually less than 3.6 mm., or if more than 3.6
mm. (up to 4.0 mm.) then length 1% times or more its greatest
Parca thin cio ce elec thats ao Ree eee OM cis Se ee ee eee 18
18. Median process attenuate distally in dorsal view, and relatively
long (more than twice its own breadth), ¥% to %4 the length of
stalk; breadth of stalk usually 34 or more length of stalk.........
BEES ee SS ne, ee ey NT A ER EN CR AERO Microtus miurus, p. 200
18’. Median process relatively blunt distally in dorsal view, relatively
short (usually less than % length of stalk), breadth of stalk usu-
allydless than: 3¢slengthvobistalics’ \lae heise ac ne oe eee ae
Se Pitymys, p. 208, Pedomys, p. 207, or Microtus mexicanus, p. 205
19. Distal processes small and firmly ankylosed to distal end of shaft
ete Vek eae vetoes Cpa ctand mecechal eee em Phenacomys longicaudus, p. 197
19’. Distal processes if present not firmly ankylosed to distal end of
SER EER Rs ee eds Ne oR geod TE yer Ce eS 20
20. Dorsal concavity of base as viewed from proximal end usually
deeper than ventral concavity.............. Microtus mexicanus, p. 205
20’. Dorsal and ventral concavities of base equal in depth or ventral
One: the vdeeper st) te nee ee io 2 eae ee 21
21. Total length of baculum more than 3.6 mm... Microtus californicus, p. 205
21’. Total length of baculum less than 3.6 mm.....Synaptomys cooperi, p. 194
DISCUSSION
Owing to shortness of lower incisors and present geographic dis-
tribution of the species, Hinton (1926:85) considered the Tribe
Lemmi (lemmings) to be more primitive than the Tribe Microti
(voles). The surviving lemmings are specialized in many features
and therefore are considered as advanced end-products of an evo-
lutionary radiation of a primitive microtine stock, of which all earlier
stages are extinct.
Hinton regarded Dicrostonyx as the most primitive of the genera
of lemmings on account of its more complex molar teeth (com-
plexity was considered to be primitive), and on account of the
presence of three primitive longitudinal rows of tubercles in unworn
molars. The other three genera were arranged in order of increasing
specialization as follows: Synaptomys, Myopus, Lemmus.
If the baculum tended to retain its primitive character while
specializations in the external anatomy developed, and if the above
arrangement is correct the most primitive bacula would be found
in Dicrostonyx and in Synaptomys. The baculum in these two
genera in comparison to that in Myopus (as figured by Ognev, 1948:
Tue BACuLUM IN MICROTINE RODENTS 213
512) and Lemmus has a slenderer stalk and smaller digital ossifica-
tions or none at all. The baculum in the genera of lemmings in-
creases in robustness and the development of processes from Dicro-
stonyx, to Synaptomys, to Myopus, to Lemmus—the same order out-
lined above for total of specialization. The two extremes in this
series are near the extremes of variation in bacula to be found in all
microtines. The baculum in lemmings as a group cannot then be
considered more primitive than in voles as a group, although the
voles are usually considered to be more advanced. The situation
in the voles, as we shall see, casts a different light on the matter.
The voles, Tribe Microti, were considered by Hinton (1926:40)
to be more advanced than the lemmings because the incisors of
the voles are longer and the root of their last lower molar is lingual
to the root of the incisor. Hinton thought also that the murine
ancestors of microtines had shorter incisors and that the backward
extension of the incisors in the voles is a more ancient feature than
the hypsodonty of the molars. A trend in the molar teeth has been
toward greater hypsodonty. The voles in which the molars are
least hypsodont are thus considered primitive. These include the
living genera Clethrionomys, Phenacomys, Ondatra, Dolomys, Ello-
bius, and Prometheomys. Therefore, the baculum, in these as-
sumedly primitive genera, would be expected to resemble the
baculum in the lemmings or at least the most primitive lemmings.
This is not the case.
The bacula that I have examined of Clethrionomys and Phenaco-
mys have well-developed digital ossifications. In this they resemble
the baculum of the genus Lemmus, the most advanced genus of
lemmings according to Hinton. The baculum of Dolomys has not
been studied. The baculum in Ondatra, and in Prometheomys as
illustrated by Ognev (1948:552), also possesses well-developed
processes. The baculum of Ellobius is small and lacks processes
(as figured by Ognev, 1950:662). No ossification was found in a
single specimen of Ellobius examined by me although the entire
glans penis was removed and cleared without dissection. So far as
known then, with the exception of Ellobius and Phenacomys longi-
caudus (Dearden, 1958:547), the primitive microtines having rooted
molars possess bacula having three well-developed ossified proc-
esses.
Voles of the genus Microtus vary in the structure of the baculum
almost as much as do the lemmings. Within the single subgenus
Microtus some individuals of Microtus mexicanus, for example, have
minute ossified lateral processes and other individuals lack these
214 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
processes; Microtus pennsylvanicus and some other species have
proportionately large lateral ossifications. If the well-developed
condition of the baculum in the microtines having rooted molars is
primitive, then within the genus Microtus those species having well-
developed bacula may be considered primitive.
The genera Lagurus and Neofiber have moderately developed or
well-developed lateral processes. Neofiber exhibits a tendency,
not prominent elsewhere, to have a proportionately smaller median
process rather than reduced lateral processes.
American species of Microtus (genus and subgenus) that have
moderately- to well-developed ossified lateral processes are M. town-
sendii, M. oeconomus, M. pennsylvanicus, M. montanus, and M.
chrotorrhinus. Microtus of other subgenera having this type of
baculum include M. (Herpetomys) guatemalensis, M. (Chilotus)
oregoni, and M. (Chionomys) longicaudus.
American species of Microtus (genus and subgenus) in which
the lateral ossifications are weakly developed or absent (although
cartilaginous lateral processes are present) include M. mexicanus
and M. californicus. In other subgenera, species of Microtus having
reduced lateral ossifications are M. (Pedomys) ochrogaster, M.
(Pitymys) pinetorum, M. (Pitymys) parvulus, M. (Pitymys) quasia-
ter, M. (Arvicola) richardsoni, and M. (Stenocranius) miurus.
The microtines are essentially holarctic in distribution. Both of
the tribes, the lemmings and the voles, as well as primitive repre-
sentatives of each tribe (not considering Ellobius) occur in both
the Old World and New World. It is not certain on which continent
(or continents) the Microtinae first differentiated. It is certain,
however, that at various times, both early and late in the evolution
of the subfamily, representatives have crossed from Eurasia to North
America or vice versa. Each of 10 or more microtines in the New
World is more closely related to some microtine in the Old World
than to any other microtine in the New World.
The similarities or differences in the baculum in Old World and
New World representatives placed in the same genus or subgenus,
or thought to be “companion species” have been commented upon
in accounts of Lemmus, Dicrostonyx, Clethrionomys, Lagurus, Arvi-
cola, Stenocranius, Chilotus, Chionomys, Pitymys, and in accounts
of Microtus agrestis as compared with M. pennsylvanicus, and
Microtus oeconomus (both Old World and New World).
The baculum in the Microtinae more closely resembles the bacu-
lum in the Cricetinae of the Old World than in the Murinae, or
than in any other rodents known to me. This resemblance suggests
relationship between Microtinae and Cricetinae.
THe BACULUM IN MICROTINE RODENTS 215
LITERATURE CITED
ARGYROPULO, A. I.
1933a. Die Gattungen und Arten der Hamster (Cricetinae Murray, 1866)
der Paliaarktik. Zeitschr. f. Siugetierkunde, 8:129-149, 27 figs. in
text.
1933b. Uber zwei neue paliaarktische Wiihlmiuse. Zeitschr. f. Saugetier-
kunde, 8:180-183, 3 figs. in text.
Cauuery, R.
1951. Development of the os genitale in the golden hamster, Mesocricetus
(Cricetus) auratus. Jour. Mamm., 32:204-207, 1 fig. in text.
CHAMBERLAIN, J. L.
1954. The Block Island meadow mouse, Microtus provectus. Jour.
Mamm., 35:587-589, 2 tables in text.
DEARDEN, L. C.
1958. The baculum in Lagurus and related microtines. Jour. Mamm.,
89:541-553, 1 fig. in text.
Dwr, R.
1943. L’os pénien des Campagnols de France du Genre Arvicola. Mam-
malia, 7:74-79, 10 figs. in text.
1954. Etude systématique de l’os pénien des Mammiféres (suite), Ron-
geurs: Muridés. Mammalia, 18:237-256, 14 figs. in text.
ELLERMAN, J. R.
1941. The families and genera of living rodents. Vol. II. Family Muri-
dae. The British Museum (Natural History), London, pp. xii +
690, 50 figs.
FRILEY, CHARLES E.
1947. Preparation and preservation of the baculum of mammals. Jour.
Mamm., 28:395-397, 1 fig. in text.
Hatt, E. R., and E. L. Cocxrum.
1953. A synopsis of the North American microtine rodents. Univ. Kansas
Publ., Mus. Nat. Hist., 5:373-498, 149 figs. in text.
Hamiiton, W. J., Jr.
1946. A study of the baculum in some North American Microtinae. Jour.
Mamm., 27:378-887, 8 figs. in text.
Hrpparp, C. W., and G. C. Rinker.
1942. A new bog-lemming (Synaptomys) from Meade County, Kansas.
Univ. Kansas Sci. Bull., 28:25-35, 3 figs. in text.
1943. A new meadow mouse (Microtus ochrogaster taylori) from Meade
County, Kansas. Univ. Kansas Sci. Bull., 29:255-268, 5 figs. in text.
Hinton, M. A. C.
1926. Monograph of the voles and lemmings (Microtinae), living and
extinct, Vol. I. British Museum (Natural History), London, pp.
xvi + 488, plus 15 plates, 110 figs. in text.
Marruey, R.
1953. Les Chromosomes des Muridae. Revue Suisse de Zoologie, 60:225-
283, avec les planches 1 4 4 groupant 84 photomicrographies, 98
figures et 5 schemas dans le texte.
216 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
1955. Nouveaux documents sur les chromosomes des Muridae. Problémes
de cytologie comparée et de taxonomie chez les Microtinae. Revue
Suisse de Zoologie, 62:163-206, avec 114 figures.
Miter, G. S.
1896. Genera and subgenera of voles and lemmings. North American
Fauna No. 12, pp. 1-85, 40 figs. and 3 plates in text.
Ocnev, S. I.
1948. The mammals of Russia (USSR) and adjacent countries (The
mammals of Eastern Europe and Northern Asia), Vol. 6. Publ.
Acad. Sci. USSR, pp. 1-587, 260 figs., 12 maps, and 11 color plates
in text (in Russian).
1950. The mammals of Russia (USSR) and adjacent countries (The mam-
mals of Eastern Europe and Northern Asia), Vol. 7. Publ. Acad.
Sci. USSR, pp. 1-736, 347 figs., 15 maps, and 10 color plates in
text (in Russian).
Rutu, E. B.
1934. The os priapi: A study in bone development. Anat. Rec., 60:231-
249, 16 figs. in 3 plates.
Situ, D. A., and J. B. Foster.
1957. Notes on the small mammals of Churchill, Manitoba. Jour. Mamm.,
88:98-115, 3 figs. and 3 tables in text.
WHEELER, B.
1956. Comparison of the Block Island “species” of Microtts with M.
pennsylvanicus. Evolution, 10:176-186, 4 figs. and 2 tables in text.
WurttE, J. A.
1951. A practical method for mounting the bacula of small mammals.
Jour. Mamm., 32:125.
ZIMMERMAN, K.
1955. Die Gattung Arvicola Lac. im System der Microtinae. Siugetier-
kundliche Mitteilungen, 3:110-112, 2 figs. in text.
Transmitted August 14, 1959.
28-774
Vol. 8.
Vol. 9.
ye
OF IU oT fe
(Continued from inside of front cover )
Life history and ecology of the five-lined skink, Eumeces fasciatus. By Henry
S. Fitch. Pp. 1-156, 26 figures in text. September 1, 4,
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.
An ecological study of the collared lizard (Crotaphytus collaris). By Henry
. Fitch. Pp. 213-274, 10 figures in text. February 10, 1956.
x field study of the Kansas ant-eating frog, Gastrophryne olivacea. By Henry
S. Fitch. Pp. 275-306, 9 figures in text. February 10, 1956.
Check-list of the birds of Kansas. By Harrison B, Tordoff. Pp. 307-359, 1
figure in text. March 10, 1956.
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.
Temperature responses in free-living amphibians and reptiles of northeastern
een By Henry S. Fitch. Pp. 417-476, 10 figures in text, 6 tables. June
1, 19
Food of the crow, Corvus brachyrhynchos Brehm, in south-central Kansas. . By
Dwight Platt. Pp. 477-498, 4 tables. June 8, 1956.
9. Ecological observations on "the woodrat Neotoma floridana. _By Henry S.
10.
Fitch and Dennis G. Rainey. Pp. 499-533, 3 figures in text. June 12, 1956.
Eastern woodrat, Neotoma floridana; Life history and ecology. By ‘Dennis
G. Rainey. Pp. 535-646, 12 plates, 18 figures in text. August 15, 1956.
Index. Pp. 647-675.
Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18
figures in text. December 10, 1955.
Additional records and extension of ranges of mammals from Utah. By
Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80.
December 10, 1955.
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.
Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming.
By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956.
The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6
figures in text. May 19, 1956.
Additional remains of the multituberculate genus Eucosmodon. By Robert
W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.
Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures
in text. June 15, 1956.
Comments on the taxonomic status of Peodemus 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.
Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp.
847-351. August 15, 1956.
A new bat (Genus Leptonycteris) from Coahuila. By Howard J.~ Stains.
Pp. 353-356. January 21, 1957.
. A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico.
iA
By Robert J. Russell. Pp. 357-361. January 21,
Geographic variation in the pocket gopher, Thomomys bottae, in Colorado.
By Phillip M. Youngman. Pp. 363-387, 7 figures in text. February 21, 1958.
New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones,
Jr. Pp. 385-888. May 12, 1958.
Pleistocene bats from San Josecito Cave, Nuevo Leén, México. By J. Knox
Jones, Jr. Pp. 389-396. December 19, 1958.
New subspecies of the rodent Baiomys from Central America. By Robert
L. Packard. Pp. 397-404. December 19, 1958.
Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405-
414, 1 figure in text, May 20, 1959.
Distribution, variation, and relationships of the montane vole, Microtus mon-
tanus. By Sydney Anderson. Pp. 415-511, 12 figures in text, 2 tables.
August 1, 1959.
Conspecificity of two pocket mice, Perognathus goldmani and P. artus. By
E. Raymond Hall and Marilyn Bailey Ogilvie. -Pp. 513-518, 1 map in text.
January 14, 1960.
Records of harvest mice, Reithrodontomys, from Central America, with de-
scription of a new. subspecies from Nicaragua. By Sydney Anderson and
J. Knox Jones, Jr. Pp. 519-529. January 14, 1960.
Small carnivores from San Josecito Cave (Pleistocene ), Nuevo Leén, México.
By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, 1960.
Pleistocene pocket gophers from San Josecito. Cave, Nuevo Leén, México.
By Robert J. Russell. Pp. 539-548, 1 figure in text, January 14, 1960.
More numbers will appear in volume 9,
(Continued on outside of back cover)
Vol. 10. 1.
2.
Sea ay
Wol.) 11 1.
(Continued from inside of back cover)
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,
Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.
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. )
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.
Birds found on the Arctic slope of northern Alaska. By James W. Bee.
Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958.
The wood rats of Colorado: distribution and ecology. By Robert B. Finley,
Jr. Pp. 218-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958.
Home ranges and movements of the eastern cottontail in Kansas. By Donald
W. Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, 1959.
Natural history of the salamander Aneides hardyi. By Richard F. Johnston
and Gerhard A. Schad. Pp. 573-585. October 8, 1959.
More numbers will appear in volume 10,
The systematic status of the colubrid snake, Leptodeira discolor Giinther.
By William E. Duellman. Pp. 1-9, 4 figures. July 14, 1958.
Natural history of the six-lined racerunner, Cnemidophorus sexlineatus. By
Henry S. Fitch. Pp. 11-62, 9 figures, 9 tables. September 19, 1958.
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.
A new snake of the genus Geophis from Chihuahua, Mexico. By John M.
Legler. Pp. 327-334, 2 figures in text. January 28, 1959.
A new tortoise, genus Gopherus, from north-central Mexico. By John M.
Legler. Pp. 335-348. April 24, 1959.
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.
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.
ee from Coahuila, México. By Emil K. Urban. Pp. 448-516. August 1,
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.
Another number will appear in volume II.
Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry
A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. July 8, 1959.
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.
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.
(MUS. COMP. 7901
UBRARY |
MAY 18 1960 |
UNIVERSITY OF KANSAS PUBLICATIONS
MuSsEUM OF NATURAL HIsTORY
Volume 12, No. 4, pp. 217-240, 12 figs.
May 2, 1960
A New Order of Fishlike Amphibia
From the Pennsylvanian of Kansas
BY
THEODORE H. EATON, JR., AND PEGGY LOU STEWART
UNIVERSITY OF KANSAS
LAWRENCE
1960
Unrversity OF Kansas PusiicaTions, MusEUM oF NATuRAL History
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson
Volume 12, No. 4, pp. 217-240, 12 figs.
Published May 2, 1960
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED IN
THE STATE PRINTING PLANT
TOPEKA, KANSAS
1960
28-2495
NWiUYd. OUI. AUUL
LIBRA:
MAY 18 1960
A New Order of Fishlike Amphibia #Aaysr |
From the Pennsylvanian of Kansps UNIVERSITY
BY
THEODORE H. EATON, JR., AND PEGGY LOU STEWART
INTRODUCTION
A slab of shale obtained in 1955 by Mr. Russell R. Camp from a
Pennsylvanian lagoon-deposit in Anderson County, Kansas, has
yielded in the laboratory a skeleton of the small amphibian Hes-
peroherpeton garnettense Peabody (1958). This skeleton pro-
vides new and surprising information not available from the holo-
type, No. 9976 K. U., which consisted only of a scapulocoracoid,
neural arch, and rib fragment. The new specimen, No. 10295 K. U.,
is of the same size and stage of development as the holotype and it
is thought that both individuals are adults.
The quarry, University of Kansas Museum of Natural History
Locality KAN 1/D, is approximately six miles northwest of Garnett,
Anderson County, Kansas, in Sec. 5, T. 19S, R. 19E, 200 yards south-
west of the place where Petrolacosaurus kansensis Lane was ob-
tained (see Peabody, 1952). The Rock Lake shale, deposited under
alternately marine and freshwater lagoon conditions, is a thin mem-
ber of the Stanton limestone formation, Lansing group, Missourian
series, and thus is in the lower part of the Upper Pennsylvanian.
Peabody (1958) placed Hesperoherpeton in the order Anthra-
cosauria, suborder Embolomeri, family Cricotidae. Study of the
second and more complete specimen reveals that Hesperoherpeton
is unlike the known Embolomeri in many important features. The
limbs and braincase are more primitive than those so far described
in any amphibian. The vertebrae are comparable to those of Ich-
thyostegalia (Jarvik, 1952), as well as to those of Embolomeri. The
forelimb is transitional between the pectoral fin of Rhipidistia and
the limb of early Amphibia. The pattern of the bones of the fore-
limb closely resembles, but is simpler than, that of the hypothetical
transitional type suggested by Eaton (1951). The foot seemingly
had only four short digits. The hind limb is not known.
The new skeleton of Hesperoherpeton lies in an oblong block of
limy shale measuring approximately 100x60 mm. After prepara-
tion of the entire lower surface, the exposed bones and matrix
were embedded in Bioplastic, in a layer thin enough for visibility
(219)
220 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
but giving firm support. Then the specimen was inverted and the
matrix removed from the opposite side; this has not been covered
with Bioplastic. The bones lie in great disorder, except that some
parts of the roof of the skull are associated, and the middle section
of the vertebral column is approximately in place. The bones of the
left forelimb are close together but not in a natural position. The
tail, pelvis, hind limbs and right forelimb are missing. Nearly all
the bones present are broken, distorted by crushing, incomplete and
scattered out of place, probably by the action of currents. The
complete skeleton, in life, probably measured between 150 and 200
mm. in length.
The specimen was studied at the Museum of Natural History,
University of Kansas, with the help of a grant from the National
Science Foundation, number NSF-G8624. The specimen was dis-
covered in the slab by Miss Sharon K. Moriarty, and was further
cleaned by the authors. Mr. Merton C. Bowman assisted with the
illustrations. We are indebted to Dr. Robert W. Wilson for critical
comments.
SKULL
Dorsal Aspect (Figs. 1, 2)
In reconstruction, the skull measures approximately 8.0 mm. dorso-
ventrally at the posterior end. The height diminishes anteriorly
to about 1.5 mm. at the premaxillary. The length is about 15.5 mm.
in the median line, or 24.0 mm. to the tip of the tabular, and the
width about 16.0 mm. posteriorly. The snout is blunt, continuing
about 1-2 mm. anterior to the external nares. Each of the tabulars
has a slender posterior process 5.0 mm. long, which probably met
the supracleithrum; the intertabular space is about 8.5 mm. wide.
The orbits are approximately 5.5 mm. in diameter and extend from
the maxillary to within about 3.0 mm. of the midline dorsally. The
pineal opening is 1.8 mm. anterior to the occipital margin of the
skull.
Reduction of bones at the back of the skull seems to have elimi-
nated any dermal elements posterior to the squamosal, while en-
largement of the orbit has removed most of the postorbital series,
leaving the squamosal as the only cheekbone. There is apparently
no jugal or postfrontal.
The squamosal of Acanthostega (Jarvik, 1952) is articulated
under the tabular and reaches forward and down, much as if it were
an opercular in reversed position. Internally, it must lie against
the otic capsule below the tabular, partially concealing the stapes.
A New Orper oF FISHLIKE AMPHIBIA 221
The bone that we suppose to be the squamosal of H. garnettense
is of similar shape, of about the same size and has internally an
articular surface at one corner, bounded by a pair of ridges in the
shape of a V. This articular surface probably fitted on a lateral
process extending from the roof of the neurocranium, over the front
of the otic capsule.
The premaxillary extends posterolaterally to a distance 5.5 mm.
from the midline and attains a width at its broadest point of about
15mm. The posterior edge is slightly concave and in part forms the
anterior border of the naris.
premaxillary nasal
maxillary external naris
lacrimal sn y prefrontal
basioccipital
opisthotic
exoccipital
supraoccipital
squamosal
Fic. 1. Hesperoherpeton garnettense Peabody. Skull, dorsal
view. Postorbital processes of the neurocranium are shown in
dotted outline. KU 10295, x 4.
The nasal is triangular and, with the lacrimal, forms the medial
border of the naris. The length of the medial side of the nasal bone
is approximately 5.0 mm., the transverse width is 3.8 mm., and the
extent of the posterolateral border is 5.5 mm.
o22 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
The maxillary meets the premaxillary lateral to the naris, borders
the naris posteroventrally, and continues posteriorly beneath the
orbit, of which it forms the external border. The maxillary is
about 8.5 mm. long, and immediately anterior to the orbit has a
maximum width of 1.3 mm.
The lacrimal fills the remaining rim of the narial opening between
the nasal and maxillary, and extends to the anterior edge of the
postorbital
supratemporal
parietal] frontal
squamosal maxillary premaxillary
Fic. 2. Hesperoherpeton garnettense Peabody. Skull, lateral view,
showing relatively large orbit and absence of smaller circumorbital
bones. KU 10295, x 4.
orbit. The length, from naris to orbit, is 4.2 mm.; the width ranges
from 1.0 mm. anteriorly to 2.5 mm. posteriorly.
The external naris is approximately 1.0 mm. in diameter. It is
slightly anterodorsal to the internal naris and 4.0 mm. lateral to
the midline.
The dorsal margin of the orbit appears to be formed by the
frontal. The anterior part of this margin, however, may be formed
by a prefrontal, which is not clearly set off by a suture. The frontal
extends 3.8 mm. in the midline, and anteriorly and laterally borders
the nasal and lacrimal, respectively. A faint pattern of pitting
radiates on the surface from the center of ossification of the frontal.
There is also a pit indicating the presence of a supraorbital sensory
pore.
The parietal bones enclose the pineal opening, approximately 2.5
mm. posterior to the suture with the frontal. The foramen is about
0.5 mm. in diameter. Laterally the parietal meets the medial angle
of the postorbital and the medial border of the supratemporal. No
bone of this animal shows the deep pitting and heavy ornamentation
characteristic of many primitive Amphibia.
A New Orpver oF FIsHLIKE AMPHIBIA 223
The postorbital meets the anterolateral corner of the parietal for
a distance of 0.5 mm., the anterior edge bordering the frontal bone
and the orbit for a combined distance of about 8.0mm. The lateral
margin is slightly convex, and is probably interrupted behind by the
anterior point of the tabular. Medially, the concave margin of the
postorbital meets the supratemporal for about 3.5 mm.
The supratemporal is thus wedge-shaped and located between
the parietal and the postorbital. The posterior edge of the supra-
temporal protrudes as a convex border slightly behind the end of
the parietal, and measures 3.0 mm. around the curve to the parietal
suture.
A B Cc D
Fic. 3. Hesperoherpeton garnettense Peabody. A, left squamosal, in-
ternal surface. B, left squamosal, external surface. C, right tabular
internal surface. D, right tabular, external surface. KU 10295, all x 4.
The squamosal (Fig. 3 A, B) is a large, somewhat rectangular
bone extending from the back of the orbit to the posterior extremity
of the cheek. It outlines almost entirely the posterior border of the
orbit, the ventrolateral portion of the cheek region, and the lateral
border of the top of the skull behind the orbit. Dorsally, the
squamosal meets the anterior half of the tabular and the lateral
border of the supratemporal. Near the anteroventral edge of the
squamosal there is a small pit, probably related to a postorbital
sensory pore in the skin.
The tabular (Fig. 3 C, D) is pointed anteriorly, where it probably
fits against the lateroposterior edge of the postorbital. The dorsal
part of the bone flares out and down, forming a small otic notch at
a point halfway back. Posteriorly, the flange attains a dorsoventral
width of 2.0 mm. at the edge of the notch. The slender posterior
224 UnIversIry OF Kansas Pusts., Mus. Nat. Hist.
process of the tabular which continues beyond the flange is ap-
proximately 0.5 mm. in diameter and 5.0 mm. long.
Ventral Aspect (Fig. 4)
The palatal view of the skull shows the paired premaxillary,
maxillary, palatine, pterygoid, and quadrate bones. The openings
for the internal nares, the ventral orbital fenestrae, and the sub-
temporal fossae are readily recognized. The quadrate processes ex-
tend posteriorly leaving a large gap medially at the posterior end of
the skull.
premaxillary
ethmosphenoid ? internal naris
maxillary
e
°
°
J
°
°
°
°
oe 9
pterygoid
quadrate
A B
Fic. 4. Hesperoherpeton garnettense Peabody. Palate recon-
structed; ventral aspect at left, showing teeth, dorsal aspect at
right. KU 10295, x 4.
A New Orper OF FISHLIKE AMPHIBIA 225
The left quadrate appears to be in place on the posterior prong
of the pterygoid. The dorsal side of the quadrate is grooved be-
tween two anterolaterally directed ridges. The groove, which prob-
ably held the end of the stapes, extends about half the width of the
quadrate itself. The width of the quadrate is 4.0 mm., the length is
4.5 mm. medially and about 2.0 mm. laterally. In ventral view the
quadrate appears to project laterally, but is incomplete and its shape
uncertain. The distance from the posterior end of the quadrate to
the visible posterior edge of the orbital fenestra, which opens
ventrally, is 10.0 mm.
This region between the quadrate and the orbit is occupied by a
pterygoid with three projections. Anteriorly, the pterygoid outlines
most of the posterior edge of the orbit (a distance of about 6.5 mm.).
A lateral process separates the orbit from the subtemporal fossa. A
posteriorly directed edge defines the fossa, which extends about 6.5
mm. anteroposteriorly. The lateral process of the pterygoid termi-
nates 10.0 mm. from the midline. Both the lateral and posterior pter-
ygoid processes are approximately 2.0 mm. wide. The greatest width
of the subtemporal fossa is about 2.0 mm. The medial border of
the orbital fenestra is missing, but apparently consisted of the ptery-
goid for at least the posterior half.
Along the posterior edge of the orbital fenestra, there is a narrow,
dorsally projecting flange of the pterygoid. The lateral opening of
the orbit is approximately 7.5 mm. wide.
The remaining border of the orbital fenestra on the anterior and
medial sides is formed by a bone occupying the position of palatine
and vomer; for convenience we designate this as palatine. When
reconstructed in its probable position in relation to the pterygoid,
the left palatine lacks a section, on its medial and posterior edges,
measuring about 2.5 mm. by 9.0 mm. The lateral margin of the
palatine is convex; about 5.5 mm. anterior to the orbit this margin
curves into a strong anteriorly pointing projection, medial to
which is seen the internal narial opening. The remaining anterior
edge is slightly convex, smoothly rounded, and meets the midline
about 9.0 mm. anterior to the pterygoid.
The void area medial to the palatine and anterior to the ptery-
goid does not fit any bone which we can recognize as the parasphe-
noid. It is thus suspected that this area is covered in part by the
missing edge of the palatine and partly by an anteromedial exten-
sion of the pterygoid. Of course a parasphenoid may also have
been present.
226 University oF Kansas Pusts., Mus. Nat. Hist.
The position, length, and shape of the premaxillary shown in
palatal view (Fig. 4) are primarily based upon the dorsal appear-
ance since ventrally most of it cannot be seen. At the point where
it forms the anterior border of the internal naris, the premaxillary is
slightly wider than the maxillary and seems to become narrower as
it approaches the midline.
The ethmosphenoid, which we cannot identify, may have been
exposed in a gap between the premaxillary and the palatine. The
gap measures approximately 8.0 mm. wide and ranges up to 1.0 mm.
anteroposteriorly.
. The maxillary begins at a suture with the premaxillary lateral
to the naris and continues posteriorly, bordering the orbit with a
width of about 1.2 mm. It then tapers to a point approximately 2.0
mm. anterior to the lateral projection of the pterygoid. The width
of the maxillary at this point is 0.8 mm. and the posterior end is
broken; probably when complete it approached the pterygoid, and
either met the latter or had a ligamentous connection with it. As
nearly as can be determined, the total length of the maxillary is
approximately 12.0 mm.
The teeth on the maxillary are small and seem to be in two
longitudinal rows. The palatine bears two large, grooved teeth
anteriorly; the first is approximately 1.0 mm. posteromedial to the
naris and the second is about 3.0 mm. posterior and slightly lateral
to the naris. The flat ventral surfaces of the palatine and pterygoid
bear numerous small teeth distributed as shown in Fig. 4.
Braincase and Occipital Region (Fig. 5)
The parts of the neurocranium are scattered, disconnected and
incomplete, but it is possible to make out a number of features of
the otico-occipital section with fair assurance. In posterior view
the notochordal canal and foramen magnum are confluent with
each other, and of great size relative to the skull as a whole. The
notochordal canal measures 2.8 mm. in diameter, and the foramen
magnum about 4.0 mm. The crescent-shaped supraoccipital rests
on the upright ends of the exoccipitals, but between the latter and
the basioccipital no sutures can be seen. Probably the whole pos-
terior surface of the braincase slanted posteroventrally; conse-
quently the rim of the notochordal canal was about 3.0 mm. behind
the margin of the parietals.
The U-shaped border of the notochordal canal is a thick, rounded
bone, comparable in appearance to the U-shaped intercentra of the
A New Orper OF FISHLIKE AMPHIBIA 227
vertebrae. This bone apparently rested upon a thinner, troughlike
piece (Fig. 5 B) forming the floor of the braincase. The latter is
broad, shallow, concave, open midventrally and narrowing an-
teriorly to form a pair of articular processes. Since no sutures can
be seen in this structure, it probably is the ventral, ossified portion
of the basioccipital. Watson (1926, Fig. 4 B) illustrates the floor
of the braincase in Eusthenopteron, with its more lateral, anterior
portion labelled prootic, but in our specimen the corresponding part
could scarcely have formed the anterior wall of the otic capsule,
foramen magnum
supraoccipital opisthotic
supratemporal
exoccipital
quadrate pterygoid
A
notochordal canal
Fic. 5. Hesperoherpeton garnettense Peabody, KU
10295, * 4. _A, occipital view of skull; B, basioccipital
bone in dorsal (internal) view.
being entirely in the plane of the floor. The two articular surfaces
anteriorly near the midline suggest that a movable joint existed be-
tween the otico-occipital part of the braincase and the ethmos-
phenoid part, as in Rhipidistia (Romer, 1987). We have found
nothing in the specimen that could be referred to the ethmos-
phenoid; it may have been unossified.
The otic capsules appear to have rested against lateral projections
of the basioccipital. The single otic capsule that can be seen (the
228 University OF Kansas Pusts., Mus. Nat. Hist.
right) is massively built, apparently ossified in one piece, with a
shallow dorsomedial excavation, probably the vestige of a supra-
temporal fossa. On the lateral face is a broad, shallow depression
dorsally, and a narrower, deeper one anteroventrally; these we
suppose to have received the broader and narrower heads of the
stapes, respectively. The posterior wall of the otic capsule we
have designated opisthotic in the figure. Anterior to the otic cap-
sule the lateral wall of the braincase cannot be seen, and may not
have been ossified.
The roof of the braincase is visible in its ventral aspect, extending
from approximately the occipital margin to a broken edge in front
of the parietal foramen, and laterally to paired processes which
overlie the otic capsules directly behind the orbits (see dotted out-
lines in Fig. 1). Each of these postorbital processes, seen from
beneath, appears to be the lateral extension of a shallow groove
beginning near the midline. Presumably this section of the roof
is an ossification of the synotic tectum. It should be noted that the
roof of the braincase proper is perfectly distinct from the overlying
series of dermal bones, and that the parietal foramen can be seen
in both. The roof of the braincase in our specimen seems to have
been detached from the underlying otic capsules and the occipital
wall.
The bone that we take to be the stapes is blunt, flattened (perhaps
by crushing), 5.0 mm. in length, and has two unequal heads; its
width across both of these is 4.0 mm. The length is appropriate to
fit between the lateral face of the otic capsule and the dorsal edge of
the quadrate; the wider head rests on a posterodorsal concavity
on the otic capsule, and the smaller fits a lower, more anterior
pit. Laterally the stapes carries a short, broad process that
probably made contact with a dorsally placed tympanic membrane.
Thus the bone was a hyomandibular in the sense that it articulated
with the quadrate, but it may also have served as a stapes in sound-
transmission. It contains no visible canal or foramen.
Mandible (Fig. 6)
The crushed inner surface of the posterior part of the left mandible
and most of the external surface of the right mandible are pre-
served in close proximity. Although the whole length of the tooth-
bearing margins is missing, some parts of six elements of the right
mandible can be seen. The pattern of sutures and the general con-
tour closely resemble those of Megalichthys (Watson, 1926, Figs.
37, 38) and other known RBhipidistia.
A New ORovER OF FISHLIKE AMPHIBIA 229
The anteroposterior length of the mandible is about 23.8 mm.,
and the depth is 3.8 mm. The dentary extends approximately 17.6
mm. back from the symphysis, and its greatest width is probably
2.0 mm. Its lower edge meets all the other lateral bones of the
jaw. The splenial and postsplenial form the curved anteroventral
half of the jaw for a distance of about 9.0 mm. The fragmented
articular, on the posterior end of the jaw, is 4.0 mm. long and 2.0
mm. deep, exhibiting a broken upper edge; presumably the surface
for articulation with the quadrate was a shallow concavity, above
the end of the articular.
articular surface dentary
articular surangular angular postsplenial splenial
Fic. 6. Hesperoherpeton garnettense Peabody. Right mandible, lateral
view, KU 10295, 4. External surfaces are pitted; broken surfaces
are coarsely stippled.
VERTEBRAE (Fig. 7)
The vertebrae that are visible from a lateral view are crushed
and difficult to interpret. It is possible, nevertheless, to see that
the trunk vertebrae resemble those of Ichthyostegalia (Jarvik, 1952,
Fig. 13 A, B), except that the pleurocentra are much larger. A few
parts of additional vertebrae can be seen, but they are so scattered
that it is impossible to be sure of their original location. Therefore
comparisons between different regions cannot yet be made.
The U-shaped intercentrum encloses the notochord and occupies
an anteroventral position in the vertebra. Anteriorly, each inter-
centrum articulates with the pleurocentra of the next preceding
vertebra by slightly concave surfaces. Dorsolaterally there is an
articular surface for the capitulum of the rib.
The two pleurocentra of each vertebra are separate ventrally as
well as dorsally, but form thin, broad plates of about the same
height as the notochord. The lateral surface appears to be de-
pressed, allowing, perhaps, for movement of the rib. Above each
pleurocentrum, on the lateral surface of the neural arch, there is a
short diapophysis for articulation with the tuberculum of the rib.
The margin of the neural spine is convex anteriorly and concave
posteriorly, the tip reaching a point vertically above the postzyga-
230 University OF Kansas Pusts., Mus. Nat. Hist.
pophysis. The prezygapophysis of each vertebra articulates with
the preceding postzygapophysis by a smooth dorsal surface. One
nearly complete neural arch shows (Fig. 7 B) a pit above the neural
canal, clearly corresponding to the canal for a dorsal ligament shown
by Jarvik in Ichthyostega. Indeed this view of the neural arch and
intercentrum together brings out the striking resemblance between
the vertebrae of Hesperoherpeton and those of the Ichthyostegids.
The rounded intercentrum in both is an incomplete ring enclosing
the notochordal canal.
neural spine
els diapophysis
« dorsal ligament
"SY prezygapo-
physis
parapophysis diapophysis
diapophysis (7 :\ parapo-
notochordal intercentrum physis
canal
notochordal canal ‘parapophysis pleurocentrum
A B Cc
Fic. 7. Hesperoherpeton garnettense Peabody. A, End view of incomplete
vertebra, probably near anterior end of column. B, Neural arch and intercen-
trum in end view, showing probable association. C, Left lateral view of
trunk vertebra. All figures: KU 10295, x4.
The shape, in end view, of a partly preserved neural arch (Fig.
7 A) seems to account for the incompleteness of the intercentrum
just mentioned; the ventral edge of the arch is emarginate in such
a way as to fit the dorsal surface of the notochord. The dorsal
portion of this neural arch is not present (either broken or not yet
TasBLE 1—AvERAGE MEASUREMENTS OF THE TRUNK VERTEBRAE (in mm.).
NUMBERS IN PARENTHESES INDICATE THE NUMBER OF PIECES AVAILABLE FOR
MEASURING
ParRTs Ant.-post. Dors.-vent. | Transv. width
Neural spine s.oiccescasecgsa,s aleve, os ese (G3) 3 OCB) y a alseeteicmerrer aera
Neural spine and arch........ 2.0 (4) Ws eat: Foe Wa baer
Neuralicanaliys fiat iseincres. 2.0 (4) PAO G0) Oh CH)
Intercentrume cies oer 1-5 16) avo (4) 3.0 (1)
Pleurocentrum. .% .:.:....0.54 +0. 1.5 (38) 30) 3(2)
Ce
A New Orper oF FIsHLIKE AMPHIBIA 231
ossified), but the opening of the neural canal is comparable in
width to the foramen magnum. Hence this vertebra may be one
of the most anterior in the column. In comparison with the trunk
vertebrae seen farther posteriorly it appears that there may be a
progressive ossification of neural arches toward their dorsal ends,
and of intercentra around the notochord, with probable fusion of
the intercentra and neural arches in the posterior part of the trunk.
The notochord seems to have been slightly constricted by the in-
tercentra, but not interrupted.
RIBS
The proximal ends of the ribs expand dorsoventrally to a width
approximately four times that of their slender shafts. The tuber-
culum and capitulum on each of the trunk ribs are separated only
by a shallow concavity. These two articular surfaces are so situ-
ated that the rib must tilt downward from the horizontal plane.
The shaft flares terminally in some ribs, and the distal end is convex.
Ribs in the trunk region differ little if any in size. Five that can
be measured vary in length from 5.0 to 7.0 mm. One short, bent
rib 8.5 mm. long perhaps is sacral or caudal.
PECTORAL GIRDLE (Figs. 8, 9, 10)
The right scapulocoracoid is almost complete, and the left one
is present but partly broken into three pieces, somewhat pushed out
of position. With the advantage of this new material, we may com-
ment on the scapulocoracoid of H. garnettense as described by Pea-
body (1958). In size and contour, the slight differences between
the type (KU 9976) and the new skeleton (KU 10295) are con-
sidered to be no more than individual variation. We have redrawn
the type (Fig. 8) in order to show the resemblances more clearly.
The small sections that were missing from the type are present
in KU 10295. The jagged edge directly posterior to the area oc-
cupied by the neural arch in the type extends 0.5 mm. farther back
in our specimen. The angle formed between the recurved dorsal
ramus and the edge of the ventral flange is seen in our specimen
to be less than 90°. The glenoid fossa, appearing as a concave
articular surface for the cap of the humerus, was in part covered
by cartilage and shows as “unfinished” bone (Peabody, 1958, p. 572);
this area is more oval than triangular, as Peabody thought. The
obstruction of a clear view of this part of the type is the result
of the accidental position of a neural arch. The raised portion
232 University OF Kansas Pusts., Mus. Nat. Hist.
MB.
Fic. 8. Hesperoherpeton garneitense Peabody. Type specimen redrawn.
Right scapulocoracoid in external view (at left), and internal view (at right).
KU 9976, x 4.
supraglenoid
foramen
articular
surface
area covered
by clavicle
9 10
Fic. 9. Hesperoherpeton garnettense Peabody. Right scapulocoracoid in ex-
ternal view, showing part of interclavicle, and position occupied by clavicle.
The specimen is flattened and lies entirely in one plane. KU 10295, x 4.
Fic. 10. Hesperoherpeton garnettense Peabody. Right clavicle in external
view. Anterior edge to right. KU 10295, x 4.
A New OrperR OF FISHLIKE AMPHIBIA 233
immediately dorsal to the glenoid fossa exhibits an unfinished sur-
face, suggesting the presence of either cartilage or a ligament.
The right clavicle is complete, and resembles a spoon having a
slender handle. The dorsal tip of the handle is L-shaped. The
expanded ventral part is convex externally, and rested upon the
anteroventral surface of the scapulocoracoid. The lateral edge next
to the “stem” is distinctly concave, abruptly becoming similar in
contour to the opposite edge, and giving the impression of an un-
symmetrical spoon. The left clavicle is present in scattered frag-
ments, its dorsal hooklike end being intact.
The posterior end of the interclavicle lies in contact with the right
scapulocoracoid. There are short lateral processes at the point
where the interclavicle was overlapped by the clavicles, but we can-
not be sure of the extent of this bone anteriorly or posteriorly.
The presumed left cleithrum, a long rectangle, is approximately
equal in length to the rodlike stem of the clavicle, and is about
as wide as the dorsal L-shaped tip of the clavicle. The posterior
end of the cleithrum presumably met the tip of the clavicle, while
the rest of it was directed anteriorly and a little dorsally. There
seems to be a small articular surface near the anterior extremity
which suggests the presence of a supracleithrum. The upper border
of the cleithrum is slightly convex and the lower concave.
FORELIMB (Fig. 11)
The left forelimb is the only one present and appears to be nearly
complete, although the elements are scattered almost at random.
The only parts of the forelimb known to be missing are two sub-
terminal and two terminal phalanges, probably of the first and
third digits, and the proximal end of the second metacarpal. The
smooth and relatively flat surfaces suggest an aquatic rather than
terrestrial limb; only the proximal half of the humerus bears any
conspicuous ridges or depressions. As we restore the skeleton of
the limb, several features are remarkable: The humerus, ulna, and
ulnare align themselves as the major axis of the limb, each carrying
on its posterior edge a process or flange comparable to those in the
axial series of a rhipidistian fin. The remaining elements take posi-
tions comparable to the diagonally placed preaxial radials in such
a fin. The digits appear to have been short, perhaps with no more
than two phalanges. There is only one row of carpals present (the
proximal row of other tetrapods). A second and third row would
be expected in primitive Amphibia; if they existed in Hesperoher-
peton they must either have been wholly cartilaginous or washed
934 Unrversiry OF Kansas Pusts., Mus. Nat. Hist.
away from the specimen. Neither of these alternatives seems at all
likely to us in view of the well-ossified condition of the elements that
are present, and the occurrence of both the proximal carpals and
the metacarpals. The space available for metacarpals probably
could not have contained more than the four that are recognized.
intermedium
ulnare
Fic. 11. Hesperoherpeton garnettense Peabody. Left forelimb,
showing characters of both a crossopterygian fin and an am-
phibian foot. KU 10295, x 4.
The proximal end of the humerus is more rounded anteriorly
than posteriorly, and has a thin articular border that bore a car-
A New Orpber OF FISHLIKE AMPHIBIA 235
tilaginous cap as the primary surface for articulation with the
scapulocoracoid. Although the unfinished surface of the head ex-
tends down the anterior margin about a third the length of the
humerus, the shaft has been broken and so twisted that the distal
part is not in the same plane as the proximal. Immediately posterior
to the cartilaginous cap is a round, deep notch bordered posteriorly
by the dorsal process of the head.
The shaft is longer and narrower than would be anticipated in a
primitive amphibian limb (cf. Romer, 1947). The distal end bears
two surfaces for articulation with the radius and ulna. The full ex-
tent of the former surface was not determined because the more an-
terior part of the expanded end is represented only by an impression.
The surface nearest the ulna was partially rounded for articulation
with that element, the remaining posterior edge being broadly con-
cave. The most striking feature of the humerus is a slender hook-
like process on the posterior edge near the distal end, probably
homologous with (1) the posterior flange on the “humerus” in Rhipi-
distia, and (2) the entepicondyle of the humerus in Archeria (Romer,
1957) and other tetrapods.
The radius is about the same width proximally as distally. The
curvature of the shaft is approximately alike on both sides. Distally
the surface is rounded for articulation with the radiale and perhaps
the intermedium.
The proximal end of the ulna is similar to that of the radius but is
slightly larger. Posteriorly, there is a short, broad expansion re-
sembling the entepicondyle of the humerus, and even more nearly
like the postaxial flanges in a crossopterygian fin.
The ends of the radiale are expanded and rounded, the entire bone
being approximately twice as long as wide. The three sides of the
intermedium are similarly convex. The surface of this bone is un-
finished, showing that it must have been embedded in cartilage.
The ulnare is conspicuously similar to the ulna in bearing a posterior
hooklike expansion, and is larger than the radiale.
The four metacarpals are slightly expanded proximally and dis-
tally. Although measurements of length and width are tabulated
below (Table 2), we are not certain of the sequence of these bones
in the row.
The dimensions of the two proximal phalanges are alike. The
shape of these elements is similar to that of the metacarpals. The
two terminal phalanges are somewhat triangular in shape, the lateral
edges being concave and the proximal convex.
236 Universiry OF Kansas Pusts., Mus. Nat. Hist.
TABLE 2.—APPROXIMATE MEASUREMENTS OF THE FORELIMB (in mm.)
Dimensions
ELEMENT Width
Length
Proximal | Midway Distal
BEUMELUS sone ca ie cite ele tore 16.0 5.0 2.0 abr
RAGUSA <4 ee ee icatic cee 9.0 4.0 1.5 Sho
nae SS os ese Cac. Geet 8.5 4.5 15 Suis
Riauislen cs. cnet ew wink Sa 3.0 2.0 15 2.0
Pntermedinum.: .4 <s ctoee cle es 5 we 2.0 ae
UN ereess® > chia coe ice Salers 325 20 2.0 245
Metacarpal 1 eee 4.5 2.5 1.0 2.0
Metacarpal 1 AR AS 4.5 3.0? £5 2.5
Metacarpal ORE ere 4.0 2.0 1.5 2.0
Metacarpal LD gehen S20 Bco 1.0 15
Brosnan Phalanye SSAn Ne oon 2.0 1.5 1.0 1.5
Proximal Phalanx B........ 2.0 5 1.0 125
Terminal Phalanx <A........ eo Was 1.0 1.0
Terminal Phalanx B........ 1.5 tas 1.0 1.0
COMPARISONS AND DISCUSSION
Apparently primitive rhipidistian characters in Hesperoherpeton
are: Braincase in two sections, posterior one containing an expanded
notochordal canal; lateral series of mandibular bones closely re-
sembling that of Megalichthys, as figured by Watson (1926); tabular
having long process probably articulating with pectoral girdle; lack
of movement between head and trunk correlated with absence of
occipital condyle; sensory pits present on frontal and squamosal.
Although we are unable to separate, by sutures, the vomers from
the palatines, the palatal surface of these bones and of the ptery-
goids is studded by numerous small teeth, as in Rhipidistia (Jarvik,
1954) and some of the early Amphibia (Romer, 1947). The stapes
apparently reaches the quadrate, and could therefore serve in hyo-
stylic suspension of the upper jaw.
The pectoral limb has an axial series of bones carrying hooklike
flanges on their posterior edges. The other bones of the limb show
little modification of form beyond the nearly flat, aquatic type seen
in Rhipidistia. No distinct elbow or wrist joints are developed.
Characters of Hesperoherpeton common to most primitive Am-
phibia, in contrast with Crossopterygii, are: Nares separated from
edge of jaw; stapes having external process that may have met a
tympanic membrane, thus giving the bone a sound-transmitting
function. Apparently none of the opercular series was present.
A New Oprper oF FIsHLIKE AMPHIBIA 237
There are two large palatal teeth, slightly labyrinthine in char-
acter, adjacent to each internal naris. The scapulocoracoid, as
shown by Peabody (1958), is Anthracosaurian in structure, as are the
long-stemmed clavicles. The limbs have digits rather than fin-lobes,
although the digital number apparently is four and the number of
bones in the manus is less than would be expected in a primitive am-
phibian. The vertebrae are similar to those of Ichthyostegids, as
described by Jarvik (1952), except that the pleurocentra are much
larger.
In addition to this remarkable combination of crossopterygian
and amphibian characters, Hesperoherpeton is specialized in certain
features of the skull. The orbits are much enlarged, probably in cor-
relation with the diminutive size of the animal, and this has been
accompanied by loss of several bones. The frontal and squamosal
nearly meet each other, and both form part of the rim of the orbit.
The bones of the posterior part of the dermal roof are greatly re-
duced, and there is none behind the squamosal except the projecting
tabular; there is no indication of quadratojugal, jugal, intertem-
poral or postparietal. The foramen magnum is enormous. The ex-
ternal surfaces of the bones of the skull are nearly smooth.
Is it possible that the “primitive” and “specialized” features of
this animal are actually larval? Are they not just the kind of char-
acters that would be expected in an immature, aquatic embolomere
of Pennsylvanian time? For several reasons we do not think this is
the case. Except for the anterior part of the braincase, there is no
indication that the skeleton was not well ossified. The postaxial
processes on the humerus, ulna and ulnare could scarcely have been
larval features only, since they are so clearly homologous with those
in adult Rhipidistia; a larval limb should indeed be simple, but its
simplicity is unlikely to involve paleotelic adult characters. The
scapulocoracoid of our specimen is of practically the same shape and
size as that in the only other known individual, the type; this would
be probable if both were adults, but somewhat less likely if they
were larvae of a much larger animal. The form of the stapes, tabu-
lar and otic notch suggest a functional tympanic membrane, which
could not have occurred in a gill-breathing larva. On the other
hand, an adult animal of pigmy size might be expected to have
large orbits, large otic capsules and a large foramen magnum.
We conclude that Hesperoherpeton lived and sought food in the
weedy shallows at the margin of a pond or lagoon, and that for much
of the time its head was partly out of water (Fig. 12). The animal
could either steady itself or crawl around by means of the paddle-
238 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist.
like limbs, but these probably could not be used in effective loco-
motion on land. Like the Ichthyostegids, it probably swam by means
of a fishlike tail.
Fic. 12. Hesperoherpeton garnettense Peabody. Probable appearance
in life. 0.5.
TAXONOMY
Evidently Hesperoherpeton is a small, lagocn-dwelling survivor
of the Devonian forms that initiated the change from Crossopterygii
to Amphibia (Jarvik, 1955). It shows, however, that this transition
did not affect all structures at the same time, for some, as the brain-
case with its notochordal canal, the mandibular bones and axial limb
bones, are unchanged from the condition normal for the Rhipidistia,
but most other characters are of amphibian grade. To express these
facts taxonomically requires that Hesperoherpeton be removed from
the family Cricotidae, suborder Embolomeri, order Anthracosauria,
and placed in a new order and family of labyrinthodont Amphibia.
Order PLESIOPODA
(plesios, Gr., near, almost; podos, Gr., foot)
Labyrinthedontia having limbs provided with digits, but re-
taining posterior flanges on axial bones as in Rhipidistia, with-
out joint-structure at elbow and wrist essential for terrestrial
locomotion; neurocranium having separate otico-occipital sec-
tion, large notochordal canal, no occipital condyle, as in Rhipi-
distia; nares separate from rim of mouth; pectoral girdle an-
thracosaurian; vertebrae having U-shaped intercentrum and
paired, but large, pleurocenira.
Probably associated with the characters of the order, as given
above, are the connection of pectoral girdle with skull, and the pres-
ence of a tympanic membrane, the stapes functioning in both sound-
transmission and palatoquadrate suspension.
A New Orver OF FISHLIKE AMPHIBIA 239
Family HESPEROHERPETONIDAE
Orbits and foramen magnum unusually large in correlation
with reduced size of animal; squamosal forming posterior mar-
gin of orbit; circumorbital series absent (except for postorbital ) ;
sensory pits on squamosal and frontal.
Characters defining the family are evidently the more specialized
cranial features, which probably evolved during Mississippian and
early Pennsylvanian times.
The definition of the genus and species may be left to rest upon
Peabody's (1958) original description and the present account,
until the discovery of other members of the family gives reason for
making further distinctions.
SUMMARY
Hesperoherpeton garnettense Peabody (1958), based on a scapu-
locoracoid and part of a vertebra, was originally placed in the order
Anthracosauria, suborder Embolomeri, family Cricotidae. A new
skeleton from the type locality near Garnett, Kansas (Rock Lake
shale, Stanton formation, Upper Pennsylvanian), shows that the ani-
mal has the following rhipidistian characters: Large notochordal
canal below foramen magnum, otico-occipital block separate from
ethmosphenoid, postaxial processes on three axial bones of forelimb,
pectoral girdle (probably) articulated with tabular. Nevertheless,
Hesperoherpeton has short digits, an anthracosaurian type of pec-
toral girdle, an otic rather than spiracular notch, nostrils separate
from the mouth, and vertebrae in which the intercentrum is U-shaped
and the pleurocentra large but paired. The stapes reaches the quad-
rate.
Hesperoherpeton is placed in a new order, PLESIOPODA, on the
basis of the characters stated above, and a new family, HESPERO-
HERPETONIDAE. Specialized characters of the family include:
Reduction of circumorbital bones, bringing the squamosal to the
edge of the orbit, loss of certain bones of the temporal region, and
relative enlargement of the orbits and foramen magnum, in cor-
relation with the diminutive size of the animal. The structural
characters of Hesperoherpeton suggest to us that it lived in the
shallow, weedy margins of lagoons, rested with its head partly out
of water, and normally did not walk on land.
240
Eaton, T.
1951.
Jarvik, E.
1952.
1954.
1955.
Moorg, R.
1944,
UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist.
LITERATURE CITED
H., Jr.
Origin of tetrapod limbs. Amer. Mid]. Nat., 46: 245-251.
On the fish-like tail in the ichthyostegid stegocephalians. Meddel.
om Gr¢gnland, 114: 1-90.
On the visceral skeleton in Eusthenopteron with a discussion of the
parasphenoid and palatoquadrate in fishes. Kgl. Svenska Veten-
skapsakad. Handl., 5: 1-104.
The oldest tetrapods and their forerunners. Sci. Monthly, 80:
141-154.
C., Frye, J. C., and Jewett, J. M.
Tabular description of outcropping rocks in Kansas. Kansas State
Geol. Surv. Bull., 52: 137-212.
PEasopy, F. E.
1952.
1958.
Romer, A.
1937.
1947.
1957.
Petrolacosaurus kansensis Lane, a Pennsylvanian reptile from Kansas.
Univ. Kansas Paleont. Contrib., Vertebrata, Art. 1: 1-41.
An embolomerous amphibian in the Garnett fauna (Pennsylvanian)
of Kansas. Jour. Paleont., 32: 571-573.
S.
The braincase of the Carboniferous crossopterygian Megalichthys
nitidus. Mus. Comp. Zool. Bull., 82: 1-73.
Review of the Labyrinthodontia. Mus. Comp. Zool. Bull., 99: 1-368.
The appendicular skeleton of the Permian embolomerous amphibian
Archeria. Univ. Michigan Contrib. Mus. Paleont., 13: 103-159.
Warson, D. M. S.
1926.
The evolution and origin of the Amphibia. Phil. Trans. Roy. Soc.
London, (B) 214: 189-257.
Transmitted January 18, 1960.
(ei
28-2495
Museum OF NATURAL HisToRY
HARVARD
Volume 12, No. 5, pp. 241-296,6 figs. |_UiUEEESITY
March 7, 1962 —
Natural History of the Bell Vireo,
Vireo bellii Audubon
BY
JON CG. BARLOW
UNIVERSITY OF KANSAS
LAWRENCE
1962
UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY
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Durrant. Pp. 1-549, 91 figures in text, 80 tables. August 10, 1952.
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87 tables: August 25, 1952.
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ar Pe ee Lewis L. Sandidge. Pp. 305-338, 5 figures in text. Aagust
8. The silky pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker.
Pp. 339-347, 1 figure in text. February -15,
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849-472, 47 figures in text, 4 tables. April 21, 1954.
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ne soe By Sydney Anderson. Pp. 489-506, 2 figures in text.
y
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Arizona. By Terry A. Vaughan. Pp. 507-512... July 23,
9. Mammals of the San Gabriel mountains of California. By Tens ‘A. Vaughan.
Pp. 518-582, 1 figure in text, 12 tables. November 15, 1954.
10. A new bat (Genus Pipistrellus) from northeastern Mexico. By Rollin H.
Baker. Pp. 588-586. 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 “he pocket gopher, Cratogeomys castanops, in Coa-
uila,, Mexico. By Robert J. Russell and Rollin. H. Baker. Pp. 501-608.
March 15, 1955.
18. 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.
Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956.
Vol. 9. 1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18
figures in text. December 10, 1955.
2. Additional records and extension of ranges of mammals from Utah. By
Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80.
December 10, 1955.
(Continued on inside of back cover)
UNIVERSITY OF KANSAS PUBLICATIONS
MusEUM OF NATURAL HIsTORY
Volume 12, No. 5, pp. 241-296, 6 figs.
March 7, 1962
Natural History of the Bell Vireo,
Vireo bellii Audubon
BY
JON C. BARLOW
UNIVERSITY OF KANSAS
LAWRENCE
1962
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
Editors: E. Raymond Hall, Chairman, Theodore H. Eaton, Jr.,
Henry S. Fitch
Volume 12, No. 5, pp. 241-296, 6 figs.
Published March 7, 1962
UNIVERSITY OF KANSAS
Lawrence, Kansas
MUS. COMP. 2001
LIBRARY
JUL 13 1962 |
HARVARD
UNIVERSITY =
PRINTED BY
JEAN M. NEIBARGER, STATE PRINTER
TOPEKA, KANSAS
1962
29-1506
Natural History of the Bell Vireo,
Vireo bellii Audubon
BY
JON C. BARLOW
CONTENTS
PAGE
PATENTS Lyle. 0) COR a) tA te, a ee 243
BERODUCTION) nite ho Oo. PO i tess de ee ee 245
Ee NOWLEDGMENNTS® ....).,. <etell ows faeiee acc Sy seer oe 245
MIPTHORS(OR STUDY ) id. 5.0.54. /agedeient ey See goatee 246
EDIT AREAS h.5 Dea ROI hn ng casa a ee ee eee 247
Wonsiderations“of Habitat «0.2 bs. 5.652... tae eee 248
SEASONAL) MOVEMENT ©; (7.0) 5 «0 -faeatitien ho sels OE Ree 250
EDV AMMA OPTI: srs Wie pee. ws Pe ol ed 250
Dalle Deparinre:i.., Suet k eet: BAS eee 251
UNERAL WOEHAVIOR whist. Ane PA ee 2) es ee oe 252
Bataleon UN ea NE oy be MINE 3 a o's 6 AT oo EO 252
Poraging and.lood) Habitse.ij7. 262 4252 22. eee 252
BARRING Ltr WeAaty : Berka A Ww) 0 wh PA eee yess eis 253
BERCATTZATIONG AS ity... Sten tye a Boies, gt h FLYt eS he CR en ae 254
Singing: Postures!) ives bess 1 eb ee. aos 255
Bight song oo Sy Coie ue, Ssh See Ri cee ee 255
mally Hrequency Of SONG -92).2020 Stee eee bee Be o>: Sa 255
Mapes of Vocalizations 55500362 ee 255
MIMEMETORTATTEY, 140) 1 (ces ies es ues Belen. dh ens a ciate eg past Re 258
Bistablishment,of Verritoryic, ct. 0.20) abies ARE ee Bad 259
SizeiOWMerritOneso een. aes Oil Ae Matise Gap Saas eee 259
Permanence Or) a Cmmitories:.. 2i2he x tsa oe a Ai, SS 260
Maintenance of Rerritonyinut 2 arate sae oe 260
Aggressive Behavior of the Female ...................... 264
interspecific Relationships). 10093550. . YudGt es 264
PPISCUSSION Wma nen occlu hate hoe oy tal nealy, We dee chee Beant 265
SE GUUSHIP ISEELAVION G20 25 25-0 rs a sre aks ela os Be ae wie he 267
Displays and: Postusest: fees eee easy ook ln eee 268
RDARCLISSIOINE 20.4 tains a ene amiie hive Be ong ne eit ae 270
244 UNIversIry OF Kansas Pusts., Mus. Nat. Hist.
PAGE
SELECTION OF NEST-SITE AND NESTBUILDING ................-. 272
Bountled imme ec eee eye eee aimee ane eee ce hee tar ore eta aaa 274
Gathering of Nesting Materials .....................-..-. 276
Length and Hours of Nestbuilding ....................-.. 277
Abortive Nestbuilding Efforts ..................--.------ 277
FRemestaing cy eerste ey ric hie dic eraee Mote eke econ Rice ss asda 277
SSNS Ee eae he oN ENE Riese eae eae Pagan 277
REGUAYINCG AND LNCUBATION) yas) fe) p wise pe aio ola aisha eye eee 278
Big glanirag ye ie eat Won Maibiaeveia ey, aieiajere leer eet 278
CltelesiZze he / eca bs tie OME ee edu ek ae tn cee eee 279
RnCubatOnn bs Chik a ase ABS a ee Bees ie a ae 280
The Roles of the Sexes in Incubation ..................-.. 280
Relief of Partners in Incubation ....................-.-.. 283
INIRSTRING PERIOD: ack sl 4 Uae bees oe Pe Ro ya a ere 283
Hatching Sequence: ..'.0.5.....4.-.).0e Beek 2h Ree 283
Development of the Nestlings ..............-..--..-+--5- 284
Parental "Behavior £042 2))s0.85anene ss 4425. eee ee eee 285
eeding of the Nestlings -..:-4)....-..:5-4-.+ BS 286
INiest"SamitatiOn . 6 hese Lb naw bakers Beelal ate ea SR ee 287
Bled ging, 2) 242i ie ce tocnee Mine gent 2034 Reel OR ree 287
West Parasites: «fick st bea snaked ee Ges AUER, SE eae 287
REDGLING AGIBE |. cinco ocak Geek Ce eeeNG Oe eee ey een 288
SECOUG BEOOUS: 4c) 4 FIN a Ee Bee eee e Sean eee 288
REPRODUGIIVE SUCCESS’... io). 6 bs ahae cele os SR ree 289
IBEhAVION FG sek kd pue Pee eee bas GEE acaps 6 Ree Re 290
Predation’. s.r ae 291
Gowhbird Parasitism «626.24 bo) hose cen SE a ee 291
GuNENTARY ie alee eis ooo j Me ciaela ce Aiecs Ma Rib ee ten et ee 292
NATuRAL History OF THE BELL VIREO 245
INTRODUCTION
The Bell Vireo ( Vireo bellii Aud.) is a summer resident in riparian
and second growth situations in the central United States south of
North Dakota. In the last two decades this bird has become fairly
common in western, and to a lesser extent in central, Indiana and
is apparently shifting its breeding range eastward in that state
(Mumford, 1952; Nolan, 1960). In northeastern Kansas the species
breeds commonly and occurs in most tracts of suitable habitat.
The literature contains several reports dealing exclusively with
the Bell Vireo, notably those of Bennett (1917), Nice (1929), Du
Bois (1940), Pitelka and Koestner (1942), Hensley (1950) and
Mumford (1952). Bent (1950) has summarized the information
available on the species through 1943. Nolan (1960) recently com-
pleted an extensive report based on a small, banded population at
Bloomington, Monroe County, Indiana. He validated for this
species many points of natural history previously based on estimates
and approximations, especially concerning the post-fledging life of
the young and the movement of the adults from one “home range”
to another in the course of a single season.
None of these reports, however, has emphasized the ritualized
behavioral patterns associated with the maintenance of territory
and with courtship. Among the North American Vireonidae, the
behavior of the Red-eyed Vireo (Vireo olivaceus) is best docu-
mented (Sutton, 1949; Lawrence, 1953; Southern, 1958). With this
species authors have concentrated on the mechanics of the breeding
season and their reports contain little discussion of the aggressive
and epigamic behavior of the bird.
The amount of information on the ritualized behavior of the Bell
Vireo and related species heretofore has been meager. I observed
breeding behavior from its inception in early May through the
summer of 1960. It is hoped the resulting information will serve
as a basis of comparison in future studies of behavior of vireos; such
ethological data are becoming increasingly important, especially as
an aid in systematics.
ACKNOWLEDGMENTS
To professors Frank B. Cross, Henry S. Fitch, and Richard F.
Johnston of the Department of Zoology of the University of Kansas
I am grateful for comments and suggestions in various phases of
the study and the preparation of the manuscript. Professor Johnston
246 UNIVERSITY OF Kansas PuBis., Mus. Nar. Hist.
also made available data on the breeding of the Bell Vireo from the
files of the Kansas Omithological Society. I am indebted to my
wife, Judith Barlow, for many hours of typing and copy reading.
Mrs. Lorna Cordonnier prepared the map, Thomas H. Swearingen
drew the histograms, and Professor A. B. Leonard photographed
and developed the histograms. Dr. Robert M. Mengel contributed
the sketch of the Bell Vireo and George P. Young prepared the
dummy Bell Vireo used in the field work. Thomas R. Barlow,
Donald A. Distler, Abbot S. Gaunt, John L. Lenz, Gary L. Packard,
A. Wayne Wiens, and John Wellman assisted in various phases of
the field work.
METHODS OF STUDY
Daily observations were made from May 11 to June 26 in 1959
and from April 15 through July 15 in 1960. Six additional visits
were made to the study area in September of 1959, and ten others
in July and August, 1960. Periods of from one hour to eleven hours
were spent in the field each day, and a total of about five hundred
hours were logged in the field.
Each territory was visited for at least five minutes each day but
more often for twenty minutes. The breeding activities of the pairs
were rarely synchronous. Consequently several stages in the cycle
of building were simultaneously available for study.
Nine young and one adult were banded in 1959. No Bell Vireos
were banded in 1960. Individual pairs could be recognized be-
cause of their exclusive use of certain segments of the study area
and by the individual variation in the song of the males. Sexes
were distinguishable on the basis of differences in vocalizations and
plumages.
Most nests were located by the observer searching, watching a
pair engaged in building, or following a singing male until the
increased tempo of his song indicated proximity to a nest. As the
season progressed and the foliage grew more dense, it became
increasingly difficult to locate completed nests. Blinds were un-
necessary because of the density of vegetation. Observations were
facilitated by a 7 x 50 binocular. Data were recorded on the spot
in a field notebook. Eggs were numbered by means of Higgins
Engrossing ink as they were laid.
Individual trees in which males sang most were marked over a
three-week period. Then the distances between the most remote
perches were paced. These distances aided in determining the
NATuRAL History OF THE BELL VIREO 247
size of the territories. The general configuration of the vegetation
within each territory determined the location of one or more bound-
aries of the territory. Each territory was given a number, 1, 2, 8,
etc., as it was discovered; consequently there is no numerical re-
lationship between the designations of the territories established in
1959 and 1960. Nests within a territory were designated as 1-a, 1-b,
1-c, etc.
Although experimentation was not a primary source of data, it
proved useful in certain instances. A stuffed Blue Jay elicited
mobbing behavior from nesting pairs. A dummy Bell Vireo elicited
both agonistic and epigamic behavior from nesting pairs, depending
on the phase of the nesting cycle.
The temperature at the beginning of each day’s work was taken
by means of a Weston dial thermometer. A hand counter and a
pocket watch having a second hand were used in determining such
data as frequency of song and periods of attentiveness by the sexes.
Histological cross-sections, prepared by A. Wayne Wiens, of the
ventral epidermis of both sexes were used to study brood patches.
STUDY AREA
The intensive field work was on a 39-acre tract (fig. 1) extending
approximately %o of a mile west from U.S. highway 59, which in
1959-1960 constituted the western city limit of Lawrence, Douglas
County, Kansas. The eastern boundary of the study area is ap-
proximately 1% miles southwest of the County Courthouse in Law-
rence. The eastern ten acres is associated with the Laboratory of
Aquatic Biology of the University of Kansas. The 15 acres adjacent
to this on the southwest is owned by the University of Kansas En-
dowment Association, but is used by Mr. E. H. Chamney for the
grazing of cattle. This portion is bounded on the west by a stone
fence, beyond which lies a 14-acre field of natural prairie owned by
Dr. C. D. Clark that is bordered on the extreme west by a narrow
thicket of elm saplings.
The principal topographic feature of the area is an arm of Mount
Oread, that rises some 80 feet above the surrounding countryside.
About 200 feet from the crest of the southwestern slope of the hill
a 40-foot-wide diversion terrace directs run-off toward the two-acre
reservoir that is the source of water for eight experimental fish ponds
of the laboratory.
The predominant shrub-vegetation consists of Osage orange
(Maclura pomifera), honey locust (Gleditsia triacanthos), and
248 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
American elm (Ulmus americana). These saplings, ranging in
height from 8 to 25 feet, grow in dense thickets as well as singly
ts
ow - »
VS
a \S >
-<S_-
S Noe
a“
)
Nae
\
3 x
- Z
PR SIG
Yip
°o
Wi.
yoy
SECOND GROWTH
TREES
(hiss Ss ST]
scale of feet
Fic. 1. Map of the study area near the University of Kansas Laboratory of
Aquatic Biology. The dashed lines mark the approximate territorial boundaries
of the original nine pairs of Bell Vireos from May 1960 to early June 1960.
and in clumps of twos and threes. Larger trees of these same species
grow along the crest of the hill, along the eastern and southeastern
boundaries of the area, and along the stone fence separating Uni-
versity land from that owned by Dr. Clark. A dense growth of
coralberry (Symphoricarpos orbiculatus) forms the understory just
below the crest of the hill. Isolated clumps of dogwood (Cornus
drummondi) and hawthorn (Crataegus mollis) are scattered
throughout the area. These species of shrubs grow densely along
the stone fence. The isolated thicket on the Clark land is composed
primarily of elm and boxelder (Acer negundo), but includes scat-
tered clumps of dogwood, Osage orange, and honey locust. Poplars
(Populus deltoides) are the only large trees in this area.
The open areas between the thickets are grown up in red top
(Triodia flava), bluestem (Andropogon scoparius), Switchgrass
(Panicum virgatum), Kentucky bluegrass (Poa pratensis), bush
clover (Lespedeza capitata) and mullen (Verbascum thapsus).
Shrubby vegetation occupies about 65 per cent of the total area,
but in the Clark portion constitutes only about 35 per cent of the
ground cover.
Considerations of Habitat
Nolan (1960:226), summarizing the available information on
habitat preferences of the Bell Vireo, indicates that this species
tolerates “a rather wide range of differences in cover.” He pointed
NATURAL History OF THE BELL VIREO 249
out that a significant factor in habitat selection by this species may
be avoidance of the White-eyed Vireo (V. griseus) where the two
species are sympatric.
In Douglas County where the Bell Vireo is the common species,
the White-eyed Vireo reaches the western extent of its known
breeding range in Kansas. At the Natural History Reservation of
the University of Kansas, where both species breed, the Bell Vireo
occurs in “brush thickets in open places” (Fitch, 1958:270) and the
White-eyed Vireo occupies “brush thickets, scrubby woodland and
woodland edge” (Fitch, op. cit., 268). Along the Missouri River
in extreme northeastern Kansas, Linsdale (1928:588-589) found the
White-eyed Vireo “at the edge of the timber on the bluff,-and in
small clearings in the timber,” while “the Bell Vireo was character-
istic of the growths of willow thickets on newly formed sand bars.”
Elsewhere in northeastern Kansas I have found the Bell Vireo in
shrubbery of varying density and often in habitat indistinguishable
from that occupied by White-eyed Vireos at the Natural History
Reservation. In the periphery of the region of sympatry the rarer
species is confronted with a much higher population density of the
common species and consequently might well be limited primarily
to habitat less suitable for the common species. This would seem
to be the case in eastern Kansas, presuming that interspecific com-
petition exists.
The Bell Vireo has followed the prairie peninsula into Indiana,
aided by the development of land for agriculture. In nearby Ken-
tucky where thousands of miles of forest edge are found, and where
little brushy habitat of the type preferred by the Bell Vireo occurs,
the White-eyed Vireo is abundant whereas the Bell Vireo is un-
known as a breeding bird (R. M. Mengel, personal communication).
In more central portions of the area of sympatry, nevertheless,
the two species do occur within the same habitat (Ridgway, 1889:
191; Bent, 1950:254) and occasionally within the same thicket
(Ridgway, in Pitelka and Koestner, 1942:105); their morphological
and behavioral differences, although slight, probably minimize inter-
specific conflict. The Bell Vireo and the Black-capped Vireo
(V. atricapillus) have been found nesting in the same tree in Okla-
homa by Bunker (1910:72); the nest of the black-cap was situated
centrally and that of the Bell Vireo peripherally in the tree. Bell
Vireos invariably place their nests in the outer portions of trees and
peripherally in thickets. This placement would further obviate
interspecific conflict with the white-eye since its nests are placed
centrally in the denser portions of a thicket.
250 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist.
A critical feature of the habitat preferred by the Bell Vireo is the
presence of water. In far western Kansas this species is restricted
to riparian growth along the more permanent waterways. This in
itself is not adequate proof of the significance of water supply be-
cause thicket growth in that part of the state is found only along
waterways. The 20 areas over the state that I have visited where
Bell Vireos were present were closely associated with at least a
semi-permanent source of water. Fifteen other areas indistinguish-
able from the 20 just mentioned, but lacking a permanent supply
of water, also lacked Bell Vireos. Nevertheless areas in which Bell
Vireos typically nest are decidedly less mesic than those frequented
by White-eyed Vireos.
Once the Bell Vireo was probably more local in its distribution
being restricted to thickets associated with permanent water. Clear-
ing of woodland for agricultural and other use, and subsequent
encroachment of second growth concomitant with the creation of
man-made lakes and ponds, has greatly increased the available
habitat for this bird. The preferred species of shrubs for nesting
are reported (Bent, 1950:254) to be various wild plums (Prunus
sp.). The widespread distribution and abundance of the exotic
Osage orange has greatly augmented the supply of trees suitable
for nesting.
SEASONAL MOVEMENT
Arrival in Spring
The subspecies of the Bell Vireo breeding in Kansas, V. b. bellii,
winters regularly from Guerrero and the Isthmus of Tehuantepec
south to Guatemala, El] Salvador, and northern Nicaragua (A. O. U.
Check-list, Fifth Edition, 1957:469-470). In the United States
migrating birds are first recorded in early March (Cooke, 1909:119).
The Bell Vireo is a relatively slow migrator, moving primarily at
night and covering little more than 20 miles at a time (Cooke,
op. cit. 119). The average date of arrival, based on 27 records, for
northeastern Kansas is May 8; the earliest record is April 22, 1925,
from Manhattan, Riley County, Kansas (fig. 2-A).
In 1959 the first bird arrived at the study tract about May 5. No
additional birds were heard singing until the third week of the
month, in which eight new males were noted. As mentioned, in
1960 field work was begun in mid-April and the study area was
traversed daily. No birds were detected until late afternoon of
May 8, when one, presumably a male, was seen foraging.
NATURAL HIsTORY OF THE BELL VIREO 251
Lawrence (1953:50) has reported that males of the Red-eyed
Vireo precede females in the breeding area by as much as two
weeks; the male Red-eyed Vireo forages but sings little in the pre-
nesting period. The male Bell Vireo arrives first at the breeding
area but precedes the female by only a few days. On the morning
of May 4 the first male was singing from a number of perches while
ranging over an area of seven acres. This area encompassed terri-
Number of Records
A
Fic. 2. Seasonal movement as indicated by the curve for spring
arrival (A), based on the earliest dates for 27 years, and the curve
for autumn departure (B), based on the latest dates for 21 years
in northeastern Kansas.
tories later occupied by three pairs, 2 (1960), 4 (1960), and 5
(1960). Late on the afternoon of May 4 the first courtship songs
were heard and the first male was seen with a mate at 6:20 p.m.
Eight additional males arrived from May 6 through May 18. A
tenth male was discovered in the vicinity of territory 9 (1960) on
June 18, 1960.
Fall Departure
The average date of departure for 21 years in northeastern Kansas
is September 3 (fig. 2-B). The earliest date is August 14 from Con-
cordia, Cloud County, Kansas (Porter, unpublished field notes).
The latest date is September 27 (Bent, 1950:262) from Onaga, Pot-
tawatomie County, Kansas. In 1959 the last vireo was seen at the
study tract on September 14. The birds do not all depart at the
252 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
same time. On September 1 there were still five singing males in
the study area; by September 10 there were three and on September
13, only one.
GENERAL BEHAVIOR
Flight
In “straight-away” flight the Bell Vireo undulates slightly. In a
typical flight several rapid, but shallow, wing beats precede a fixed-
wing glide of from 1 to 15 feet in length. Because the wings are
extended horizontally during the glide, the bird does not move dis-
tinctly above or below the plane of flight. The White-eyed Vireo
generally appears to be slower and more lethargic in flight than the
Bell Vireo. In the breeding season most flights of the Bell Vireo
are no longer than a few feet between adjacent shrubs and trees,
but occasional sustained flights are as long as 300 feet. The birds
fly as low as 2 feet above ground, but have often been observed as
high as 70 feet above the ground.
In courtship and protracted territorial disputes, where chase be-
tween sexual partners or a pair of antagonists occurs, looping flights
are observed. The wings are beaten as the birds climb and many
aerial maneuvers are performed in the course of the glide.
Foraging and Food Habits
The Bell Vireo has been characterized as a thicket forager (Ham-
ilton, 1958:311; Pitelka and Koestner, 1942:104), but in my ex-
perience it is not restricted to low level strata; birds forage from
ground level upward, both in thickets and isolated trees ranging in
height from 8 feet to 65 feet. The tendency to forage at higher
levels is in part dictated by the presence of tall trees within the
various territories.
Territories 1 through 7 (1960) contained from three to ten trees
surpassing 40 feet in height. They grew singly or in small groves.
The Bell Vireos foraged fully 20 per cent of the time in these trees.
Food was sought throughout the leaf canopy.
Behavior in foraging in larger trees followed a routine pattern.
Typically a pair alighted in a tree at a height of 15 feet. Then the
female hopped to a perch a foot above the one upon which she
landed. The male succeeded her to the perch she had previously
occupied. The pair in effect spiraled around some large, essentially
upright, branch, in foraging. The birds usually reached higher
NAaATuRAL History OF THE BELL VIREO 253
perches in this manner rather than by flying upward 10 to 15 feet
to them. This manner of progression within a tree is reminiscent
of a similar habit of the Cyanocitta jays. Presumably, the habit of
the Bell Vireo of foraging in higher strata is facilitated by the ab-
sence of other species of arboreal foraging vireos.
Chapin (1925:25) found the Bell Vireo to be more insectivorous
in its food habits than any other North American vireo. He found
99.3 per cent of all food contained in 52 stomachs to be of animal
origin. Only three times have I seen a Bell Vireo take food of
vegetable origin. On September 9, 10, and 14, 1959, I noted a male
eating wild cherries over a period of 65 minutes of observation.
Chapin (1925:27) noted that beginning in July vegetable matter
represented 1.57 per cent of the bird’s subsistence, and thereafter
slightly more until fall migration.
Animal food, consisting primarily of insects and spiders, is actively
sought along branches and under leaves. Often a foraging bird will
leap to the underside of a branch and hover, mothlike, beneath a
cluster of leaves while extracting some insect. Some individuals
hung upside down on small branches, paridlike, while foraging.
Lawrence (1953:710), and Southern (1958:201) have recorded
similar behavior of the Red-eyed Vireo. Occasionally, I have seen
a Bell Vireo fly from a perch and capture an insect in the manner
of a flycatcher. The birds do not appear to be adept at this type
of food-getting. Nolan (1960:242) mentions Bell Vireos holding
hard-bodied insects by means of their feet while breaking the
exoskeleton with the beak to obtain the soft parts. Southern (1958:
201) recorded a female Red-eyed Vireo foraging on the ground; I
have seen a Bell Vireo on the ground but once, and it was gathering
nesting material.
Bathing
On May 14, 1960, in a rill that empties into the northeastern edge
of the reservoir a female flew down from a perch six inches above
the surface, barely dipped into the water, flew to a perch 12 inches
above the water, violently shook her ruffled body feathers, quivered
her wings, and rapidly flicked her fanned tail. The entire procedure
was repeated three times in five minutes. She was accompanied by
a singing male that did not bathe.
Nolan (1960:241) reports a male Bell Vireo bathing by rubbing
against leaves wet with dew; he notes that the White-eyed Vireo
bathes in a similar manner. Southern (1958:201) twice observed
954 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Red-eyed Vireos bathing in water that dropped from wet leaves.
In my study area in 1960, only territories 7, 8, 9, and 10 were not
immediately adjacent to permanent water. The pairs of Bell Vireos
in those territories presumably had to reply on wet vegetation for
bathing.
VOCALIZATIONS
The male Bell Vireo begins to sing regularly soon after its ar-
rival in spring. Some daily singing continues following the cessa-
tion of breeding activities until departure of the species in late
summer or early fall. The highest sustained rate of song occurs
on the first and second days of nest building. Because careful rec-
ords of meteorological data were not kept, I cannot significantly
correlate rates of song and specific temperatures and other weather
conditions. Frequency of song was reduced when the temperature
rose above 90° F., as it did on many days in June, 1960. Nice
(1929:17) mentions a similar decrease in singing when the tempera-
ture exceeded 85° F.
Passerine birds typically sing at a high rate throughout courtship
and nestbuilding, but at a markedly lower rate thereafter. Most
vireos are atypical in this respect. In the study area in 1960 Bell
Vireos sang more often than Robins, Mockingbirds, Field Sparrows,
Brown Thrashers, Catbirds, and Doves breeding in the same habitat,
about as often as the Meadow Larks in the adjacent fields, and less
often than Painted Buntings.
The Bell Vireo seems to sing less often in the undisturbed state
than when aware of the presence of an observer. Observations from
my car, at a site approximately equidistant from territories 1 (1960),
2 (1960), 4 (1960), and 6 (1960) indicate that the rate of song
during incubation is decidedly less when no disturbing influence
is present. Normally, in this period, song aids in maintaining con-
tact between the members of a pair, serving to locate the male as
he forages. Mumford (1952:230) noted that the males often came
out to meet him as he entered their territories, singing as they ap-
proached, The male typically continues to sing for some time after
the intruder has departed. Here the song acquires the additional
functions of alerting the female to danger and threatening the tres-
passer. Even after allowance is made for this reaction to disturb-
ance, Bell Vireos sing more often than most of their nesting asso-
ciates, and, on a seasonal basis, they are vocal for a much longer
time.
NATURAL HisToRY OF THE BELL VIREO 25D
Singing Postures
In the normal singing posture the body of the Bell Vireo is main-
tained at an angle of 85° to the horizontal. Occasionally, during
nest building, I have observed the body held at angles as severe as
80° from the horizontal.
The head of the White-eyed Vireo is distinctly bobbed up and
down, two or three times, during the utterance of a song phrase.
A bob involves a deliberate withdrawal of the head towards the
body and subsequent sharp, almost vertical, extension of the neck.
The head of the Bell Vireo does not bob, although it vibrates as the
song is delivered.
Flight Song
The Bell Vireo does not have a distinctive flight song; in fact,
it rarely sings or calls while in flight. Nolan (1960:240) has re-
corded a male singing the normal song while in flight. Sharp
scold-notes are uttered in mid-air when a bird is agitated or actually
attacking an enemy. These notes and songs recorded by Nolan
hardly qualify as flight song, for this term implies use of a dis-
tinctive vocalization not uttered in other circumstances.
Daily Frequency of Song
In the morning, Bell Vireos usually began singing a few minutes
before sunrise. Their songs were invariably preceded in the study
area by those of Western Kingbirds, Robins, Mourning Doves,
Mockingbirds, Cardinals and Meadow Larks. Bell Vireos sang rela-
tively little after 6:30 p.m., even on the longest days of the year.
The latest daytime singing that was recorded was seven songs at
7:18 p.m. on June 20, 1960. A Cardinal in the vicinity sang for
a full hour after this.
Types of Vocalizations
Six vocalizations were readily distinguishable in the field. These
are divisible into songs and call notes.
1. Primary song. It has been described by Pitelka and Koestner
(1942:103) as an “irregular series of harsh and sharp, but slurred
notes preceded by a few distinct notes of the same quality and
ending with a decided ascending or descending note of similar
harshness.” The terminal note may also be somewhat abbreviated
and intermediate between an ascending or descending note. The
song is sometimes delivered as a couplet that consists of a phrase
ending on a descending note. This delivery is typical of incubation
and later renesting. During early season activities, the bird utters
256 UnIveErSITY OF Kansas PuBLs., Mus. Nat. Hist.
a phrase ending on the descending note as many as 15 times before
a phrase ending on an ascending note is heard.
A sonagram of a single phrase, one of several recorded on May
9, 1960 (the third day of building of nest 1-b 1960), consists of
10 notes, the first of which is distinct. The remaining notes are
slurred. This phrase is 1.4 seconds in length.
Songs are delivered most rapidly in the course of territorial dis-
putes and defense. The song is loudest in times of nestbuilding and
periods of aggressive behavior. At these times, on clear, calm days,
the songs are audible 100 yards away. Singing in the nestling period
and post-breeding season is audible at distances of no more than
50 feet; such notes have been termed “whisper songs.” Table 1 sum-
marizes singing rates at different periods of the nesting cycle in
several situations and under various weather conditions.
Songs are of equal frequency in the immediate vicinity of the
nest and elsewhere in the territory. Nice (1929:17) also found this
to be true. Perches can be almost at ground level or as high as 60
feet. Forty per cent of my data on song concern singing at heights
of more than 20 feet. As indicated in foraging, the lack of competi-
tion from aboreal species of vireos presumably contributes to the
use of higher perches by Bell Vireos.
No female song was recorded in 1959, but on May 26, 1960, a
female was heard to sing once. She appeared at nest 1-f (1960)
shortly after the male arrived. Unlike him, she did not participate
in building, but seemed to be inspecting the nest. After 30 seconds
she sang once—a low garbled phrase—and also scolded once. After
this she left. In the meantime the continuously singing male moved
two feet away from the nest, then back to it and resumed con-
struction.
The song of the female signaled to the male her departure.
Pitelka and Koestner (1942:103) heard a female sing twice after
she replaced the male on the nest. Females of three other species
of vireos, the Black-capped Vireo, V. atricapillus (Lloyd, 1887:295),
the Philadelphia Vireo, V. philadelphicus (Lewis, 1921:33), and the
Latimer Vireo, V. latimeri (Spaulding in Pitelka and Koestner,
1942:103) have been heard singing. Lewis and Spaulding also sug-
gest that the song of the female functions as a signal prior to
exchange at the nest.
The primary song identifies the singer as a male Bell Vireo. It
aids in securing a mate and in warning potential adversaries; also,
the song is a signal in certain situations and serves to locate the male.
NATURAL HisTORY OF THE BELL VIREO 257
TABLE 1. REPRESENTATIVE SINGING RATES OF BREEDING BELL VrrEos. ALL
Rates WERE AT AIR TEMPERATURES LEss THAN 86° F. Eacu INSTANCE REp-
RESENTS APPROXIMATELY 380 MINUTES OF OBSERVATION.
Average
Circumstance Instances | rate per
minute
PUPA CHIONGOR SAGE! bs) ie ete. Rca ace ahs Wyn. e Huck 2 6.3
Mernvorialidispute sapere Seep ei ae chsiamisls leer «alors 5 12.8
esthurdine ees. O27. TS Qak. 215. 2ROkS Aaa. Ae 6 7.0
LAG EA 90-2 ae pe PROUR REISS SOR IRA FR MLA A ne 1 3.0
MCHOMINOH Miers Crictioh Seise tistics ccs cece ck ne ateee one 6 3.9
Exchange of partners:in the incubation period......... 1 4.0*
FORA GIT ers cee sence Merete Re rivs. cess Rays cha Sees er atcuats, crete 2 2.2
SeIVIOPMIT Peg RONG Ae eiorsete aicie ote Staite aus sesh oyalsiohoe lone auei ets 1 28 .6*
REVI, HONEY OS. CNOA. oes AB. Bea. OH Baie. 1 IR OE
Over-all average rate per minute 6.3
* Not sustained; data representative of periods less than 5 minutes in length.
2. Courtship song. It is here termed the “congested” song and
is comparable to the adult “run-on” song mentioned by Nolan (1960:
240). The congested song is a squeaky version of the primary song
and is given when birds are engaged in pair-formation, nestbuilding,
and egglaying. The delivery is rapid and the sound can be likened
to that made by rapidly scraping a bow across a taut violin string.
Nolan (in Mumford, 1952:230) is probably speaking of this song
when he describes a “tuneless” song that “had a jerky, sputtering
quality that characterizes part of the song of the Ruby-crowned
Kinglet (Regulus calendula).” More recently (1960:240) he applies
the adjectives “twanging,” “Bobolink-like,” “bubbling,” “jerky,” and
“squeaky.” This song is often blended with the primary song and
is audible for 75 feet.
A specialized version of the congested song is associated with pre-
and post-copulatory display but differs from the typical squeaky
performance in terminating in two ascending notes reminiscent of
the ascending phrase of the primary song.
3. Distress call. It was heard only once, when a captured bird
was being freed from a net. When the bird was almost disentangled
it uttered 10 high-pitched, plaintive notes. The quality of the notes
suggested a relationship to the song phrase rather than to other
types of vocalization. A nesting pair of Bell Vireos, 10 feet away,
became extremely excited when they heard the distress notes. They
“scolded” vigorously and flew around my head at a distance of
six feet.
2—1506
258 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
4. Alarm note. This is a specialized, three-note call of the male
and was heard only from the onset of pair-formation through
early nestbuilding. This whinnying, flickerlike call; phonetically
eh-eH-EH, each succeeding note of which is louder than the one
before, is given whenever the male is disturbed by an unfamiliar
object. This call is generally succeeded by the chee, but occasion-
ally blends into an extended “whinny,” and is typically given from
some perch affording an unobstructed view of the offending object.
The male stretches his neck and cocks his head, the wings and tail
are not flicked or fanned, and no feather tracts are erected. The
bird, nevertheless, flits nervously from perch to perch when uttering
the call.
5. The zip. The male has a special “scold” note of his own that
is heard when an intruder first approaches the nest. Phonetically
it is zip-zip-zip. It is not so loud as the chee, and the delivery is
more deliberate than that note. If the intruder remains near the
nest, the zip is usually replaced by the chee.
6. The generalized call note or chee. The call notes associated
with several situations are combined under this subheading since
all can be rendered in English by the same phonetic equivalent—
chee. The chee associated with nestbuilding is of moderate pitch
and delivered deliberately at a rate of about 40 per minute. The
feeding call of the adults is a soft slurred chee, while that of the nest-
lings has a mewing quality. In general, the chee utilized in signal
situations consists of a few repetitions of the basic note emitted at
a moderate pitch. The chee associated with hostile and courtship
behavior is higher pitched and the delivery is much more rapid,
approximately 200 per minute. Nolan (1960:240) reports a con-
tinuous rate of 25 per five seconds when an adult Bell Vireo is
alarmed. The chee of extreme anxiety is a loud emphatic buzz,
phonetically ZZ-ZZ-ZZ-ZZ.
TERRITORIALTY
The Bell Vireo exhibits “classic” passerine territoriality. Within
a specific area, a pair of this species carries out pair-formation,
courtship activities, copulation, nesting, rearing the young, and
foraging. With the cessation of reproductive activities, a pair con-
tinues to restrict its other daily activities to the same general area,
NATURAL History OF THE BELL VIREO 259
Establishment of Territory
In early May the segment of the total suitable habitat within
which a Bell Vireo restricts its activities is not rigidly defined and
the first male of the season ranges over an area too large to be
maintained permanently—one that seems greatly to exceed the
needs of breeding. Male 1 (1960), for instance, was first seen forag-
ing over an area of approximately seven acres. With the influx
of other males, portions of this large tract were usurped and the
territory of the original male was gradually reduced to an area
of little more than an acre.
In this initial period, a male becomes identified with a large
area but is restricted to an area of nearly typical size by the en-
croachment of other males. Territorial disputes in this period often
involve physical contact, as well as protracted sessions of high-
intensity singing at rates exceeding three hundred song-phrases
per hour.
Eventually the carrying capacity of the habitat is reached and
no further partitioning occurs. The beginning of nestbuilding
coincides with this relative stabilization of the territorial boundaries.
Through the remainder of the cycle of behavior associated with any
one nest, all activity is that of the occupant pair within its terri-
tory.
Size of Territories
The nine original territories established in 1960 varied in size
from .26 acre to 3.1 acres (Table 2). Fitch (1958:270) found the
territories of several pairs of Bell Vireos at the University of Kansas
Natural History Reservation to vary from .4 to 1 acre. Hensley
(1950:243) estimated the size of the territory of a pair of Bell
Vireos observed in Piatt County, Illinois, at 3.1 acres. Nolan
(1960:227) records home ranges of 2 to 3 acres. The pairs that
he studied were sole occupants of fields several acres larger than
the portions actually utilized. His description of the vegetation
indicates that most of the second growth was not much taller than
7 feet. As indicated elsewhere, the second-growth in my tract aver-
aged 15 feet tall. The smaller average size of territory (1.25 acres)
that I found is probably a function both of this greater vertical
range of available foraging area and the much higher gross density
of birds (40 pairs per 100 acres).
260 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hisr.
Permanence of Territories
Most pairs remain in their original territories throughout the
summer, although some shift certain territorial boundaries. In
1960 pairs 2 and 6, in the course of selecting a site for a replace-
ment nest, annexed adjacent areas previously occupied by other
pairs. Pair 2 relocated in a space that originally included territories
1 and 4, and pair 6 built a nest in an area formerly occupied by
pair 7. Males 1 and 4 were sacrificed for specimens and pair 7
probably was destroyed by a predator. Owing to the presence of
TABLE 2. SIZE OF THE NINE ORIGINAL TERRITORIES OCCUPIED IN 1960.
Date first
Territory occupied Dimensions
1 Dy ies Pe Oa NY le I ote Cae ae May 3, 1960 1.6 acres
DEPOT PIRES are REDE ERNE i om oh clatat s May 5, 1960 .6 acre
5 tl eS oe nop a ah RK ie A oh ed een May 7, 1960 .26 acre
BT ig Ny PN ois Sky a Ee eS May 11, 1960 1.03 acres
| ee Ale ete PE A (tks ER eeand(BAD oar te May 12, 1960 2.07 acres
Gates aw cae beech Capone one May 14, 1960 3.1 acres
EAC es sR TA Boh a a oe ik May 13, 1960 1.7 acres
ic Mein MULIURY, ANE SESE RIE Me SP e rere tte Rao Adore May 14, 1960 .46 acre
(2 RMON ee ste Un SAS ida 8. Res RE oe RE May 14, 1960 .4 acre
Average 1.25 acres
a nest, the annexed area becomes the focal point of the activities
of a pair, but the original area is regularly visited and may be re-
turned to in a later renesting.
Maintenance of Territory
Except in the early stages of nesting, territory is maintained
primarily by song. In the period of incubation a male regularly
patrols his territory between sessions of sitting on the eggs. He
sings several songs from each of several perches. A male follows
a predictable path, rarely traveling more than 150 feet from the
nest. Incipient patrolling is seen early in the breeding season
when territorial boundaries are in a state of flux.
The male White-eyed Vireo travels a semi-predictable route, as
does the Solitary Vireo (R. F. Johnston, MS). According to Law-
rence (1953:50), the male Red-eyed Vireo has a distinct singing
area completely divorced from the nest area dominated by the fe-
male. Southern (1958:109), working with this same species in
Michigan, did not recognize separate areas, but found that the male
wandered randomly over the territory.
NATURAL HistoRY OF THE BELL VIREO 261
In a species so highly active as the Bell Vireo, the degrees of
hostile action associated with an encounter overlap in such a
fashion that no clearcut distinction can be drawn among the various
displays. Nevertheless, certain generalized patterns are characteris-
tic of all situations in which members of this species are in a state
of anxiety. The threat displays described in the succeeding para-
graphs may all be utilized within as little as two minutes; mutual
agonism may be terminated at any stage by concerted attack of the
dominant bird.
1. Vocal threat. When an intruder is discovered the resident
male markedly increases his rate of singing. The alarm note, eh-
eH-EH, is the first call uttered during the nestbuilding and egg-
laying periods.
2. Head-forward threat. If the intruder does not flee, the resi-
dent male adopts a specific threat posture. The head and neck
are extended. The feathers of the crown are erected, but those of
the body are sleeked. The bird crouches slightly and the tail is
flicked laterally, but not fanned. The intensity of the singing in-
creases and is supplemented by scolding, also delivered at a rapid
rate. The intruder normally retreats at this juncture.
3. Wing-flicking and submaximal tail-fanning. If the interloper
remains, the anxiety of the resident male increases, He slightly
depresses the tail and, at the same time, rapidly fans and closes it.
The tail is only partially fanned. The wings are held slightly away
from the body and rapidly flicked above the back. This flicking
should not be confused with quivering of the wings associated with
begging and other solicitory postures. Song is now almost com-
pletely replaced by high-intensity scolding. Associated with this
high degree of anxiety are displacement behaviorisms, including
bill-wiping, reversal of direction on a single perch, and a nervous
hopping from one perch to another.
4. Ruffing and maximum tail-fanning. This display is most
often seen in conjunction with the harassment of predators, but
occasionally it is observed in territorial disputes occurring at the
boundary of adjacent territories where neither male is strictly
dominant and in which there is much vacillation prior to attack.
The feathers of the abdomen are ruffled. The term “ruffled” per-
tains to a full erection of the feathers, giving a ragged appearance
to the body outline (Morris, 1956:80). Ruffling of the abdominal
feathers emphasizes their yellow color and seemingly heightens
the intimidatory effect. The tail is fully fanned, and so maintained,
262 UNIVERSITY OF KANSAS PUBLS., Mus. Nar. Hist.
for a few seconds at a time; it is held at a 45° angle to the body.
The scold becomes an extremely intense, stacatto buzz, ZZ-ZZ-
ZZ-ZZ.
5. Supplanting attack. The attack directed against a trespassing
male is initiated as a lunge that results in a collision with the op-
ponent in mid-air or on his perch. The bird attacked is struck by
his adversary’s open beak or body.
Hinde (1952:71-72) indicates four courses of action followed by a
Great Tit (Parus major) when attacked under similar circum-
stances. “(a) It flies away: The attacker usually flies after it and
a chase ensues. (b) It shifts its perch a few inches: the attacker
lands in its place, and both usually show head-up postures. (c)
It remains where it is, but adopts a head-up posture: the attacker
usually then shows upright flight. (d) It may fly up and meet
the attacker in mid-air: in that case an actual combat may result,
or both combatants may show upright flight.”
Head-up posturing and upright flight are not presently recognized
components of the behavior of the Bell Vireo. The behavior of the
attacked Bell Vireo is similar to that described in (a), (b), and
(d) above, and is clearly dictated by the proximity of his own
“home base.”
Eleven disputes among occupants of adjacent territories were
witnessed between May 6 and June 3, 1960, in which some or all
of the described threat displays were manifest (Table 3). In each
instance, patrolling males were gradually attracted to each other.
As they approached, their rates of song increased from an average of
six repetitions per minute to 15 per minute. Eight of the disputes
involved physical combat.
On May 6, 1960, when male 2 (1960) was in the process of usurp-
ing an eastern segment of the original territory of male 1 (1960),
a violent, protracted dispute was observed. By this date male 1
(1960) had obtained a mate and had begun construction of nest
l-a (1960); male 2 (1960) had not yet acquired a mate. At first
the two males were singing vigorously, from one to 10 feet apart.
Female 1 (1960) followed her mate closely and scolded, at the same
time partially fanning her tail. In the course of vocal dueling the
males had traveled to within 50 feet of nest 1-a (1960), when male 1
(1960) suddenly lunged at 2 (1960). The males plunged to the
ground, locking bills and clutching at each other with their feet as
they fell. As soon as they touched the ground they separated.
NATURAL Hisrory OF THE BELL VIREO 263
Male 2 flew east with male 1 in pursuit. This conflict lasted three
minutes.
Additional physical combat was witnessed several minutes later.
This again involved striking with the bill, wings and feet. A high
pitched squeaky chee was uttered by both combatants. The female
scolded from a nearby perch. Upon separating, the males engaged
in a wild, looping flight. At about 350 feet from nest l-a (1960),
the chase abruptly ended. For ten minutes thereafter, both males
sang at a high rate from perches about 10 feet apart. This termi-
nated the physical combat, but three additional protracted, vocal
duels occurred in the remainder of the morning.
TABLE 8. INTRASPECIFIC DISPUTES IN MAINTENANCE OF TERRITORY.
Behavior
Number Average
of ee Combat length of
conflicts 8 disputes
Prenesting ssh. oes; 3 3 2 6 min. 40 sec.
Bilding eee. ae 8 8 6 3min. 8 sec.
HMCUDAtIONS cuties cy. oe 1* 1 aN ares Te 20 min
Ota Sf. oytiscie Ree 12 12 8 5 min. 30 sec.
* Directed against a stuffed Bell Vireo.
Probably as a direct result of these conflicts, a neutral zone ap-
proximately 300 feet wide developed between the two territories.
By May 14 this intervening area was occupied by male 4 (1960).
By this date both 1 (1960) and 2 (1960) were involved in nest-
building and 4 (1960) was not challenged for several days.
Maximum tail-fanning prior to attack also appears as an element
of aggressive behavior in White-eyed Vireos. A brief skirmish be-
tween a male of this species and a small, greenish passerine was
observed at the Natural History Reservation on May 25, 1960. The
White-eyed Vireo was singing from a perch 30 feet high in a dead
elm, when the unidentified passerine landed 10 feet distant. The
white-eye ceased regular song and uttered several catbirdlike calls,
and at the same time slightly depressed and fully fanned the tail.
After 10 seconds, the white-eye lunged at the intruder, striking it in
mid-air. A brief looping flight ensued through the branches of the
elm before the intruder was able effectively to retreat.
264 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
Aggressive Behavior of the Female
The female Bell Vireo is concerned primarily with the defense of
the nest and the young and she rarely assists the male in defense
of distant parts of the territory. She employs the same threat dis-
plays as the male.
Interspecific Relationships
A number of meetings between Bell Vireos and other species were
observed in the course of the study (Table 4). Resident pairs of
this species exhibited different degrees of tolerance toward other
TABLE 4. INTERSPECIFIC CONFLICT OBSERVED IN 1959 anp 1960.
Numb ee Bee Behavior of Bell Vireos
Species re breeding
POURS eycle HFT*| S| TF | A
Coccyzus americanus. . 1 Nestling period....}......] x
Cyanocitia cristata. ... 3t Nestling and incu-
bation period..| x x x x
Parus atricapillus... . | Prenesting #5 occ eden x
Molothrus ater....... 1 Nestling period....]...... Xv eeee x
Dendroica petechia... . 1 iPrenesting feck + ocak ees Sih ecg Pic x
Geothlypis trichas..... 1 Nestbuilding......}...... Ku fawn x
Pituophis catenifert... 1 Post-Hedging 222 aloes ban [eee x
* HFT = head-forward threat; S = scolding; TF = tail-fanning; A = attack.
+ Includes attack against a dummy Blue Jay.
syne Bull Snake is here included because the vireos directed typical aggressive displays
towards it.
species. Many birds, including Cardinals, Field Sparrows, Painted
Buntings and Mourning Doves were ignored completely. Chicka-
dees evoked responses characterized by slight increase in song and
some anxiety; this was perhaps owing to similarity in size, motion
and call notes. Warblers, when met with, were invariably chased.
They may be momentarily mistaken for rival vireos.
Blue Jays were vigorously attacked, especially late in incubation
and throughout the nestling period of the Bell Vireo. I did not see
a jay struck, but a vireo would circle one closely as it perched and
pursue it when it flew, following as far as 100 yards beyond terri-
torial bounds. The buzz, ZZ-ZZ-ZZ-ZZ, was uttered in conjunction
with this harassment.
A stuffed jay placed eight feet from a nest elicited threat display
and displacement behavior from the owners of the nest, but no
NATURAL HIsTORY OF THE BELL VIREO 265
attack. Incubation had just begun at this nest. A dummy Bell Vireo
placed close to another nest only momentarily disturbed the male,
and the female completely ignored it. Incubation had also recently
begun at this nest. At this same general stage, moreover, nesting
pairs showed little inclination to harass me.
Discussion
Hinde (1956:341-342) indicates that territory has been defined
in a number of ways by many workers. All of the definitions involve
modification of Howard’s classic “defended area.” Pitelka (1959:
258) has reacted against this behaviorally-oriented concept. He
thinks that the concept of territory should be based on exclusive
use of an area by its occupants, and not so much the defense by
which they maintain it.
Methods of treating territoriality in the Bell Vireo seemingly
incorporate features of both schools of thought. The area used
exclusively for all biological needs by a single pair of Bell Vireos
is vigorously defended both physically and vocally early in the
breeding season and vocally as the season progresses.
In the period of territorial establishment a relatively large area
is actively defended. The building of a nest establishes a focal point
of activity in a somewhat more restricted area than that originally
occupied. After the success or failure of a nest, a new site is selected
to which the focal point of activity is shifted. If suitable habitat
adjacent to the extant territory is unoccupied by other Bell Vireos
the unoccupied area may be annexed in the course of searching for
a new site. Such annexation occurs only when pairs formerly oc-
cupying adjacent suitable habitat disappear from this territory;
possibly the size of the territory of any one pair is dictated by the
density of population of the species as well as by the presence of
suitable habitat. This may not always be true as indicated by
Kliujver (1951:40), who in studying the Great Tit, found no appre-
ciable difference in the size of territory in two different habitats
even though there was a marked difference in population density
of the birds.
Fluctuation of territorial boundaries is not uncommon in passer-
ines, especially when no rivals exist to contest movement. Hinde
(1956:351) indicates that fluctuations in size of territory are to be
expected although the territories of different species of birds have
different mean sizes.
Once nesting activities commence there is a marked reduction in
266 UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist.
the amount of territory utilized and a distinct decrease in the
aggressive tendencies of the male; it would seem that energy pre-
viously utilized in regular fighting is rechanneled for nestbuilding,
incubation and care of the young. Further, contraction of the area
of activity obviates high-intensity territorial defense, as adjacent
males, even in regions of high population density, are isolated from
one another by an area no longer regularly traversed.
With cessation of breeding activities physiological mechanisms
governing maintenance of territory seemingly are no longer active
and yet the pairs of Bell Vireos remain within a restricted area which
they alone use. Earlier definitions of territory as a “defended area”
do not adequately cover such situations and yet from the standpoint
of Pitelka the area still retains the characteristics of true territory.
In fact, territory as defined by Pitelka is clearly manifest at this
time. Whether the birds remain in an area through “force of habit”
is of little consequence.
I have retained the term “territory” in preference to the term
“home range” used by Nolan (1960:227). His failure to observe
territorial defense is responsible for his terminology, although it is
readily understandable that such defense would be lacking in a
population of relatively low density in which pairs were isolated
from one another by areas of unfavorable habitat. This isolation in
itself would tend to preclude territorial conflict but territories were,
in fact, maintained.
The marked similarity in the essential features of aggressive
behavior in North American vireos attests to their close relationship.
Flicking and fanning of the tail are distinct components of the hostile
behavior of the Bell Vireo, White-eyed Vireo, Red-eyed Vireo (Law-
rence, 1953:69), and the Black-whiskered Vireo (Vireo altiloquus;
Bent, 1950:319), and, presumably, of the remaining species of the
genus. The occurrence of these same displays as intrinsic behavioral
elements of interspecific hostility suggests a common derivation.
Moynihan (1955:256) indicates that all intraspecific hostile displays,
and probably most interspecific hostile displays, evolved originally
as social signals having the same general function. Further, Hinde
(1956:344) points out that there is a fundamental similarity in the
motor patterns used in fighting in different contexts, including both
interspecific and intraspecific fighting.
NATURAL HIstToRY OF THE BELL VIREO 267
COURTSHIP BEHAVIOR
The precise mechanism of pair-formation in the Bell Vireo is not
known. My experience has been to find a male one day and then
one or two days later to discover that it has a mate. Lawrence
(1953:53), tells of a male Red-eyed Vireo singling out a female
from a flock of migrants passing through his territory and violently
driving her to the ground. Shortly after this attack the pair was
seen searching for a nest site. But such an incident has not been
reported for other vireos, nor have J witnessed such behavior myself.
Early courtship activities of the Bell Vireo are characteristically
violent affairs, with the male directing strong aggressive attacks
toward the female. Rapid, looping flights through the thickets
occur, the female leading the male. Occasionally he deliberately
collides with her in mid-air, but the pair quickly separate. This
violent sexual chasing is manifest prior to the inception of nestbuild-
ing. With commencement of this activity, sexual chases through
the territory subside.
Absence of sexual dimorphism in the Bell Vireo obviously suggests
that behavioral criteria are used by the birds in sex-recognition. The
lack of aggression by the female upon initial aggression by the male
is an essential component of recognition of sex; she is clearly sub-
ordinate. Such subordination is also the significant feature of
continued sex-recognition. Courtship display by a resident male,
directed toward a stuffed male and a wounded male which sat
motionless, supports the contention that a subordinate or submissive
attitude of the female is a key factor in sex-determination.
Nestbuilding and courtship are intimately associated in this
species. The male constructs the suspension apparatus of the nest,
the completion of which coincides with the assumption of nestbuild-
ing activity by the female. Roles of the sexes in nestbuilding are
described in the section on nestbuilding. The male frequently in-
terrupts construction to court the female. This, in combination with
perpetual song as he works, serves to strengthen the pair-bond and
stimulate nestbuilding tendencies of the female.
It is doubtful that any attempts at copulation are successful up
to this time. The female is singularly unresponsive to the advances
of the male; a female retreats before most violent attacks and is
seemingly oblivious to less vigorous behavior. After the female
268 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
assumes the responsibility of building, the tempo of courtship
activities increases.
The female becomes increasingly more receptive and her work is
often interrupted by advances of the male. Copulation occurs fre-
quently from about the third day of nestbuilding through the first
day of egglaying, a period of four to six days. Male displays and
vocalizations associated with courtship continue through the fourth
or fifth day of incubation.
Displays and Postures
The principal courtship displays and postures that were seen
throughout the nestbuilding phase are as follows:
1. Greeting ceremonies. Both birds are crouched from one to five
inches apart. The feathers on one (the male?) are sleeked, and on
the other are fluffed. Fluffing (Morris, 1956:80) denotes partial
erection of the body feathers producing a rounded, unbroken body
line and is not to be confused with ruffling, mentioned in the sections
pertaining to territoriality and pre- and post-copulatory display.
Fluffing is generally considered to be an appeasement display and
it is seen in a variety of situations involving a dominant-subordinate
relationship. Both birds flick wings and tails rapidly and reverse
directions on their perches frequently. A low, rapid chee is uttered
during this performance. This ceremony is repeated often in the
first three days of nestbuilding, but less frequently thereafter. It
usually occurs after building by one or both partners and prior to
another trip in search of nesting material. It lasts from 10 to 50
seconds and is not immediately followed by any additional courtship
activities. Nolan (1960:228-229) observed mutual displays between
periods of violent sexual chase that suggest that the greeting cere-
monies that I have described are an integral part of pair-formation
as well as a component of continued maintenance of the bond.
2. “Pouncing.” The female rapidly quarter-fans and partially
depresses her tail. She utters a high pitched scold (chee). The
male, from a perch within two feet of the female, fans the tail fully
and depresses it vertically, and, with mouth open, lunges at the
female; or, with similar tai] mannerisms, the abdominal feathers
ruffled, the wings held horizontally, and the primaries spread, he
sways from side to side from four to six times, and then lunges at
the female. The male is silent when he pounces; the chee or the
courtship song is emitted when swaying precedes pouncing. The
male strikes the female with his breast or with his open beak. The
female rarely flees although she is usually displaced several inches
Narurau History OF THE BELL VIREO 269
along the branch upon which she is sitting. However, the female
may fly several inches to a new perch. The failure of the female to
adopt a solicitation posture presumably indicates sexual unreadi-
ness. Instances of the male deliberately colliding with the female
as she flies in the course of gathering nesting material are probably
analogous to pouncing. In none of the above situations are females
observed to fight back in any way. Nice (1943:174) believed pounc-
ing to be analogous to sexual chasing found in such species as the
Red-winged Blackbird. In the Song Sparrow, pouncing is observed
most often in the first and second days of nestbuilding.
3. “Leap-flutter.” The male, in the course of displaying with the
tail fanned before the female, suddenly leaps eight inches to ten
inches vertically and flutters in mid-air several seconds, before drop-
ping to the original perch. This display occurs in full view of the
female. It is often associated with pouncing and is also seen prior
to copulation. In the latter instance it is probably pragmatically
functional, for it permits the male to orient above the female before
dropping to her back to copulate. No vocalization is uttered during
the leap-flutter.
4, Pre-copulatory display (Fig. 3). The male faces the female.
The tail is fanned fully and depressed at a sharp vertical angle to
the body. Body feathers, both dorsal and ventral, are ruffled, almost
tripling the apparent volume of the thorax. The head is withdrawn
and slightly thrown back. Feathers of the head are not erected.
Fic. 8. A single male Bell Vireo in the pre-copulatory
display. Note the ruffled dorsal and ventral body feath-
ers. The male on the left has reached the zenith of a
single swing. The male on the right has nearly reached
the low point of a swing.
270 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
The mouth is opened wide. The legs are slightly flexed and the
body is swayed laterally. Horizontally, the head and body traverse
an arc of about 100°; vertically, they traverse an arc slightly less
than 180°. At the low point of any one swing, the delivery of the
courtship song begins. At the termination of the swing the two
normal, ascending notes are emitted. This performance may last
as long as three minutes.
The pre-copulatory display of the male elicits receptive behavior
in the female. She crouches in a solicitous manner, with the body
feathers fluffed and the tail raised slightly, and utters a muted chee.
5. Copulation. The male abruptly terminates his swaying display
with a leap-flutter that positions him above the female’s back. He
then descends and copulation occurs. The male continues to flutter
his wings to maintain balance throughout the two seconds of cloacal
contact. Following an unsuccessful copulation on June 23, 1960,
displacement preening and bill wiping were performed by both
sexes.
6. Post-copulatory display. On June 25, 1960, after a second
attempt at copulation with a stuffed bird in which semen was
actually deposited on the dummy’s back, male 10 (1960) performed
a swaying display. In this instance, however, instead of addressing
the dummy from the front, the male alighted one inch to the right
of the stuffed bird. When swaying to the left (toward the dummy)
the head of the displaying male actually passed above the neck of
the stuffed bird. This ritualized behavior could conceivably be
derived from hetero-preening.
Discussion
Within the scope of my research it was difficult to detect the
over-all sequence of epigamic displays that result in synchronization
of the physiological states of the sexes throughout the period of
courtship. Possibly all displays, except the post-copulatory one,
occur in no particular order in the courtship period. However, each
ritualized display seemingly strengthens the pair-bond.
Swaying has been recorded in a variety of situations of a sexual
and semi-sexual nature for the Solitary Vireo (V. solitarius; Town-
send, 1920:158) and the Red-eyed Vireo (Tyler, 1912:280; Bent,
1950:342). In every instance the body feathers of the swaying
birds were sleeked. Courtship behavior in any species of North
American vireo seems closely to resemble that of any other; pairing
NATURAL HIsToRY OF THE BELL VIREO oral
and nestbuilding of a female V. solitarius and a male V. flavifrons
as reported by Hauser (1959:383) support the idea of close re-
semblance.
A marked similarity will be detected between certain basic ele-
ments of aggressive and epigamic displays. These basic elements
are wing- and tail-flicking, tail-fanning, and high-intensity delivery
of the chee. Pouncing and supplanting attacks are essentially simi-
lar. Such similarities suggest either a common origin for certain
aggressive and epigamic displays or the derivation of one from the
other.
High-intensity cheeing is obviously a function of excitement,
whether in conjunction with hostility or sexual behavior. According
to Andrew (1956:179), flicking of wing and tail in passerines are
intention movements of flight. These actions have been emanci-
pated from incomplete take-offs and incorporated in ritualized court-
ship and agonistic behavior. In incipient courtship behavior the
male is governed by three conflicting tendencies; to flee, to attack,
or to behave sexually before his mate (Tinbergen and Hinde, 1958:
256). When pairing, Bell Vireos interrupt sexual chase with
“greeting ceremonies,” the male’s tendency to attack and the fe-
male’s tendency to flee are momentarily reduced, and the forming
bond is strengthened. Thus, the intention movements become an
integral part of courtship.
In situations where attacking and fleeing are the two conflicting
tendencies, wing-flicking and tail-flicking are incorporated into
threat display, but do not lose all of their original function, for
they facilitate attack. Tail-fanning, as a display element, increases
the awesome aspect of the threatening bird and in courtship pre-
sumably makes the sexes more attractive to one another.
Courtship feeding has not been recorded for the Bell Vireo. In
general, it is unknown in North American vireos, with the excep-
tion of the red-eye (Lawrence, 1953:53). It would serve no “prac-
tical” purpose in the Bell Vireo since the male regularly relieves the
female during incubation, thus allowing her ample opportunity to
forage. In the Red-eyed Vireo, only the female regularly incu-
bates, and courtship feeding is definitely functional. Nolan (1960:
228) described a brief pecking or pulling with their bills between
pairing birds. This may be incipient “symbolic” courtship feeding,
or perhaps mutual preening.
272 Universtry OF Kansas Pusts., Mus. Nat. Hist.
SELECTION OF NEST-SITE AND NESTBUILDING
As far as can be determined, the nest-site is selected by the
female, Typically, the pair makes short, low-level flights from tree
to tree with the female invariably in the lead. The birds usually
forage within each tree; the female interrupts this activity to inspect
small forks of low, pendant branches and the male occasionally
pauses to sing. The singing is loud but not particularly regular,
as it is later when the male accompanies the female during actual
nestbuilding. Method of selection of site resembles that described
by Lawrence (1953:53) for the Red-eyed Vireo.
Nests are suspended from lateral or terminal forks about 27 inches
high in bushes and small trees that, in the study area, averaged
11 feet, four inches in height (Table 5). The height above ground
of the nests does not vary appreciably as the season progresses as
is the case with nests of Red-eyed Vireos, for which Lawrence
(1953:54) noted that late nests were placed higher than those
built earlier in the season.
Most nests are so situated that they are protected and concealed
by the dense foliage of trees. Where nests are placed in low bushes,
as coralberry or dogwood, the bush is invariably overhung by the
foliage of a much taller shrub or tree.
The nest tree or shrub was in every instance situated at the edge
of a thicket or isolated from adjacent trees by several feet. Pref-
erence for open situations is characteristic of the species. In con-
trast, the nest of the White-eyed Vireo (Bent, 1950:229) is placed
toward the center of thickets.
In the choice of sites in the study area, the Bell Vireos were
almost unopposed by other avian species, owing to the size of the
TABLE 5. Nest-stres UTILIZED In 1960.
Number Average Average
Plant of height of height of
nests plant nest
|
Ulmus americana... .......-. 4 Git Gin.s-: 2 ft. 3 in.
Maclura pomifera........... 20 | 13 ft 11 in......| 1 fe. 11 in.
Crataegus mollis... 2.0... 0... 1 1 ya ey ee tae 3 ft. lin.
Glediisia triacanthos.......... 2 15 G- On. ee et
Ater'neyuntdo. ¢ Ps SSS SS 4 S'toSin ts = 2 ft. 5 in.
Cornus drummondi.......... 2 SE cpscn. SSE 2 ft. 8 in.
Symphoricarpos orbiculatus... .| 3 = St ee! 1 ft. 10 in.
|
Ie woe we si Sek Gite Cee 36 11 ft. 4im.......] 2 ft. 3 in.
NATuRAL History OF THE BELL VirnEO 273
fork utilized and the fact that the nests are located peripherally,
rather than centrally, in the bush or tree. This lack of competition
for a nest-site provides a Bell Vireo with an ample supply of nest-
sites within any one territory.
Selection of the first nest-site may take as long as two days,
possibly owing to incomplete development of the nesting tendency,
but more likely to a general lack of familiarity with the territory.
Red-eyed Vireos require five to six days to choose the first nest-site
(Lawrence, 1953:54). Later sites of the Bell Vireo are chosen in
as little as three hours. Nest l-c (1960) was abandoned at about
11:00 a.m, on May 14, 1960, when part of the thicket on the edge
of which this nest was located was removed by brush-cutters clear-
ing a power line right-of-way. By 2:00 p.m. this pair had begun
construction of J-d (1960) in an Osage orange 110 feet southwest of
l-c (1960).
This particular site is of further interest because it is the same
one utilized for nest l-a (1960). In all, four instances of utiliza-
tion of a nest-site a second time were recorded. Two-a (1960) and
2-d (1960) were built in the same fork; 1-c (1960) and 1-h (1960)
were in the same tree, but not the same fork. It should be men-
tioned that l-a (1960) and 2-a (1960) were abortive attempts that
did not progress beyond the suspension apparatus. Nice (1929:16)
recorded a similar instance of the re-use of a nest tree, but different
forks were used.
Re-use of an exact nest-site would ordinarily be impossible if
the initial attempt were not abortive, because the presence of a
completed nest would pose problems in construction with which
the birds would probably be unable to cope. (A report by Morse
in Bent, 1950:256 of a double nest indicates that this may not al-
ways be true. At the time of discovery one nest contained two eggs
and the other nest contained young.) Since nests are used only
once there would be no tendency to adopt the old nest. However,
abortive nests, usually little more than a few strands of nesting
material secured to the fork, might stimulate the birds to continue
building. Re-use of a single nest-site in 15.8 per cent of 38 nests
built in 1960 seems to be more than fortuitous circumstance. This
re-use may have physiological benefits in conjunction with appor-
tionment of energy for other nesting activities, because rapid loca-
tion of a nest-site would mean that energy normally expended in
searching and selecting could be rechanneled for actual construc-
tion. In each of the instances of rebuilding, the new nest was
3—1506
274 Untversiry OF Kansas Pusts., Mus. Nat. Hist.
begun on the same day that the previous nest was abandoned.
The re-nesting of pair 9 (1960) is worthy of note. These birds
were established in the elm thicket on Clark land. Elm was by
far the most abundant tree, with dogwood, Osage orange and honey
locust also relatively common, There were only six boxelders in
the territory and yet the four nests built by this pair were placed
in them. This is the only instance of seeming preference.
Building
Nestbuilding by Bell Vireos can be best discussed in terms of
the phases of construction described for the Red-eyed Vireo, Law-
rence (1953:57), which are: (1) construction of the suspension
apparatus, (2) construction of the bag, (3) lining of the bag and
smoothing and polishing of the exterior, and (4) adornment of the
exterior. Red-eyes (Lawrence, 1953:59) may continue adornment
far into the period of incubation. Both the male and female Bell
Vireo have been observed to add spider egg sacs and other silk
to the exterior of the nest as late as the sixth day of incubation.
Nice (1929:16) recorded only the female Bell Vireo building,
but she did recall, from previous studies, having seen males aiding
somewhat. Pitelka and Koestner (1942:102) wrongly concluded
that the female Bell Vireo builds unaided, but Hensley (1950:243)
observed that both sexes participated in nestbuilding, and Mum-
ford (1952:229) reported two instances of building by both adults.
His description of the activities viewed in mid-May suggest that
they were of the transitional period between the first and second
phases. On the second occasion he recorded both adults building
during the second phase. Since no details accompany this second
observation I assume that it pertained to activity not necessarily
typical of this phase of construction. Whereas both sexes of the
Bell Vireo cooperate in building the nest, only the female Red-
eyed Vireo builds according to Lawrence (1953:56). But Common
(1934:242) saw both Red-eyed Vireos building a nest.
The suspension apparatus is constructed by only the male on the
first day. He punctuates each trip to the nest with song. The single
song phrase is given from three to eight times when the male, carry-
ing nesting material in his bill, arrives in the tree. Typically, he
alights on several perches within the nest tree before flying to the
nest. He often interrupts his work with several songs; when he
has finished adding a load of material he sings from several perches
NATURAL HisTorRY OF THE BELL VIREO 275
within the nest tree before departing. The male periodically stops
building to court the female.
In eight hours (494 minutes) of observing the first phase of con-
struction at five different nests, I saw the female come to the nest
28 times; the male made 95 trips. The female came alone only once,
and brought nesting material ten times, but did not build; on the
other 18 occasions her visits were brief and she usually confined her
activities to an inspection of the nest. Twenty of the visits by the
female were made late in the first phase, marking a gradual transi-
tion to her assumption of building responsibility. (The delay by
the female in beginning to build is puzzling; because all evidence
indicates that she helps select the nest-site, I would expect her to
help with the initial building. There seems to be no clear advantage
in her delay in beginning to build.) The courtship and building
activities of the male plus the presence of a partly completed nest
seem to stimulate the female to commence building. Her visits
become more frequent as construction of the suspension apparatus
nears completion. At a time early in the second day the transition
has taken place, and the female becomes the sole worker.
On May 7, 1960, male 2 (1960), at the time unmated, was ob-
served as he came upon a nest of the previous year. The nest, after
a year’s weathering, suggested in appearance perhaps an early
second-day nest. The bird flew to the nest and tugged and wove
loose strands of grass for three minutes. Before leaving the site, the
bird sang twice from different perches. This observation suggests
that a partly constructed nest can elicit nestbuilding behavior, even
in an unmated male.
The techniques of building by the male consist primarily of laying
pieces of grass or bark across the fork, or along one of its branches,
and then fastening them in place with pieces of animal silk. Once
a “racket” has been formed, spider egg cases and plant down are
emplaced among the fibers. The male employs weaving, twisting,
and pecking motions of the head to emplace material.
As previously indicated, the female is the principal worker in the
second and third phases of construction. The male infrequently
visits the nest, but regularly visits the nest tree. The molding of the
bag is accomplished by piling leaves, grasses and plant down onto
the suspension apparatus. This material is also bound in with animal
silk. As the amount of material accumulates, the female begins to
trample it and gradually the bag takes shape. When trampling is
276 UNIvERSITY OF Kansas Pusts., Mus. Nat. Hist.
first attempted, the nest often fails to support the female and she
falls through the bottom of the nest. Such an occurrence was ob-
served on May 23, 1960, on three consecutive trips by female 1
(1960), in constructing nest l-e (1960). As the bag deepens, addi-
tional strands of grass are added to the wall and woven into place.
The male is extremely attentive during this and the following
phase. He follows the female as she gathers nest-material ac-
companying both this activity and her building with rapid song;
he may give an average of seven song phrases per minute. The
male brings to the nest a strand of grass, or some other material,
about every twentieth trip. He frequently inspects the nest and
the activities of the female from perches near the nest. Con-
struction of the bag is ordinarily completed in the third day.
The third phase, the lining of the interior and the smoothing
of the exterior, involves an additional one and one-half to two days.
Smoothing of the exterior refers to tightening of the grasses woven
into the bag and addition of more animal silk. In lining the nest,
the female stands on one of the branches of the fork and emplaces
one end of a long, thin strand of some relatively stiff piece of
grass or strip of bark. She then jumps into the bag and, while slowly
turning around, pecks it into place, thus coiling the strand neatly
around the interior of the bag.
As previously mentioned, the fourth phase overlaps the periods
of lining, smoothing, egglaying, and incubation. The principal
activity is the addition of white spider egg sacs to the exterior.
The trips are infrequent; but, occasionally, birds will interrupt
an hour of incubation with three or four minutes of active adorn-
ment, during which several trips may be made. Both sexes par-
ticipate in this phase.
Gathering of Nesting Material
Nesting materials were gathered anywhere within the territory.
Occasionally materials were collected from within the nest tree,
but usually they were obtained 20 to 200 feet from the nest-site.
On several occasions I observed birds inspecting stems or branches
where bark was frayed. Loose ends are grasped in the beak and
torn free with an upward jerk of the head. Possibly the notch near
the distal end of the upper mandible aids in grasping these strands,
plant down is first extracted and then rolled into a ball by means
of the beak while held with the feet before being transported to
the nest.
NATuRAL History OF THE BELL VIREO 277
Length and Hours of Nestbuilding
As indicated by Nolan (1960:230), accurate determination of
the length of nestbuilding is difficult because of continued adorn-
ment and polishing after the nest is functionally complete. Most
of the early nests for which I have records took from four and one-
half to five days to construct. A four- to five-day period of building
is reported by other observers (Nice, 1929:16; Pitelka and Koestner,
1942:99; Hensley, 1950:242; Nolan, 1960:230).
One instance of protracted building was recorded. Nest 6-d
(1960) was begun on May 29, 1960, and not completed until nine
days later on June 6, 1960. In contrast nest 1-g (1960) begun on
May 31, 1960, was finished three days later on June 2, 1960. Nest-
building occurs between the hours of 6:00 a.m, and 5:30 p.m.
Heavy rain in the early morning may delay building.
Abortive Nestbuilding Efforts
Eight of 38 nests started in 1960 were never completed (Table 6).
Six of these abortive attempts were abandoned during, or shortly
after, the completion of the suspension apparatus. Five of these
nests were abandoned because the female did not begin building
following the end of work by the male. The early abandonment of
the other three nests l-a (1960), 2-c (1960) and 6-e (1960) was
attributable to the interruption of building by the male because of
heavy rain and protracted territorial conflicts. The occurrence of
these abortive nests at any time within the nesting efforts of a single
pair indicates that such attempts are not examples of “false nest-
building.”
Renesting
Renesting after desertion or successful fledging occurs within two
to thirty-six hours. Young were fledged from l-a (1959) on June
19, 1959, and nest 1-b (1959) was discovered when late in the
second phase of construction on June 22. If the nest was started on
June 20, then renesting took place within 15 hours after fledging.
The Nest
Several authors have described various aspects of the nest of the
Bell Vireo, notably Goss (1891:535); Simmons (in Bent, 1950:256),
Nice (1929:13) and Nolan (1960:230-231). I can add but little to
these descriptions.
The nest itself is a compact structure composed of strips of bark
and strands of grasses that are interwoven and tightly bound with
278 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
TABLE 6. ABORTIVE NESTING ATTEMPTS IN MAY AND JUNE OF 1960.
Length
Nest of time Cause of abandonment
worked on
Lae we caine ee ee oe 1 day Heavy rain
MDa Sroqelin wae so. ality ie 2 days Female failed to build
Da cea elitist Stal stone eh ate caietekohd Coulee day Female failed to build
7 7 anata ec aa Paar ARP a 1 day Protracted territorial dispute
Pe eR CR a RRR CE ICR Aa 1 day Female failed to build
DB sca Un athe War dgsl earcio tense eterota oe 1 day Female failed to build
GHC a Perera ee aate, Sote raeke 1 day Heavy rain
Cations Cobden gece Wevotere sistigayl ieee 2 days Female failed to build
animal silk. The floor of the cup is first lined with a layer of small
leaves and then the entire interior is lined with fine stems or strips
of bark. Feathers are occasionally used to pad the bottom prior to
lining, as are pieces of wool and milkweed down. Nest 2-e (1960)
had been packed with small pieces of soil bearing moss prior to
lining.
Early nests tend to be bulkier, having thicker walls and bottoms
than later efforts. However, nests in May were found to have 16
per cent thicker bottoms and 41 per cent thicker walls than nests
in June (Table 7). Standard nest measurements do not show this
to be so, for the exterior and interior diameters at the rim are gov-
erned by the angle between the two branches of the fork.
TABLE 7. DIMENSIONS OF NEsTs IN May (1960) AND JUNE (1960).
Measurements May (N 10) June (N 8)
EGXCernalpdenUns ois tiers cists tsos oye sie 0% wie 61.6 mm. 59.3 mm.
Depthiol(eup ss. 2 os teres eels aks eee 45.5 mm. 46.3 mm.
Qutside;diameter. = wa.2; srs. Sess bide, 57.3/55.5 mm. | 54.3/53.5 mm.
Insideidiametens se wee ei eae 43.4/42.2 mm. | 45.5/43.9 mm.
Thickness of forward wall 1 inch below rim | 13.8 mm. 7.6 mm.
‘Dhicknessiofibottomi: ast ene. ties cee. 11.3 mm. 4.6 mm.
EGGLAYING AND INCUBATION
Egglaying
Egglaying begins the first or second day after completion of
the nest. The female sits in the nest occasionally for periods of
five to twenty-five minutes on the day the nest is completed. This
is interrupted by periods of nest-adornment and foraging; such ac-
tivities sometimes keep the female off the nest for several hours.
Prior to the laying of the first egg, only the female is seen on the
NATURAL History OF THE BELL VIREO 279
nest, although the male is often seen sitting quietly within the nest
tree a few feet from the female. The infrequency of the “con-
gested” song and the alarm (eh-eH-EH) after the inception of
“broodiness” indicates the waning of courtship behavior. As later
in incubation only the “normal” song and the scold are heard.
Eggs are laid early in the morning prior to 5:30 a. m. according
to Nolan (1960:232). The nest is usually left unoccupied for con-
siderable periods after the first egg is laid, but, on the first day of
laying, both sexes have been observed sitting for brief periods aver-
aging ten minutes in length. Eggs are laid at one-day intervals
until completion of the clutch. I found incubation to begin with
the second egg.
Clutch-size
The average clutch-size of the Bell Vireo in Kansas, based on
thirty-three records, is 3.39 eggs (Table 8). Seasonally, the largest
average clutches are produced in the middle of the breeding season,
that is, in June. Lack (1947:308-309) indicates that in European
TABLE 8. AVERAGE NumBERS OF Eccs PER Nest (NUMBER OF RECORDS IN
PARENTHESES ) *,
Mean
Year May June July annual
clutch-size
——————— | |
LUG a Al SP Ae Bs 320) (C7) 3.2 (12) 3.0 (1) 3.06
BOGOF at. 2 as ft S3- 3.3 (6) 3.83 (5) 4.0 (2) 3.72
BOBO — LOGO =... 55; oe teyoncs 3.17 3.52 3.5 3.39
* These data have been supplemented from the literature pertinent to Kansas.
passerines the highest seasonal average clutch-sizes likewise occur
in June. The largest average clutch-size in the Bell Vireo is pre-
sumably related to some aspect of the availability of food.
Caution is necessary in determining mean clutch-size in the Bell
Vireo. Eggs occasionally disappear from the nest prior to or dur-
ing incubation, without subsequent addition of cowbird eggs. Un-
familiarity with the history of such a nest on the part of the observer
would lead to an inaccurate determination of clutch-size.
Complete clutches are not replaced with the same regularity as
are nests. I have recorded intervals of six to thirty days between
successive clutches. Successful replacement of clutches is deter-
mined by a number of factors: nest-site, completion of a nest,
weather, predation, and parasitism by the cowbird. The difference
280 UNIVERSITY OF Kansas PuBsts., Mus. Nat. Hist.
between the number of renesting attempts and the successful re-
placement of clutches seems to indicate that different physiological
processes are responsible for these two phenomena and that there
is lack of synchrony between them. The development of the ovarian
follicle requires a specific number of days that is not always coinci-
dent with the building of replacement nests. If, in the Bell Vireo,
replacing a nest were solely a responsibility of the female, instead
of involving the male to a considerable extent, it would seem likely
that replacement of nests and the replacement of clutches would
be more closely coordinated.
Incubation
Nice (1954:173) considers the incubation period to be the elapsed
time between the laying of the last egg in a clutch and the hatching
of that egg, when all eggs hatch. My data indicate that, normally,
intensive incubation begins when the second egg is laid and lasts
fourteen days in the Bell Vireo. Nice (1929:99) also considered
the incubation period in this species to be fourteen days but be-
lieved it to commence when the third egg was laid. Pitelka and
Koestner (1942:99) noted that the first and second eggs hatched
fourteen days after laying of the second egg. However, they
thought incubation began with the first egg. This would mean a
fifteen-day period for this egg. All the eggs that Nolan (1960:234)
marked hatched in approximately fourteen days. Eight eggs arti-
ficially incubated by Graber (1955:103) required an average of
15.01 days to hatch. As Van Tyne and Berger (1959:298) indicate,
periods of sitting on the nest, even all night, do not necessarily mean
that incubation has begun, for it has been demonstrated in several
species that birds may sit on an egg without actually applying heat.
My own observations demonstrate that the first egg may be left un-
attended for several hours at a time on the day that it is laid.
The Roles of the Sexes in Incubation
Both the male and female sit on the eggs in the daytime. My
study of histological sections of ventral epidermis indicates that
the male does not possess a brood patch; the increased vasculariza-
tion typical of the brood patch in females is not evident in males.
But, the male loses most of the down feathers of the ventral ap-
terium. Also, according to Bailey (1952:128), the male Warbling
Vireo that sits on the eggs lacks a brood patch.
Bailey (1952:128) suggests that male passerines lacking brood
patches that habitually sit on eggs do not heat the eggs. Thus it
Natura History OF THE BELL VIREO 281
cannot be considered true incubation since no increase of tem-
perature in the eggs is effected by such means. He further notes
that it is at night when eggs are likely to experience a drop in tem-
perature that embryonic development will be impaired. I have no
data directly pertaining to which sex sits at night, but it is pre-
sumably the female, because she is always seen on the nest early in
the morning and late in the evening.
If a highly-vascularized brood patch is essential for true incuba-
tion, then it is surprising that males take regular turns on the nest in
cold, rainy weather. On May 20, 1960, male 3 (1960) sat on the
eggs longer than did the female (fig. 4). The temperature during
A. B.
Ge SY
4:30 p.m.
4:30 p.m.
4:00 p.m.
4:00 p.m.
3:30 p.m.
3: 30p.m.
3:00p.m.
2:30p.m.
Fic. 4. Comparison of periods of incubation by both sexes
in cold (54° F.) rainy weather (A) and in warm (82° F.)
sunny weather (B).
282 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
this hour and a half of incubation was 54° F. One solution to this
problem is supplied by Skutch (1957:74). He indicates that, “the
type of the incubation is determined largely by innate factors, so
that it persists through fairly wide fluctuations in weather, although
it may break down in extreme conditions.” Obviously then, in the
example described above, the weather conditions do not qualify as
“extreme.” Sitting by the male is certainly functional to some extent
for it relieves the female to forage; furthermore, the eggs are shel-
tered from inclement weather and protected from predators. No-
lan (1960:232) suggests similar reasons for incubating by the male
and adds the “conservation of heat supplied to the eggs by the fe-
male.”
My data, based on incubation beginning with the second egg,
indicate that the female incubates more often daily than the male
(fig. 5). The male sits on the eggs only occasionally in the morn-
ing, but almost as often as the female in the afternoon. Nolan
(1960:233) found that 95.5 per cent of the male’s time on the nest
and only 40 per cent of the female’s time were attributable to the
early hours of the day. Although I lack data on the critical hours
of 5:00 a.m. to 6:59 a.m., I have enough observations (20) from
7:00 a. m. to 9:00 a. m. to indicate that the males sit on the eggs in-
frequently (8 of 20 instances) in those hours. The discrepancy in
the two sets of data, which may be merely an artifact of sampling
techniques, does suggest two possible alternatives: (1) the male
15
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observer.
NATuRAL History OF THE BELL VIREO 283
sits on the eggs in the morning and gives the female, who sits on
the eggs throughout the night, an extended rest and an opportunity
to forage; (2) the female continues to sit throughout the morning,
especially during the early hours of daylight, a time of day when
the temperature may still be low enough to impair development of
the embryo.
Relief of Partners in Incubation
Relief of partners involves some ceremony. When the female is
incubating, the male sings several times as he approaches the nest
tree; the female responds with several chees, but otherwise remains
immobile. The male sings several more times upon alighting in the
nest tree whereupon the female chees again and flies directly from
the nest. A few seconds later the male appears at the edge of the
nest and, after inspecting the eggs, hops in and settles upon them.
When the male is sitting he is notably anxious prior to an exchange
with the female, often arising and craning his neck as he surveys the
surrounding vegetation, seemingly searching for his mate. The
singing of the male and the calling of the female serve as signals,
coordinating the exchange.
NESTLING PERIOD
Hatching Sequence
As indicated earlier, hatching normally occurs fourteen days after
the second egg is laid. Hatching of the young was staggered at
three nests under observation. In nest 2-b (1959) the first young
hatched on June 8, 1959, the second on June 10. In 3-b (1959)
one young hatched each day from the 12th through the 14th of
June. In 5-a (1959) two young hatched on June 15, the third on
June 16, and the fourth on June 17. Size of the young differed
notably for about three days as a result of staggered hatching, but
after that day the younger birds tended to catch up in size with
their older brood-mates. The fourth young in nest 5-a (1959)
grew steadily weaker and was missing from the nest on June 28,
1959. Staggered hatching is usually thought to be related to the
availability of food that will insure survival of at least some of the
nestlings when a shortage of food exists. It is doubtful that stag-
gered hatching has adaptive significance in the Bell Vireo, since
there seems to be no shortage of food for the young. In small
passerines such as the Bell Vireo the principal problem is to insure
fledging as quickly as possible because of the danger from predators.
284 UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist.
Development of the Nestlings
Young are pinkish at hatching and devoid of visible natal down.
Du Bois (in Wetherbee, 1957:380), inspected day-old nestlings by
means of a magnifying glass and was unable to detect any down.
Nolan (1960:236) also indicates that the young are naked at birth
and that the “body color is between flesh and rufous except
where folds of the straw yellow skin obscure the underlying colors.”
The Hutton Vireo (Vireo huttoni) is essentially naked at birth,
save for sparse hairlike down on the head and back (Wetherbee,
1953:380). The Red-eyed Vireo, according to Lawrence (1953:
67) is naked at birth save for a sparse covering of greyish natal
down, on the head, shoulders, and back.
In the Bell Vireo the pterylae darken slightly on the second day
and the color becomes more intense daily until the quills of the
dorsal tracts, the wings, and the tail break from their sheaths on
the sixth day. In Red-eyed Vireos the pterylae darken by the end
of the first day and the quills break through the skin on the fifth
day, erupting from the sheaths by the seventh day (Lawrence,
1953:67).
From the first day the young are able to squeak. Poking a young
bird was sufficient to elicit this sound, phonetically a nasal peek.
The only other vocalization noted throughout the nestling period
was an abbreviated chee.
For the first three days tapping the nest or even movement of it
caused by wind would elicit begging. By the fifth day at nest 2-a
(1959) only vigorous agitation of the branch to which the nest was
TABLE 9. MATURATION OF NESTLING BELL VinEOS. THE First Day THaT AN
Activiry Was OBSERVED Is SHOWN.
Day of nestling life
Hiyes Open. ot... lice sie hase is utters] 910.5 oflee a8 x
Feathers erupt009, So oicesl. san |e cs oie: oleic: x
Sound: Baten bis eiete as x
£
Head scratching and
Peering soi hs ioe [ote diesel rations] sfeyetel| Stalovelflaverscet| eyarema lleehe de x
Hopping to rim of nest3)2 see cfe a erste eis)] cee Sicilia edo 56
MICA PINE. 55 fe. swat cele he CHENG gale Sie oscil age ele eee al eackoneld ta verall eh arerelp stator xe
* This is the commonest fledging day.
NATURAL HisTORY OF THE BELL VIREO 285
attached evoked any response. At this nest on June 16, 1959, one
young begged while the other cowered. Cowering is correlated
with opening of the eyes, as the young bird that begged had its
eyes only partly open. Both young cowered on June 19, 1959.
Table 9 summarizes the maturation of the nestling Bell Vireos.
Parental Behavior
No eggshells were found in nests on the days of hatching. Pre-
sumably they had been removed by the parents. Nolan (1960:234)
indicates immediate disposition of the eggshell after hatching. Law-
rence (1953:62) suggests that conspicuous removal of eggshells by
the female Red-eyed Vireo informs the male that the young have
hatched.
Both sexes brood and the exchange of partners resembles that
described for the incubation period. Decrease in brooding in the
daytime begins about the sixth day of nestling life. Nolan (1960:
235) reports a sharp decrease in brooding when the oldest nestlings
are seven days old. Brooding decreases notably on the sixth day
of nestling life in the Red-eyed Vireo (Lawrence, 1953:62). Nice
(1929:17), Hensley (1950:244), and Nolan (1960:235) report that
the female Bell Vireo assumes a slightly greater role in brooding
than the male.
Apparent sun-shading was noted at nest 3-b (1959) at 2:00 p.m.
on June 17, 1959, on the fifth day of the nestling period. The nest
contained three young. An adult flew to the nest; while standing
on its rim the bird dipped its head into the nest six times, afterward
appeared to be eating a fecal sac, than shifted position to the un-
attached portion of the rim, gaped three times, thereupon spread
its wings, and sat motionless 85 minutes. In this attitude it formed
an effective shield sheltering the young from direct sunlight pene-
trating the thin foliage of the honey locust in which the nest was
situated. The temperature at this time was 95° F., but the sky was
partly cloudy. By 2:30 p.m. the sky had become overcast and the
sun passed behind a cloud. Although sunlight no longer fell di-
rectly upon the nest, the bird remained in the shielding posture for
another five minutes before flying from its perch. Sun-shading was
not observed at either of the other nests containing young; dense
overhead vegetation protected those nests. Sun-shading has been
noted in other species where the nest was poorly protected from the
sun. Lawrence (1953:62) observed this behavior at two Red-eyed
Vireo nests in conifers. The “sun-shield” posture of the Bell Vireo
does not correspond to any of the sunning postures described by
Hauser (1957).
286 UNIVERSITY OF Kansas PuBLs., Mus. Nat. Hist.
Feeding of the Nestlings
Both sexes fed the young, and presumably began shortly after the
first nestling hatched. My data indicate that the female does more
feeding than the male (Table 10); in about eight hours of observa-
tion a total of 67 morsels were brought, 43 by the female and 24 by
the male, for an average of once every 7.6 minutes. Nice (1929:17),
however, observed a male to bring food 53 times as compared to
21 visits by the female. In five and one-half hours of watching,
meals were brought once every 4.9 minutes. Du Bois (in Bent,
1950:257) recorded seven trips in an hour and forty minutes, or one
every fourteen minutes.
At three nests containing young the adults were sometimes silent
and sometimes vocal on their approach. The female often emitted
a subdued chee which, coupled with the vibration of the nest caused
by her arrival, elicited begging behavior from the young. None of
the males was heard to utter such a call, but I have the impression
that they often did call although I failed to hear the sounds. The
TABLE 10. FEEDING OF THE NESTLINGS.
Adult involved
Day of Length of
nestling period observation
Male Female
Yo est Peet Settee PRS rate te 30 min. 3 5
PEERS Rae ts, eee Ons OTE TOIOT. 60 min. 1 4
yes apa ics Sie Mf ube axplli bela ed 60 min. 2 5
AG) OA AU nietn Reid Bene ER EE TES 30 min. 1 4
ea Pein deen Renee re ofits ton 60 min. 4 7
Del koe: lar oA hE Sy ak a J aa 60 min. 3 3
Gis ed Mates Se eee he les 60 min. 3 6
TEE Ree ORS ID o COT oTE 30 min. 3 3
OAEAIYE) TAN Cote ieee ene a oer. Snel a 5 60 min. 4 6
Motalsin eee ke See eeeic 510 min. 24 43
males did, on occasion, sing several songs as they approached, even
with food held in their beaks. Such singing elicited begging from
the nestlings. Once the eyes of the young were open they often
began begging when a silent adult was within two or three feet of
the nest; begging behavior probably is elicited by tactile, auditory
or visual stimuli in that order, or, as the nestling period proceeds,
by any combination of these stimuli.
Not all trips made by parents resulted in successful feeding of
young; some visits seemed to be purely for inspecting the young.
NATURAL HisToRY OF THE BELL VIREO 287
On other occasions the adults experienced difficulty in transferring
food to the young, and, thus thwarted, would themselves eat the
food. Nice (1929:17) estimated that from five to twelve of a total
of seventy-five meals were eaten by adults.
Nest Sanitation
Both parents regularly removed fecal sacs from the nest, eating
them for the first five days and thereafter carrying them off and
presumably dropping them. It is doubtful that fecal sacs were
actively removed in the last two days of nestling life as the bottoms
of nests from which young flew away were invariably covered with
excrement.
On several occasions a parent brought food to the nest and then
remained perched on the rim alternately peering into the nest and
then preening. Once bill swiping was observed and another time
an adult male sang once. The adult remained at the nest from
twenty seconds to a full minute.
Fledging
Eight young were fledged from the four nests in 1959. The
nestling period lasted from nine to twelve days. Human interfer-
ence may have been largely responsible for the fledging of the
young at nine days. Pitelka and Koestner (1942:100) found nestling
life to last eleven days. Nolan (1960:235) reports nestling periods
varying from 10.5 to 12 days. The young Red-eyed Vireo is ready
to leave the nest at ten days but often remains an additional day
before departing (Lawrence, 1953:68).
The oldest nestling at nest 2-a (1959) hopped out on June 17,
1959, when I disturbed the parents. On this date the juvenal
plumage was only partly developed and the young bird was in-
capable of flight. By the tenth day of nestling life the young in all
the nests were observed to hop to the rim, flutter their wings, hop
back into the nest and also to preen and scratch their heads, The
young at fledging are usually completely feathered, but have notably
short tails and relatively short, stubby wings. According to Ridge-
way (1904:205) the juvenal plumage is much like that of the adult.
Nest Parasites
Pitelka and Koestner (1942:103) found that incubating adults
and later the young suffered infestation of the northern fowl mite,
Ornithonyseus sylviarum, Nolan (1960:241) reports a heavy infes-
tation of this mite at four nests. Unidentified mites were noted at
four nests in my study area in 1959. Incubating adults were ob-
288 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
served to peck at their breasts and scapulars from the eleventh
through the fourteenth day of incubation. Serious infestations were
not noted at the nests until the ninth day of nestling life. At this
time the young were observed to scratch their heads and peck at
their breasts, scapulars, and the base of their tails. On the day of
fledging the nests were a seething mass of crawling mites; the mites
also extended well up the branches to which the nests were at-
tached, Nest l-a (1959), which was discovered on June 18, 1959,
presumably on the day after fledging, was densely covered with
mites. Some mites were still crawling on this nest on June 20, 1959.
FLEDGLING LIFE
On June 20, 1959 I located one young 80 feet northeast of nest
2-a (1959), about five hours after it had left the nest. One parent
was observed to feed it once. No young were seen thereafter from
this or any other nest. Extreme agitation on the part of one or both
parents on several occasions shortly thereafter, however, sug-
gested the proximity of the young. Search in the immediate
vicinity on each of these occasions proved fruitless, Three days
after fledging their young, pair 2 (1959) was primarily occupied
with courtship activities. Pair 1 (1959) was involved in courtship
and nestbuilding one and one-half days after the apparent fledging
of their young. Nolan (1960:238) indicates that the young remain
within the territory and perhaps are fed by the parents up until an
age of about 40 days. Sutton (1949:25) and Lawrence (1953:68)
present contradictory reports on fledgling-parent relationships in
the Red-eyed Vireo. Sutton concluded that the young quickly
took leave of their parents whereas Lawrence reported a young
bird being fed 35 days after fledging.
Second Broods
The curve based on 66 nesting records of the Bell Vireo repre-
senting the breeding activity in northeastern Kansas demonstrates
a tendency toward double-broodedness (fig. 6). The peak of the
breeding season is from May 20 to June 20. The large number
(20) of replacement nests built in late May of 1960 tends to distort
the curve of the breeding data; a second peak about 35 days after
the first is evident.
I am of the opinion that the vast majority of vireos are single-
brooded solely by virtue of the limited success of early nesting
efforts, and that in “good” years most pairs would be double-
NATURAL History OF THE BELL VIREO 289
Number of Records
TGP ik 1 Oat del 1 Me MG | am 0 ml EM} 2 el 0
May June July
Fic. 6. Breeding season in northeastern Kansas based on the number of com-
pleted clutches in each 10-day period from May through July.
brooded. Each of the four pairs that successfully raised one brood
in my study area in 1959 renested within a day or two after the
fledging of the young. I do not know the fate of these nests. Nolan
(1960:237) reports at least one instance of a second brood in the
course of his study. Nolan (op. cit.) notes that the literature, in
general, indicates that vireos are double-brooded, but that his
evidence, mentioned previously, is the only evidence based on
banded birds.
REPRODUCTIVE SUCCESS
Only four nests were successful; all of these were observed in
1959. The principal external factors responsible for nesting failure
were severe weather, predation, parasitism by Brown-headed Cow-
birds (Molothrus ater) and human interference (Table 11).
In late winter and early spring of 1960 heavy snow, continuously
at a depth of at least 10 inches, covered most of the Mid-west from
February 20 through March 20, Consequently, the growing season
was some two weeks behind that of 1959. Of all the species in the
4—1506
290 UNIVERSITY OF Kansas PuBLs., Mus. Nat. Hist.
study area, the Bell Vireo is the most dependent on dense foliage
for cover and concealment for its nests. Consequently the tardiness
of the season seemingly negatively influenced reproductive success
of this more than any other species of bird in the study area.
Behavior
Several aspects of the behavior of the Bell Vireo tend to contribute
to nesting failure. They include:
1. Nest-site. Nests are occasionally suspended from exposed
branches. Occurrences of this sort suggest that the dimensions of
the fork are more important in the choice of a site than availability
of cover.
2. Song. The loud, continuous song of the male during nest-
building alerts cowbirds and predators to the presence of a nest.
The incongruous habits of the male of singing in the nest tree and
while sitting on the nest may facilitate location by some enemies,
particularly cowbirds.
TABLE 1]. Ecco Morratiry IN BELL ViIREOs.
Mortalit t eee 1960
Mortality agents
Per cent Per cent
Predationes ty cise ern: : 10
Weathers taro ee ; 16
Cowbirds i one se 74
Notals. stones ee 100
* Number of eggs out of the total number laid lost to mortality agents.
$+ In 1959 nine eggs were successful (ultimately gave rise to fledglings).
I am not fully convinced that song from the nest is simply a
“foolish” habit, since snakes, the principal predators with which
this species has to contend, are deaf. My own field observations
and the circumstances of the innumerable instances recorded in the
literature of male vireos singing from the nest suggest that this is a
function of the proximity of the observer. As mentioned elsewhere,
vocal threat is the initial as well as the primary means by which
territory is maintained. Song from the nest evoked by an enemy
also serves to alert the female to danger.
3. Flushing. The Bell Vireo normally relies upon cryptic be-
havior to avoid detection at the nest. Most sitting birds, especially
the females, either flush silently when an enemy is about forty feet
NATURAL HistTORY OF THE BELL VIREO 291
from the nest or remain sitting upon the nest tenaciously, refusing
to flush even when touched or picked up. Some birds flushed at
intermediate distances of from three to fifteen feet. In so doing
they revealed the location of their nests. Since none of these
“intermediate flushers” enjoyed nesting success there is possibly
some correlation between these two factors.
Predation
Several complete clutches being incubated disappeared from
nests that were unharmed. Absence of eggshells in the vicinity sug-
gests predation by snakes.
On May 25, 1960, I found a Peromyscus climbing toward nest l-a
(1960). The mouse moved to within two inches of the nest where-
upon I removed the mouse. Such small rodents constitute another
potential source of predation.
Cowbird Parasitism
In this study the failure of 12 of 35 nests can be directly attributed
to cowbird interference. It is well established that the incidence
of cowbird parasitism of Bell Vireo nests is high (Friedmann,
1929:237; Bent, 1950:260-261). Nolan (1960:240) found only one
nest of eight studied to be parasitized by cowbirds. He indicates
that this is surprising in view of the heavy molestation of the Prairie
Warbler (Dendroica discolor) in the same region. A possible
explanation of this phenomenon seems to lie in the much greater
abundance of the Prairie Warbler in comparison to that of the
Bell Vireo. In my study area the incidence of cowbird parasitism
on Bell Vireos in 1959 and 1960 greatly exceeded that of all other
nesting species that were parasitized (Table 12).
As indicated previously, the female Bell Vireo leaves the nest
unoccupied several hours at a time in the transition period between
completion of the nest and the start of egglaying. Such behavior
early in the morning certainly would facilitate deposition of cow-
bird eggs. Early in the nesting period the mere presence of a
cowbird egg in the nest prior to the laying of the host’s first egg
leads to abandonment of the nest. This seems to be correlated
with the relative strength of the nesting tendency; anyhow cowbird
eggs laid in later nests prior to the appearance of the host’s own
eggs did not cause the nesting birds to desert. The Bell Vireo does
abandon the nest when all but one of its own eggs have been re-
moved by the cowbird. Mumford (1952:232) records the removal
of a cowbird egg by the host birds and I recorded a similar instance
292 UNIVERSITY OF KANsAs Pusts., Mus. Nat. Hist.
TABLE 12. INCMENCE OF COWBIRD PARASITISM OF THE BELL ViEO CoMPARED
WiTH OTHER PASSERINES IN THE STuDy AREA IN 1959 Aanp 1960.
involving nest 2-b (1960). On May 14, 1960, I found one punctured
cowbird egg on the ground about 10 feet west of this nest. Oc-
casionally a cowbird egg is buried beneath the lining of a nest.
Mumford (1952:23) observed this in mid-May in 1951 and I ob-
served pair 8 (1960) actively covering with building material a
cowbird egg on July 5, 1960. Covering a cowbird egg constitutes
effective removal. Since the egg cannot be turned, an adhesion de-
velops.
The percentage of cowbird eggs hatched in relation to the number
laid is relatively low. For instance, Mumford (1952:231) has only
one record of a young cowbird successfully raised by a Bell Vireo.
The data available in Bent (1950:260-261) also indicate that the
percentage of cowbird eggs hatched is small. The Bell Vireo is
less tolerant of cowbird parasitism than are many of the species
so victimized, but is not so intolerant as the Robin, Catbird, and
the Yellow-breasted Chat (Friedmann, 1929:193).
SUMMARY
1. The behavior of a small population of Bell Vireos was studied
in the spring and summer of 1959 and again in 1960 in Douglas
County, Kansas, and results are compared with previous studies
elsewhere.
2. The Bell Vireo sings more often daily and throughout the
nesting season than do the majority of its avian nesting associates.
Six types of vocalizations are readily distinguishable in the field:
primary song, courtship song, distress call, alarm note, specialized
male call note or zip, and the generalized call note or chee.
3. Territories are established in early May and occupied through-
out the breeding season and post-breeding season. The average
NATURAL History OF THE BELL VIREO 293
size of the territories in 1960 was 1.25 acres. Shifting of territorial
boundaries occasionally occurs after nesting attempts.
4, Territory is maintained primarily by song, but at least five
aggressive displays are manifest in the early phases of territorial
establishment. These include: (a) vocal threat, (b) head-forward
threat, (c) wing-flicking and sub-maximal tail-fanning, (d) ruffling
and maximum tail-fanning, and (e) supplanting attack.
5. The precise mechanism of pair-formation in the Bell Vireo
is not known. Early courtship activities are characteristically violent
affairs. Absence of sexual dimorphism suggests that behavioral
criteria are used by the birds in sex-recognition; the male is dom-
inant and the female is subordinate.
6. The principal displays associated with courtship include:
greeting ceremonies, “pouncing,” “leap-flutter,” pre- and post-copula-
tory displays, and the posture, copulation. The marked similarity
between elements of courtship display and aggressive display sug-
gests common origin or the derivation of one from the other.
7. The nest-site probably is selected by the female. Nests are
suspended from lateral or terminal forks about 2 feet 3 inches high
in small trees and shrubs averaging 11 feet 2 inches in height.
8. Nestbuilding is intimately associated with courtship and is a
responsibility of both sexes. The male builds the suspension ap-
paratus and the female constructs and lines the bag. Both sexes
participate in adorning the exterior. Construction lasts from four
and one-half to five days.
9. The nest is compact, pendant, and composed of strips of bark
and strands of grasses that are interwoven and tightly bound with
animal silk. Nests built in May are bulkier than those constructed
later in the season.
10. Egglaying begins on the first or second day after the nest
is completed. The eggs are deposited early in the morning. The
average clutch-size of the Bell Vireo in Kansas is 3.39 eggs.
11. Both sexes sit on the eggs, but only the female truly incubates
because the male lacks a brood patch. Incubation lasts fourteen
days.
12. The Bell Vireo is double-brooded in “good” years.
13. Nesting failure resulted from severe weather, predation,
parasitism by cowbirds, and human interference. Behavior that
contributes to nesting failure is selection of an unfavorable nest-site,
singing on and near the nest, and the tendency to flush from the nest
in view of potential enemies.
294 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
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1960. Breeding behavior of the Bell Vireo in southern Indiana. Condor,
62:225-244,
PITELKA, F, A.
1959. Numbers, breeding schedule, and territoriality in Pectoral Sand-
pipers of northern Alaska. Condor, 61:223-264.
PITELKA, F. A., and KorEstnenr, E. J.
1942. Breeding behavior of Bell’s Vireo in Illinois. Wilson Bull., 54:97-
106.
296 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Riweway, R.
1889. The ornithology of Illinois. [Illinois State Nat. Hist. Survey, 1:
viii + 520 pp.
1904. The birds of North and Middle America. U.S. Nat. Mus. Bull.,
50, pt. 3; xx + 801 pp.
Sxutcu, A. F.
1957. The incubation patterns of birds. Ibis, 99:69-93.
SOUTHERN, W. E.
1958. Nesting of the Red-eyed Vireo in the Douglas Lake Region, Michi-
gan. The Jack-Pine Warbler, 36:105-130, 185-207.
Sutton, G. M.
1949. Studies of the nesting birds of the Edwin S. George Reserve. Part 1.
The Vireos. Misc. Pub. Univ. Michigan Mus. Zool., 74:5-36.
TOWNSEND, C. W.
1920. Supplement to the birds of Essex County, Massachusetts. Mem.
Nuttall Orn. Club, No. 5:196 pp.
TyLer, W. M.
1912. A vireo courtship. Bird Lore, 14:229-230.
Van Tyne, J., and BERGER, A. J.
1959. Fundamentals of omithology. John Wiley & Sons, Inc., New York.
xi + 624 pp.
WETHEBBEE, D. K.
1957. Natal plumages and downy pteryloses of passerine birds of North
America. Bull. Amer. Mus. Nat. Hist., 113:5:339-436.
Transmitted November 8, 1961.
O
29-1506
8.
4,
5.
6.
wf.
8.
9.
_ 10.
11.
12.
13.
14,
15.
16.
17.
18.
19.
20.
21.
22.
28.
(Continued from inside of front cover )
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.
Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming.
By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956.
The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6
figures in text. May 19, 1956.
Additional remains of the multituberculate genus Eucosmodon. By Robert
W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.
Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures
in text. June 15, 1956.
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.
Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp.
847-351. August 15, 1956.
A new bat (Genus Leptonycteris) from Coahuila. By Howard J. Stains.
Pp. 353-3856. January 21, 1957.
A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico.
By Robert J. Russell. Pp. 857-361. January 21, 1957.
Geographic variation in the pocket gopher, Thomomys bottae, in Colorado.
By Phillip M. Youngman. Pp. 363-387, 7 figures in text. February 21, 1958.
New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones,
Jr. Pp. 885-388. May 12, 1958.
Pleistocene bats from San Josecito Cave, Nuevo Leén, México. By J. Knox
Jones, Jr. Pp. 389-396. December 19, 1958.
New subspecies of the rodent Baiomys from Central America. By Robert
L. Packard. Pp. 897-404. December 19, 1958.
Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405-
414, 1 figure in text, May 20, 1959.
Distribution, variation, and relationships of the montane vole, Microtus mon-
tanus. By Sydney Anderson. Pp. 415-511, 12 figures in text, 2 tables.
August 1, 1959.
Conspecificity of two pocket mice, Perognathus goldmani and P. artus, By
E. Raymond Hall and Marilyn Bailey Ogilvie. Pp. 518-518, 1 map. Janu-
ary 14, 1960.
Records of harvest mice, Reithrodontomys, from-Central America, with de-
scription of a new subspecies from Nicaragua. By Sydney Anderson and
J. Knox Jones, Jr. Pp. 519-529. January 14, 1960. :
Small carnivores from San Josecito Cave (Pleistocene), Nuevo Leén, México.
By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, 1960.
Pleistocene pocket gophers from San Josecito Cave, Nuevo Leén, México.
By Robert J. Russell. . Pp. 539-548, 1 figure in text. January 14, 1960.
Review of the insectivores of Korea. By J. Knox Jones, Jr., and David H.
Johnson. Pp. 549-578. February 23, 1960.
Speciation and evolution of the pygmy mice, genus Baiomys. By Robert L.
Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960.
Index. Pp. 671-690.
Vol. 10. 1.
2.
10.
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.
Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.
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 $1, 1956.
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.
Birds found’ on the Arctic slope of northern Alaska. By James W. Bee.
Pp. 168-211, plates 9-10, 1 figure in text. March 12, 1958.
The wood rats of Colorado: distribution and ecology. By Robert B. Finley,
Jr. Pp. 213-552, 84 plates, 8 figures in text, 85 tables. November 7, 1958.
Home ranges and movements of the eastern cottontail in Kansas. By Donald
W. Janes. Pp. 553-572, 4 plates, 3, figures in text. May 4, 1959.
Natural history of the salamander, Aneides hardyi. By Richard F. Johnston
and Gerhard A. Schad. Pp. 578-585. October 8, 1959.
A new subspecies of lizard, Cnemidophorus sacki, from Michoacin, México.
By William E. Duellman, Pp. 587-598, 2 figures in text. May 2, 1960.
A taxonomic study of the middle American snake, Pituophis deppei. y
William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960.
\ Index. Pp. 611-626.
Vol. 11. 1. The systematic status of the colubrid snake, Leptodeira discolor Giinther.
2.
3.
By William E. Duellman. Pp. 1-9, 4 figures. July 14, 1958.
Natural history of the six-lined racerunner, Cnemidophorus sexlineatus. By
Henry S. Fitch. Pp. 11-62, 9 figures, 9 tables. September 19, 1958.
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; 8 tables. December 12, 1958.
A new snake of the genus Geophis from Chihuahua, Mexico. By John M.
Legler. Pp. 327-334, 2 figures in text. January 28, 1959.
(Continued on outside of back cover)
Vol. 12.
(Continued from inside of back cover)
A new tortoise, genus Gopherus, from north-central. Mexico. By John M.
Legler. Pp. 335-343. April 24, 1959.
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.
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.
aan from Coahuila, México, By Emil K. Urban. Pp. 443-516. August 1,
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, ©
Natural history of the omate box turtle, Terrapene ornata omata Agassiz. By
John M. Legler. Pp. 527-669, 16 pls., 29 figures in text. March 7, 1960.
Index Pp. 671-703.
AM
2.
8.
4,
5.
Vol. 18. 1.
Vol. 15.
2.
|
.
Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry
A. Vaughan. Pp. 1-158, 4 plates, 24 figures in text. July 8, 1959.
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.
The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49
figures in text. February 19, 1960,
A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By
Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in
text. May 2, 1960. ‘
Natural history of the bell vireo. By Jon C. Barlow. Pp. 241-296, 6 figures
in text. Mareh 7, 1962.
More numbers will appear in volume 12.
Five natural hybrid combinations in minnows (Cyprinidae). By Frank B.
Cross and W. L. Minckley. Pp. 1-18. June 1, 1960.
A distributional study of the amphibians of the Isthmus of Tehuantepec,
México. By) William E. Duellman. Pp. 19-72, pls. 1-8, 8 figures in text.
August 16, 1960.
A new subspecies of the slider turtle (Pseudemys scripta) from Coahuila,
Merce By John M. Legler. Pp. 73-84, pls. 9-12, 8 figures in text. August
Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, pls. 13-20,
26 figures in text. November 30, 1960.
Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and
Rocky Mountains. By Henry S. Fitch and T. Paul Maslin. Pp. 289-308,
4 figures in text. February 10, 1961.
Fishes of the Wakarusa river in Kansas. By James E. Deacon and Artie L.
Metcalf. Pp. 309-322, 1 figure in text. February 10, 1961.
Geographic variation in the North American cyprinid fish, Hybopsis gracilis. —
By Leonard J. Olund-and Frank B. Cross. Pp. 323-348, pls. 21-24, 2 figures
in text. February 10, 1961.
Descriptions of two species of frogs, genus Ptychohyla; studies of Ameri-
can hylid frogs, V. By William E. Duellman. Pp. 349-357, pl. 25, 2
figuresin text. April 27, 1961.
Fish populations, following a drought, in the Neosho and Marais des Cygnes
rivers of Kansas. By James Everett Deacon. Pp. 359-427, pls. 26-80, 8 figs.
August 11, 1961.
Recent soft-sheHed turtles of North America (family Trionychidae). By
ee Webb. Pp. 429-611, pls. 31-54, 24 figures in text. February
Neotropical bats. from western México. By Sydney Anderson. Pp. 1-8.
October 24, 1960.
Geographic variation in the harvest mouse, Reithrodontomys megalotis, on
the central Great Plains and in adjacent regions. y J. Knox Jones, Jr.,
and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961.
Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson. —
Pp. 29-67, pls. 1 and 2, 3 figures in text. July 24, 1961. \ :
A new subspecies of the black myotis (bat) from eastern Mexico. By E.
Paes Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December
North American yellow bats, “‘Dasypterus,” and a list of the named kinds
of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr.
Pp. 73-98, 4 figures in text. December 29, 1961. ‘
Natural history of the brush mouse (Peromyscus boylii) in Kansas with
description of a new subspecies. By Charles A. Long. Pp, 99-111, 1 figure
in text. December 29, 1961. q
Taxonomic status of some mice of the Peromyscus boylii group in eastern
Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 118-
120, 1 figure in text. December 29, 1961.
A new subspecies of ground squirrel (Spermophilus spilosoma) from Ta-
maulipas, Mexico: By Ticul Alvarez. Pp. 121-124. March 7, 1962.
Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J.
Knox Jones, Jr., and Ticul Alvarez. Pp. 125-133, I figure in ‘text. March 7,
1962.
More numbers will appear in volume 14.
The amphibians and reptiles of Michoacan, México. By William E. Duell-
man. Pp. 1-148, pls. 1-6, 11 figures in text. December 20, 1961.
Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox
Jones, Jr., and George W. Byers. Pp. 149-173. January 31,1962.
A new species of frog (Genus Tomodactylus) from western México. By
Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962.
More numbers will appear in volume 15.
UNIVERSITY OF KANSAS Pree {UUL 6
| a 4 af
JUL-6 = 19592
MusEuM OF NATURAL History
A
Volume 12, No. 6, pp. 297-307, 6 aks UNIVE
May 21, 1962
Two New Pelycosaurs from the Lower Permian
of Oklahoma
BY
RICHARD C. FOX
UNIVERSITY OF KANSAS
LAWRENCE
™
1962
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HisTORY
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 12, No. 6, pp. 297-307, 6 figs.
Published May 21, 1962
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY
JEAN M. NEIBARGER, STATE PRINTER
TOPEKA. KANSAS
{
Two New Pelycosaurs from the Lower Permian’
of Oklahoma
1 ,
BY } 1 AUG Lie
RicHARD C. Fox
In the course of examining material from fissure deposits of early
Permian age collected from a limestone quarry near Fort Sill, Ok-
lahoma, the author recovered several tooth-bearing fragments of
small pelycosaurs. The fragments were examined, compared with
descriptions of known kinds appearing in the literature, and deter-
mined to be new genera within the Nitosauridae (Edaphosauria)
and Sphenacodontidae (Sphenacodontia ).
Appreciation is expressed to Prof. Theodore H. Eaton, Jr., for
permission to examine the collections of the University of Kansas
from Fort Sill, and for the financial assistance furnished by his Na-
tional Science Foundation grant (NSF-G8624). I am grateful both
to Prof. Eaton and Mr. Dale L. Hoyt for their suggestions regarding
this manuscript. The accompanying figures have been drawn by the
author.
Family NITOSAURIDAE
Delorhynchus priscus new genus and new species
(delos, Gr., evident; rhynchos, Gr., neuter, nostril; priscus, L., ancient. Delorhynchus
is masculine because of the ending that it acquires when transliterated into Latin. )
Type specimen.—Fragmentary left maxilla, bearing four teeth, KU 11117.
Referred specimens.—Fragmentary right maxilla having four teeth, KU
11118; fragmentary left maxilla having four teeth, the most posterior of which
has been broken, KU 11119.
Horizon and locality—A fissure deposit in the Arbuckle limestone at the
Dolese Brothers Limestone Quarry, approximately six miles north of Fort Sill,
in sec. 31, T. 4 N, R. 11 W, Comanche County, Oklahoma. These sediments
are of early Permian age, possibly equivalent to the Arroyo formation, Lower
Clear Fork Group of Texas (Vaughn, 1958: 981).
Diagnosis —Small; marginal teeth conical, slender and recurved at tips;
marginal tooth-row without caniniform enlargement; narial opening enlarged
and bordered dorsally, posteriorly and ventrally by maxilla; maxilla with fora-
men opening laterally at posteroventral corner of naris.
Description (based on 8 maxillary fragments, see Table 1).—
Each of the maxillary fragments bears four thecodont teeth. These
are conical, slender and sharply pointed; in their distal third they
are slightly recurved, laterally compressed, and have anterior and
posterior non-serrated cutting edges. In medial aspect at their
bases, the teeth are longitudinally striated. The bases of the teeth
(299)
300 UNIVERSITY OF Kansas Pusxs., Mus. Nat. Hist.
are circular in cross-section and are slightly bulbous. There is no
caniniform enlargement of any of the teeth, the longest tooth of
each fragment being differently placed in the series of teeth and
little longer than the others. There is no swelling on either the
internal or external surfaces of the maxillae. The teeth are in a con-
tinuous series; no diastema or maxillary step is evident.
Ficures 1-3. Delorhynchus priscus, lower Permian, 6 miles north of Fort Sill,
Comanche County, Oklahoma. All x 8.
Fic. 1. KU 11117 (type specimen), lateral view of left maxilla.
Fic. 2. KU 11118, lateral view of right maxilla.
Fic. 8. KU 11119, lateral view of left maxilla.
The fragments have been broken along similar lines of fracture,
and each is approximately rhomboidal in shape. The maxilla en-
circles the posterior border of the naris and extends dorsally above
the naris to an extent sufficient to indicate the probable exclusion of
the lacrimal bone from the narial border. At the posteroventral
corner of the naris a foramen opens onto the lateral surface of the
maxilla. The opening is the entrance to a canal that runs posteriorly
TABLE 1.—DIMENSIONS, IN MILLIMETERS, OF THREE MAXILLARY FRAGMENTS OF
DELORHYNCHUS PRISCUS
~ = ran
me | BS Se deeeh tale =
‘BS 5 2 | bs = =
CaTALOGUE NuMBER| << §& rants 32 a 6 bp a
AND MEAN 5s Gis are So iinea ole aa
eo Se ee ones es | 5S
eK) 2S BAS 5.5 o 8 =I)
< Ay 4 fa a <
DSU i 10 Gi Wn PCa eee ere pee ke 6.0 8.0 6.0 8.0 3.0 3.0
DG OA es i ee suena 6.0 6.0 9.0 8.0 2 3.0
JEQUINT LTH eee croioe oe 6.6 8.0 10.0 1150 ts 4.6
MWieami aeons. cee 6.2 hee) 8.3 9.0 2.5 4.5
New PELYCOSAURS FROM LOWER PERMIAN 3801
above the tooth-row throughout the length of each specimen. Be-
neath the naris the maxilla extends as a broad tapering shelf, the
ventral surface of which articulates with the premaxilla. The narial
rim is wide, but wider ventrally than dorsally. The plane of the
narial rim is oblique to the lateral surface of the maxilla. The ex-
ternal surface of each fragment is grooved and pitted. The ossifi-
cation of each fragment appears to have been complete.
Discussion.—The Nitosauridae are small primitive edaphosaurs
with a moderately elongate face, sharp subisodont teeth, little de-
velopment of canines and few specializations. The jaw is of a
primitive type and articulates on a level with the tooth-row. The
palatal dentition is primitive (Romer, 1956:280). The nitosaurids
are thought to be related to the later Caseidae, and the most obvious
structural similarities are found in the postcranial skeleton ( Vaughn,
1958:989). Cranial resemblances between the families are fewer,
but nevertheless indicate that a nitosaurid-caseid relationship exists.
Vaughn (1958) described a small pelycosaur, Colobomycter pho-
leter (Eothyrididae, Ophiacodontia) that structurally resembles the
Caseidae. This individual also was obtained from the Fort Sill lo-
cality. In Vaughn’s opinion the features of Colobomycter indicate
a close relationship between eothyridids and caseids and the possi-
bility that the caseids may well have been of eothyridid rather than
nitosaurid derivation.
In view of this historical uncertainty of the relationships between
the Nitosauridae, the Eothyrididae and the Caseidae, it is well to
consider how the maxillary fragments described above differ from
and resemble representatives of each of these three families as re-
ported in the literature.
Delorhynchus resembles Colobomycter in size. The mean extra-
maxillary length of the undamaged teeth of the three fragments is
2.5 mm., equal to that reported by Vaughn (1958:985) for teeth
about midway in the postcanine series of Colobomycter. None of
the teeth of Delorhynchus extends beyond the maxillary rim as far
as does the canine of Colobomycter (3.5 mm. ).
The teeth in both genera are conical and sharply pointed. The
naris in each is enlarged, and the lacrimal is excluded from the
narial margin in each (by inference in Delorhynchus. )
The differences between the maxillae of Colobomycter and De-
lorhynchus are most striking in the lack of canines in the latter and
the correlated absence of modifications of the maxillary for support
of canines. Additionally, Delorhynchus bears an infraorbital canal
302 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
in contrast to the groove in similar position in Colobomycter. The
recurvature of the four teeth present in the fragments of Delorhyn-
chus differs from that in the teeth of Colobomycter in which only
the canine and precanine are recurved. Vaughn implies that an-
terior and posterior cutting edges extend the length of the teeth in
Colobomycter; these are restricted to the distal third of the teeth
in Delorhynchus. The external surfaces of the maxillae of De-
lorhynchus are pitted and ridged; Vaughn was unable to discern
sculpturing of the corresponding surfaces in Colobomycter.
Delorhynchus resembles the nitosaurids in size, the shape and
sharpness of the teeth, their recurvature and the slight enlargement
of their bases, the exclusion of the lacrimal bone from the narial
margin (in Mycterosaurus) and the apparent lack of a special ca-
nine pair of teeth. Resemblances to the caseids are to be noted in
the enlargement of the naris (4.5 mm. in height as opposed to 1.7
mm. in Colobomycter), lack of development of canines, presence of
an infraorbital canal (in Cotylorhynchus) and absence of many re-
placement gaps in the marginal row of teeth.
The absence of caniniform enlargement and the extension of the
maxilla dorsad of the naris exclude Delorhynchus from the Eothy-
rididae (Ophiacodontia) but are no bar to its inclusion in the Nito-
sauridae (Edaphosauria). The marginal teeth of Delorhynchus
are simple and primitive, being much like those of the nitosaurids
that are described in the literature.
The large narial opening and its posterior, dorsal and ventral en-
closure by the maxilla, the infraorbital canal, and the sculptured
external surfaces of the maxillary fragments indicate that Delorhyn-
chus, in these features at least, is close to achieving the caseid grade.
Family SPHENACODONTIDAE
Thrausmosaurus serratidens new genus and new species
(Thrausmosaurus is formed from the neuter Greek noun, thrausma, meaning fragment,
and the masculine Greek noun, sauros, meaning reptile. The specific name, serratidens, is
formed from the Latin serratus, meaning serrate, and the masculine Latin noun, dens, mean-
ing tooth. The specific name is used as a substantive in apposition with the generic name.)
Type specimen.—Fragmentary left dentary, bearing five teeth, the most pos-
terior of which is broken at the base, KU 11120.
Referred specimens.—Fragmentary Pleft maxilla, having two teeth, KU
11121; fragmentary left dentary having two teeth, KU 11122.
Horizon and locality—From the early Permian fissure deposits in the Ar-
buckle limestone of the Dolese Brothers Limestone Quarry, approximately 6
miles north of Fort Sill, in sec. 31, T. 4N, R. 11 W, Comanche County, Ok-
lahoma.
New PELYCOSAURS FROM LOWER PERMIAN 303
Diagnosis.—Small; teeth thecodont, compressed laterally, recurved distally,
and bearing anterior and posterior cutting edges; anterior serrations limited to
recurved portions of teeth, posterior serrations extending nearly entire length
of teeth; lateral compression of teeth more pronounced medially than laterally;
bases of teeth expanded.
Description—The type specimen is 16 mm. long. It bears five
teeth that are implanted in a straight row. Empty sockets are pres-
ent between the first and second teeth, and the third and fourth
teeth. The first tooth is 3.0 mm. long, the middle two are each
2.5 mm. long, and the fourth tooth is 2.0 mm. long. The fifth tooth
is broken off at its base.
The empty sockets are large. The mouth of each is circular and
approximately 2.0 mm. in diameter. Both sockets are 1.25 mm.
deep. The bases of the teeth are expanded to fill the sockets, al-
though the blades of the teeth arise from only the lateral portions of
the bases. The edge of the dentary rises above the bases of the
teeth medially, thereby producing a small depression at the junction
of each base with the dentary bone.
The lateral compression of the teeth is pronounced but asym-
metrical, in that the lateral surface of each blade is more convex
than the medial surface.
Ficures 4-6. Thrausmosaurus serratidens, lower Permian, 6 miles north of Fort
Sill, Comanche County, Oklahoma. All x 8.
Fic. 4. KU 11120 (type specimen), lateral view of left dentary.
Fic. 5. KU 11121, lateral view of Pleft maxilla.
Fic. 6. KU 11122, lateral view of left dentary.
The recurvature of the anterior cutting edges is much more se-
vere than that of the posterior edges, but the recurvature of both
is limited to the distal half of each tooth.
The serrations of the cutting edges are not visible to the naked
eye and are limited on the anterior edges of the teeth to those por-
tions of the blades that are recurved. The posterior serrations ex-
tend nearly to the junction of the blade of each tooth with its base.
The serrations tend to be more nearly crenulate than cuspidate.
304 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
A portion of the lateral wall of the dentary surrounding the Meck-
elian canal is present. The external surface of the wall is gently
convex and smooth, without sculpturing. The internal surfaces of
the canal are unmarked either by muscle scars or foramina.
The fragment is a piece from the posterior portion of the dentary,
since the decrease in height from the first tooth to the fourth is pro-
nounced.
KU 11122, a fragment of the left dentary bearing two teeth, is
7.5 mm. long. The anterior tooth is 3.0 mm. long; the posterior
tooth is 3.5 mm. long. The shape of the teeth and their implanta-
tion conform to the description of the type specimen. The lateral
surface of the fragment is smooth and gently convex. What little
is present of the surface bordering the Meckelian canal is unmarked.
The Pmaxillary fragment bears two teeth which are 3.0 mm. long,
and which conform in their characters to the type. The lateral,
medial and ventral surfaces of the fragment have been sheared off,
so that an exact identification of the bone is impossible. Presum-
ably the fragment is too deep dorsoventrally to be a piece of the
dentary, and no sign of the Meckelian canal is present.
Discussion.—The implantation, lateral compression, recurvature
and cutting edges of the teeth borne by these fragments make clear
their sphenacodontid nature. The characters of the fragments are
too few to determine subfamilial affinities, however. That the frag-
ments are the remains of adult animals can be only surmised from
the lack of bones or teeth of large pelycosaurs in the extensive col-
lections of the University of Kansas from the Fort Sill locality.
If Thrausmosaurus is, in fact, adult, the genus is an unusually
small sphenacodontid, and of significance both on that account and
because of the resemblance of the teeth presently known to those
of its far larger relatives.
The Fort Sill Locality—Peabody (1961) suggested that the fis-
sures of Fort Sill had been used as dens by predatory animals in the
early Permian, and that the unusually abundant bones in the fissures
were the remains of animals eaten there by these occupants. Evi-
dence now known to me affords an alternative explanation that is
presented here as a preliminary to a more complete study of the
fauna and paleoecology of these deposits currently being under-
taken.
The suggestion that the skeletal material found in the fissures is
the remnant of the prey of other animals is questionable because of:
Nrw PELYCOSAURS FROM LOWER PERMIAN 305
1. The absence of tooth marks on the fossils.
2. The recovery from the matrix of skulls and portions of articulated skele-
tons that are undamaged or damaged only by pressure after burial.
3. The rarity in the deposits of animals of larger body size than Capto-
rhinus, the exceptions being a few limb fragments and skull fragments
of labyrinthodont or pelycosaurian nature.
4. The absence of coprolites in the matrix.
If the fissures were the dens of predators, at least some and prob-
ably many of the bones would show tooth marks. A predator feed-
ing on other animals would be expected to leave some evidence of
its habits on the bones of its prey. No such evidence is known to
me, either from my own examination of several thousand bones or
from the reports in the literature by others who have studied aspects
of the early Permian fauna of Fort Sill.
If the predators were larger than Captorhinus and occupied the
fissures for a long enough time to account for the accumulation of
the tremendous numbers of individuals that are represented, a con-
siderable amount of the skeletal material of the larger animals
would be present in the fissure deposits. Even if for some reason
the predators died in areas other than within the fissures, thereby
accounting for the absence of large bones, coprolites should appear
in the deposits if, in fact, the fissures were feeding places. In view
of the nearly undamaged condition of many of the bones recovered
from the fissures, it is reasonable to expect that fecal material would
be preserved.
The character of the matrix of the deposits varies from a homo-
geneous clay to clay interrupted by layers of soft, limey, conglomer-
atic rock, to a hard, well-cemented, calcareous conglomerate. In
general the bone in each kind of matrix is colored characteristically
and exhibits a characteristic degree of wear. The bones entrapped
in the homogeneous clay are relatively few, black, usually disar-
ticulated, little worn and not unduly fragmented; consequently the
discovery of undamaged limb bones, for example, from this kind of
matrix is not unusual. The bones found in the stratified portion of
the matrix are more numerous within the layers of conglomerate
than between. The bones are black, brown or white, highly frag-
mented and waterworn to a variable degree. The fragments recov-
ered from the hard, calcareous matrix are numerous, range in color
from white through various shades of brown, to black, are highly
fragmented, and are usually worn by water.
These categories for bone and matrix, however, are not mutually
306 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
exclusive, since bones of any of these colors and exhibiting any de-
gree of wear and fragmentation are found in any of the kinds of
matrix described above. That water was the agent of wear is sug-
gested by the highly polished appearance of the worn bones and
pebbles that are found in the matrix.
The variability of the matrix and of the color and condition of
the bones indicates that the agencies of burial and fossilization dif-
fered from time to time and that the agency of transportation of the
bones from the site of burial to the fissures was running water. One
can easily visualize a stream coursing the early Permian landscape
that was subject to periodic flooding and droughts. Along the banks
of the stream and in its pools lived a variety of microsaurs, capto-
rhinids, small labyrinthodonts and small pelycosaurs. Some of the
animals, after they died, were either buried near the site of their
death or were swept along and buried in sediments further down-
stream. Burial was for a length of time sufficient to impart a color
to the bones characteristic of the site in which they were buried.
Later floods reexposed the sites of burial, picked up the bones and
carried them to the openings into the fissures. Presumably, too, a
proportion of the bones was carried to the fissures without previous —
burial.
The differences in wear exhibited by different bones within the
same block of matrix is attributable to differences in distance that
the bones were transported before final deposition. The final sites
of deposition, the fissures, were inundated occasionally by floods
alone, or because of changes in location of the channel of the stream
at the time of flooding. The periodicity of deposition of the sedi-
ments within portions of the fissures is indicated by the stratification
of the bone conglomerate mentioned earlier.
In summary, it seems that there is little or no evidence beyond
the numbers of bones involved to support the hypothesis that the
concentration of bones in the fissures of Fort Sill represents the re-
mains of food of predators, and that the fissures were used as dens
by their predatory occupants. On the contrary, the evidence indi-
cates that the deposition of the bones in the fissures was secondary |
and that the agency of transportation, deposition and accumulation |
of the bones was an early Permian stream characterized by periodic |
flooding.
NEw PELYCOSAURS FROM LOWER PERMIAN 807
LITERATURE CITED
Peapopy, F. E.
1961. Annual growth zones in living and fossil vertebrates. Jour. Morph.
108(1):11-62, 69 figs., January.
Romer, A. S.
1956. Osteology of the reptiles. The University of Chicago Press, Chi-
cago. xxi-++ 772 pp., 248 figs.
Romer, A. S., and Price, L. I.
1940. Review of the Pelycosauria. Geol. Soc. America, Spec. Pap., 28:
x + 538 pp., 71 figs., 46 pls., 8 tables, December 6.
Vaucun, P. P.
1958. On a new pelycosaur from the lower Permian of Oklahoma, and
the origin of the family Caseidae. Jour. Paleont., 32:981-991, 1
fig., September.
Transmitted March 15, 1962.
29-3001
UNIVERSITY OF KANSAS PUBLICATIONS
MusEeuM OF NATURAL HISTORY
Volume 12, No. 7, pp. 309-345, pls. 5-8
June 18, 1962
Vertebrates from the Barrier Islands
of Tamaulipas Mexico
BY
ROBERT K. SELANDER, RICHARD F. JOHNSTON, °
B. J. WILKS, AND GERALD G., RAUN
UNIVERSITY OF KANSAS
LAWRENCE
1962
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HiIsToRyY
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 12, No. 7, pp. 309-345
Published June 18, 1962
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY
JEAN M. NEIBARGER, STATE PRINTER
TOPEKA, KANSAS
en eer >
mance — ~ —caret |
}
Vertebrates from the Barrier Island
of Tamaulipas, México
BY { 13
ROBERT K. SELANDER, RICHARD F. JOHNSTON, B. J. WILKS, and —
GERALD G. RAUN
Lying between the Gulf of Mexico and the Laguna Madre de
Tamaulipas is a narrow barrier island extending from the delta of
the Rio Grande south for 140 miles to within 185 miles of Tampico,
Tamaulipas (Plate 5). This island, like most of coastal Tamauli-
pas, has been all but neglected by zoological collectors. Conse-
quently, little is known of the kinds, distribution, and seasonal status
of the vertebrates occurring there. The present paper is a report
on land vertebrates collected and observed on the northern part
of the barrier island of Tamaulipas from July 6 to 10, 1961. Our
collection, which has been deposited in the Museum of Natural
History, The University of Kansas, consists of 63 reptiles, 83 mam-
mals, and 97 birds (58 skins, 19 skeletons, and 20 alcoholics).
Acknowledgments
We are especially indebted to Dr. Charles H. Simpson of Sinton,
Texas, who generously placed at our disposal his truck, a four-wheel
drive “Land Rover,” without which travel on the island would have
been difficult. We also acknowledge a loan of field equipment
provided by Dr. Clarence Cottam, Director of the Welder Wildlife
Research Foundation, Sinton, Texas.
Financial support for the present research was provided by grants
from the National Science Foundation to The University of Texas
(G 15882) and to The University of Kansas (G 10048).
Permits to collect vertebrates in México were supplied by Ing.
Luis Macias Arellano, El Director General, Departamento de Con-
servacion de la Fauna Silves, México, D. F.
We are indebted to Dr. Richard H. Manville for arranging a loan
of specimens of Geomys personatus tropicalis in the United States
National Museum. Dr. Marshall Johnston kindly identified speci-
mens of plants from the barrier island. Several bones of birds and
mammals were identified by Dr. Pierce Brodkorb and Dr. E. L.
Lundelius. Mr. J. Knox Jones identified some of the mammalian
material, and Dr. W. E. Duellman verified the identifications of the
lizards; we thank all of these men for their willing assistance.
(311)
312 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
The Ecological Setting
The barrier island of Tamaulipas geologically and ecologically
resembles Padre Island, of the Gulf coast of lower Texas, north of
the mouth and delta of the Rio Grande. South of the delta, the
island in Tamaulipas is a narrow strip of sand less than a mile in
average width and is broken by a series of narrow inlets or “passes”
through which water from the Gulf of Mexico mingles with that
of the Laguna Madre de Tamaulipas. The passes are subject to
recurrent opening and closing. North of the mouth of the Rio Soto
la Marina, eight passes are designated by local fishermen, but only
three, the Third, Fourth, and Fifth, were open at the time of our
visit.
The Laguna Madre de Tamaulipas is described by Hildebrand
(1958) in connection with a preliminary study of the fishes and
invertebrates there. The average depth is probably less than 70 cm.
and the waters are hypersaline. In the time of the recent drought
in Texas and northeastern México, salinity varied from 108 to 117
parts per thousand in the northern part of the laguna near Arroyo
del Tigre (measurements taken in March, 1955) to from 39 to 48
parts per thousand in the southern part near Punta Piedras (meas-
urements taken in October and November, 1953, and in March,
1954). Discussions of the geologic history, ecology, and zoogeogra-
phy of the lagoons of the Gulf coast of the United States are given
by Hedgpeth (1947; 1953).
Localities in coastal Tamaulipas mentioned in the text of this
paper are shown on Plate 5.
The principal animal habitats are found in three vegetational
associations (plates 6 and 7). On flats and low dunes lying between,
and partly sheltered by, larger active dunes, small clumps of
Croton punctatus and a sedge (Fimbristylis castanea) are the only
conspicuous plants. Near the western edge of the dunes, Ipomoea
pescaprae var. emarginata is mixed with Croton, and there are scat-
tered clumps of shrubby wolf-berry (Lycium carolinianum var.
quadrifidum), and mesquite (Prosopis juliflora).
The dunes are relatively stabilized on the western side of the
island, and there we found moderately dense stands of mesquite
trees reaching heights of from eight to 10 feet. Prickly-pear cactus
(Opuntia lindheimeri) was common in those stands of mesquite,
and we saw an occasional yucca tree. A fairly dense ground cover
was formed by blanket-flower (Gaillardia pulchella), marsh-elder
(Iva sp.), Flaveria oppositifolia, Enstoma exaltatum, and Croton
capitatus var. albinoides.
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 313
A more open, xeric expression of the mesquite-cactus vegetation
occurs on exposed, low clay dunes (see description by Price, 1933)
located on alkaline flats bordering the laguna. At the time of our
visit, most of the mesquites in these stands were dead or dying, the
cactus was abundant, and the ground cover, which was sparse,
included drop-seed (Sporobolus virginicus), ragweed (Ambrosia
psilostachya), and Commicarpus scandens.
On alkaline flats flooded by hypersaline waters of the laguna fol-
lowing heavy rains, Batis maritima is found in the lower areas, but
on the slightly elevated areas there is low and almost continuous
cover of Monanthochloé littoralis, in which can be found Batis,
Borrichia fructescens, Salicornia sp., Iva sp., and sea-lavender (Li-
monium carolinianum).
Near Third Pass, sea oats (Uniola paniculata), evening primrose
(Oenothera sp.), and cordgrass (Spartina sp.) are present on the
dunes, and on alkaline flats we collected Conocarpus erectus, Leu-
caena sp., and Cassia fasciculata var. ferrisiae.
Itinerary
We reached Washington Beach from Matamoros on July 6, and
drove to a point approximately 33 miles south on the beach, where
we made Camp 1 on the east side of large dunes 400 yards from the
surf. From this camp we worked the beach and dunes and also
visited alkaline flats adjacent to the Laguna Madre. On the after-
noon of July 8, we drove south along the beach and established
Camp 2 on the south side of the Third Pass, approximately 73 miles
south of Washington Beach. We had intended to go farther south
but were unable to cross the Fourth Pass, an inlet three miles south
of the Third Pass. We left the barrier island on the afternoon of
July 10, after driving north from Camp 2 to the mouth of the Rio
Grande, 11 miles north of Washington Beach.
Mexican fishermen camped at the Fourth Pass told us that, had
we been able to cross the Fourth Pass, it would have been possible
to drive south on the beach all the way to La Pesca, a fishing village
near the mouth of the Rio Soto la Marina, approximately 150 miles
south of Washington Beach.
Summary of Previous Work in the Area
The ornithologist H. E. Dresser (1865-1866) worked in southern
Texas and at Matamoros, Tamaulipas, in 1863, and on one occasion
reached the mouth of the Rio Grande (“Boca Grande”). He did
not visit the barrier island or the Laguna Madre de Tamaulipas.
314 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
In their extensive travels through México, E. W. Nelson and E. A.
Goldman made collections at three localities in the coastal region
of Tamaulipas but did not reach the barrier island (Goldman, 1951).
Goldman collected at Altamira, near Tampico, from April 2 to 24,
1898, and from May 15 to 20 of the same year both he and Nelson
made headquarters at Altamira. Nelson and Goldman also collected
in the vicinity of Soto la Marina, 25 miles from the coast, from March
1 to 10, 1902, and, from February 18 to 15, they visited Bagdad,
described by Goldman (1951:260) as “a village at very low eleva-
tion on the Rio Grande about 6 miles above the mouth of the river.”
In March, 1950, C. von Wedel and E. R. Hall collected four species
of mammals and one bird on the barrier island at Boca Jésus Maria
(Eighth Pass). A report of this work published by Hall (1951)
contains descriptions of three new subspecies of mammals from the
island.
A few records of birds from the southern end of the barrier island
and from other parts of coastal Tamaulipas were reported by Robins,
Martin, and Heed (1951). In 1953, R. R. Graber and J. W. Graber
made ornithological studies in the vicinity of Tampico and also
reached the western edge of the Laguna Madre de Tamaulipas.
Several papers on this work have appeared (Graber and Graber,
1954a, 1954b; Graber, 1955), but a comprehensive account of their
observations and specimens was not published. Finally, J. R. Alcorn
collected some sandpipers 20 miles southeast of Matamoros, on
August 21, 1954, obtaining the first record of the Semipalmated
Sandpiper (Ereunetes pusillus) in Tamaulipas (Thompson, 1958).
Accounts of Species
Catalogue numbers in the following accounts are those of the
Museum of Natural History, The University of Kansas.
Reptiles
Gopherus berlandieri Agassiz: Texas Tortoise.—A pelvic girdle
and complete shell with a few attached scutes (63494) were found
in stabilized dunes at Camp 1 on July 7, and tracks were seen in
the same area. Fragments of two other shells (63493, 63495) were
found on sand flats between active dunes at Camp 1.
Holbrookia propinqua prepinqua Baird and Girard: Keeled Ear-
less Lizard.—This lizard was abundant on dunes and in pebble-
strewn blow-out areas between dunes at Camp 2, but it occurred
in smaller numbers in the less stabilized dunes of sparser vegetation
at Camp 1. Breeding was in progress at both localities, as evidenced
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 315
by the presence of eggs in the oviducts of several females, by the
heightened coloration of both sexes, and by mating behavior.
The mating behavior of this species has not been described in
the literature, and the following observations, made by Raun at
Camp 2 on July 8, may be of interest. A male was seen to circle
a female as the latter remained motionless with tail curved upward
and to the side, exposing a patch of bright pink-orange color on
the ventral surface of the tail. At times the male approached the
female from the rear and slightly to the side, biting the dorsal part
of her neck and simultaneously attempting to effect intromission.
The female several times reacted to this approach by running for-
ward a few steps, thereby freeing her neck from the grasp of the
male. When the male did not attempt to approach again, the
female appeared to invite copulation by moving in front of him
with tail elevated and the colored ventral surface prominently dis-
played. At the time of copulation, the male mounted from the rear
on the right side of the female, grasped her neck, and circled his
tail beneath her tail; at the same time the hindquarters of the female
were arched upward.
To confirm the presumed sexes of the two individuals under ob-
servation, both were collected while in copulation. Examination
of the still-coupled specimens showed that both hemipenes of the
male were everted and the left one had been inserted.
Apparently the pink-orange subcaudal patch of females is present
only in the mating season. It was not present on specimens of this
species taken by Raun and Wilks on Padre Island, Texas, in autumn,
and it is not mentioned in taxonomic descriptions by Axtell (1954)
and Smith (1946).
Measurements of adult specimens in our series indicate that
females are of smaller average size than males, and, as previously
noted by Smith (1946:132), females of this species have dispropor-
tionately shorter tails than do males (Table 1).
Holbrookia propinqua was previously collected on the barrier
island by Axtell (1954:31; see also Axtell and Wasserman, 1953:2),
who took specimens at Boca Jésus Maria, at a locality six to seven
miles south of Boca Jésus Maria, and at a point 20 miles east-south-
east of Matamoros. Axtell (Joc. cit.) also lists specimens in the
Museum of Zoology, University of Michigan, from Tepehuaje and
from one mile north of Miramar Beach (Tampico).
Specimens (56): 3 g g adult, 1 g subadult, 63433-436, Camp
1, July 7. 33 g g adult, 63437-440, 63443-445, 63447, 63448, 63450-
316 UNIVERSITY OF Kansas Pusxs., Mus. Nat. Hist.
TABLE 1.—MEASUREMENTS IN MILLIMETERS OF ADULT SPECIMENS OF
Holbrookia propinqua FROM THE BARRIER ISLAND OF TAMAULIPAS
Number Ratio:
Sex of Bho Eve nt Tail length snout-vent
specimens 8 to tail
Male...... 33 56.0+0.5* 77.0+0.7 0.731+0.001
(49-62) (69-85) (0.682-0.817)
Female.... 14 50.9+0.5 62.2+0.9 0.825+0.001
(47-53) (57-68) (0.735-0. 877)
* Mean + standard error; range indicated in parentheses.
456, 63458, 63460, 63462, 63468, 63465-468, 63470-478; 138 ¢ 9 adult,
63441, 63446, 63449, 63457, 63459, 63469, 63479-485; 6 juv., 63442,
63461, 63464, 63486-488; Camp 2, July 9-July 10.
Cnemidophorus gularis Baird and Girard: Whip-tailed Lizard —
At both camps we found this species in the same general habitat in
which Holbrookia occurred, but in numbers decidedly fewer than
the latter.
Specimens (4): 2 ¢ 9 adult, 63489, 63490, Camp 1, July 7.
1 g adult, 63491, 1 ¢ adult, 63492, Camp 2, July 9.
We failed to take specimens of snakes on the barrier island, but
tracks of snakes were noted on two occasions in dunes near Camp 1;
one trail led into a burrow of a kangaroo rat.
Birds
Unless otherwise indicated, specimens taken were not molting.
For birds undergoing postnuptial or postjuvenal molt, the degree
of advancement of the molt is indicated by recording the number
of primaries of the old plumage that have not been dropped. For
example, the designation “4 P old” signifies that all primaries except
the distal four have been molted.
Table 2 presents results of a strip census of birds along the strand,
made by three of us from the moving truck on the morning of July
10. Birds characteristically found on sand near the surf were thus
conveniently counted in accurate fashion. Birds not ordinarily found
on the strand could not be treated this way; most were considerably
less abundant than the eight most numerous species listed in Table 2.
Over-all, the numbers of individuals listed are a good index of
abundance of the Great Blue Heron and of the common charadrii-
form birds on the beach in early July. The Black Tern is an excep-
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 317
TABLE 2.—Brirps * RECORDED ALONG 17 MILEs OF BEACH } ON THE
BARRIER ISLAND OF TAMAULIPAS
SPECIES Number Birds per mile
GreatuBlue Heron os. csiis ye istis eae 9 0.5
Oveter-calenenes. bch. a) Paks tae.) baa, 1 0.1
Black-bellied'Plower:)5:fécc 2a hs ieselois SE 20 1.2
WilsonUPlover.. cso ti si ci a cusrs 53 3.1
Wallies iirc aetna moiehse ele aa etuess 43 2.5
Speenlerling Os ce tah. aud Al ho 5 ee 55 3.2
Baughing< Gulls seen sae ee eee 136 8.0
IRC KPPeR cht eect te eae eT ae 19 VA
@asmiane bern fonts cc acbeiscie ls Ete taka 82 4.8
eartylerns sassris Hie sce: AAS nae 221 13.0
Royalhlerrese: se sooth te eee ee 301 Wea?
Cabot UGnne ge ticcnae unst eared Va ee oe kel 122 ise
Total: 1062 Total: 62.4
* Common Tern, Forster Tern, and Long-billed Curlew also seen but not counted.
sean Between 56 and 73 miles south of Washington Beach, 11:00 to 11:45 a.m., July 10,
tion, however, and this is discussed in the account of that species
on page 327.
Pelecanus erythrorhynchus Gmelin: American White Pelican.—
A flock of approximately 800 individuals was seen resting at the
edge of the Laguna Madre near Camp 2 on July 9. When disturbed
by gunshots, the birds circled high over the laguna and flew to the
west. Among bones found on sand flats at Camp 1 are a left tarso-
metatarsus and a pedal phalanx of an American White Pelican.
Supposedly the only breeding colony of this species on the north-
ern Gulf coast is one in the Laguna Madre near Corpus Christi
(Peterson, 1960:8), but the possibility of one or more such colonies
existing in northeastern Tamaulipas has been suggested by Amadon
and Eckelberry (1955:68) on the basis of their observations of
individuals seen soaring near the coast 15 to 20 miles south of
Brownsville on April 15 and June 5, 1952. According to Hildebrand
(1958:153, and personal communication, August 14, 1961), small
318 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
colonies of white pelicans do breed in some years on two small
islands, in the Laguna Madre of Tamaulipas, located at 25° 26’ North
and 93° 30’ West.
In Veracruz the species is recorded as a winter visitant and tran-
sient (Loetscher, 1952:22; Amadon and Eckelberry, 1955:68). Cof-
fey (1960:289) reports the following observations for Veracruz and
Tamaulipas: a flock of 52 between Tlacotalpan and Alvarado, May
29, 1951; 80 near Cacaliloa, April 20, 1958; 180 birds north of Al-
varado, April 24, 1958; four at Altamira, May 28, 1955; flocks of
three, 13, and 87 “south” of Matamoros, May 20, 1951; 72 at Lomas
del Real, November 20, 1956.
Pelecanus occidentalis Gmelin: Brown Pelican——Three indi-
viduals flew north over the surf near Camp 1 on July 7, and a Jone
bird was seen diving into the Gulf a short distance beyond the surf
near Camp 2 on July 9. Birds seen by us probably were of the
population named P. o. carolinensis, which is resident along the Gulf
coast (Mexican Check-list, 1950:21).
Phalacrocorax sp.: Cormorant.—From 80 to 100 adult and ju-
venal cormorants were on the laguna at Camp 2 on July 8 and 9.
Probably they were Common Cormorants (P. olivaceus), but, be-
cause specimens were not taken, we cannot eliminate the possibility
that some (or all) were Double-crested Cormorants (P. auritus).
The former breeds in coastal lowlands of eastern México, whereas the
latter is known in eastern México only as a winter visitant and has
not been recorded in Tamaulipas (Mexican Check-list, 1950:24).
Fregata magnificens Mathews: Magnificent Man-o-war Bird.—
An observation of a lone bird circling high over the laguna at Camp
2 on July 9 seemingly constitutes the third record of this species in
Tamaulipas. Previous records were reported by Robins, Martin, and
Heed (1951:336), who found “large numbers” in the Barra Trinidad
region (8 miles north of Morén) on April 27 to 29, 1949, and men-
tioned an immature male taken at Tampico on April 23, 1923; this
specimen has been identified by P. Brodkorb as F. m. rothschildi.
Ardea herodias Linnaeus: Great Blue Heron.—Our records of
this heron are limited to the following observations: four individuals
on the beach and seven in the laguna at Camp 1, July 7; one on the
beach 52 miles south of Washington Beach, July 8; one 74 miles
south of Washington Beach, July 8; two at Third Pass, July 8; 41
standing on mud-flats at the edge of the laguna near Camp 2, July 9;
nine on the beach 56 to 73 miles south of Washington Beach, July 10;
one on the beach 42 miles south of Washington Beach, July 10.
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 319
The status of the Great Blue Heron in coastal Tamaulipas remains
to be determined. The subspecies A. h. wardi (considered a syno-
nym of A. h. occidentalis by Hellmayr and Conover, 1948) is resi-
dent and breeds on the Gulf coast of Texas and is to be expected as
a resident in Tamaulipas (Mexican Check-list, 1950:27). The
species may breed south to Veracruz, where Loetscher (1955:22)
reports it is “regular at nearly all seasons, chiefly on the coastal
plain”; he records an observation near Tamds on July 1. The sub-
species A. h. herodias and A. h. treganzai winter through much of
México and have been recorded in Tamaulipas (Mexican Check-list,
1950:27).
Florida caerulea (Linnaeus): Little Blue Heron—We saw a
white (immature) individual feeding with Reddish Egrets along
an inlet at Camp 2 on July 8.
Dichromanassa rufescens rufescens (Gmelin): Reddish Egret —
This egret was recorded only about the inlet at Camp 2, where 15
individuals were feeding, either singly or in small groups, on July 8
and 9. We noted frequent use of the “Open Wing” method of
foraging, as described by Meyerriecks (1960:108).
Specimen: 9 juv., 38899, ovary inactive, 587 gm., Camp 2, July
8. This specimen is referable to the nominate subspecies, which is
resident along the Gulf coast. Our record seems to be the first for
the species in Tamaulipas.
Leucophoyx thula (Molina): Snowy Egret—Ten individuals of
this species were feeding in association with Reddish Egrets in the
inlet at Camp 2 on July 9.
Hydranassa tricolor (P.L.S. Miller): Tricolored Heron.—An
observation of one individual flying along the margin of the laguna
near Camp 2 is our only record of this species.
Nycticorax nycticorax (Linnaeus): Black-crowned Night Heron.
—This heron was found only at the edge of the laguna near Camp 2;
ten individuals were noted on July 8, and 20 were seen perched in
a clump of mesquite trees on July 9. Perhaps half the birds seen
were in juvenal plumage. A juvenile was shot and examined on
July 9 but was not preserved as a specimen.
There appears to be no definite evidence of breeding by this
species in Tamaulipas (Mexican Check-list, 1950:32), but such may
be expected because the species breeds locally in Texas (Peterson,
1960:19) and in Veracruz.
Ajaia ajaja (Linnaeus): Roseate Spoonbill.—On July 9 at Camp
320 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
2, 38 spoonbills flew up from the edge of the laguna where they had
been resting near a large flock of white pelicans.
Cathartes aura (Linnaeus): Turkey Vulture-—One Turkey Vul-
ture was seen flying east at a point 2 miles west of Washington Beach
on July 10. It is noteworthy that we saw no Yellow-headed Vultures
(C. burrovianus), a species recently recorded in the region of
Tampico north to Lomas del Real (Graber and Graber, 1954a).
Colinus virginianus texanus (Lawrence): Bob-white.—This spe-
cies was seen only in or near clumps of mesquite near Camp 1, where
three covies (7, 13, and 18 individuals) were flushed on July 7.
Specimen: ¢ juv., 38900, testis 3 mm., 100 gm., 6 P old, Camp 1,
July 7.
Porzana carolina (Linnaeus): Sora Rail.—On sand flats at Camp
1 we found a left humerus and several other post-cranial skeletal
elements that have been identified by Dr. Pierce Brodkorb as be-
longing to this species. All the bones are of Recent age. We have
no other record of the Sora Rail on the barrier island, but in all
probability it occurs as a migrant and winter visitant along margins
of the laguna.
Haematopus ostralegus Linnaeus: Oyster-catcher——One _indi-
vidual was seen at Camp 2 on July 8, three were noted at the same
locality on July 9, and one was present on the beach 72 miles south
of Washington Beach on July 10. The only previous records of this
species in Tamaulipas are a specimen ( 7, 29348) taken by E. R.
Hall 10 miles west and 88 miles south of Matamoros on March 20,
1950 (herewith reported for the first time), and three seen on the
beach near Tepehuaje on May 9, 1949 (Robins, Martin, and Heed,
1951).
Squatarola squatarola (Linnaeus): Black-bellied Plover.—Plov-
ers of this species were uncommon but regular on the beach; fre-
quently two individuals were seen together, sometimes in associa-
tion with one or more Willets. Specimens (4): ¢, 38915, testis 4
mm., 231 gm.; 3g, 38914, testis 4 mm., 221 gm.; ¢, 38916, testis 3
mm., 209 gm., Camp 1, July 7. Male, 38917, testis 4 mm., 186 gm.,
Camp 2, July 9. The specimens were molting (3-4 P old) into win-
ter plumage and showed little or no subcutaneous fat.
Our specimens and records probably pertain to nonbreeding in-
dividuals summering on the coast, as the species is known to do in
Texas (Hagar and Packard, 1952:9) and elsewhere in its range
(Eisenmann, 1951:182; Haverschmidt, 1955:336; A.O.U. Check-list,
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 321
1957:174). In any event, our dates (July 6 to 10) are unusually
early for autumnal migrants; they do not reach Texas until August
(Peterson, 1960:94), and Loetscher (1955:26) gives August 7 as the
earliest date for southbound migrants in Veracruz.
Charadrius hiaticula semipalmatus Bonaparte: Ringed Plover.—
We have a single record, an adult male (38913, testis 2x1 mm.,
heavy fat, 47.0 gm., 4 P old) taken on a sandbar at Camp 2 on July 9.
The bird was feeding in company with a flock of Sanderlings.
There is no previous record of the Ringed Plover in Tamaulipas.
In Texas, Hagar and Packard (1952:8) indicate that the first au-
tumnal migrants reach the central Gulf coast in the last week of
July. In coastal México, the species has previously been recorded
from August 23 to May 12 (Mexican Check-list, 1950:91). There-
fore, the present record must represent an exceptionally early south-
bound migrant, or, more probably, a nonbreeding, summering
individual. According to the A.O.U. Check-list (1957:166), non-
breeding birds are found in summer in coastal areas south to Cali-
fornia, Panama, and Florida. Many individuals spend the northern
summer along the coast of Surinam (Haverschmidt, 1955:336).
Charadrius wilsonia wilsonia Ord: Wilson Plover.—This small
plover breeds commonly on the beach and on alkaline flats adjacent
to the laguna. Previous evidence of breeding in Tamaulipas con-
sisted only of a report of a male with brood patches and an enlarged
testis taken near Tamés on May 30, 1947 ( Loetscher, 1955:26).
We saw many pairs of adults and a large number of well-grown
juveniles, and, at a point 4 miles south of Washington Beach, we
collected a brood of three small juveniles that had only recently
hatched. The breeding season apparently was drawing to a close,
for several adults in our collection were in postnuptial molt and
showed marked gonadal regression. From July 6 to 9, a few small
groups of birds were noted, but large groups were not seen until
July 10, when several flocks of up to 60 individuals were found along
the coast 3 to 7 miles south of Washington Beach.
Specimens (12): 3, 38904, testis 4.5 x 2 mm., 58 gm., 3 P old,
brood patches refeathering; ¢ , 38905, testis 5 x 2 mm., 59 gm., 4 P
old, brood patches refeathering; ¢ juv., 38903, 6.2 gm.; 2 sex?,
38901, 38902, 5.7 and 6.2 gm., 4 miles south of Washington Beach,
July 6. Male, 38907, testis 5 x 2 mm., 56 gm., 7 P old, brood patches
refeathering; 9 , 38906, ova to 1 mm., 61 gm., 3 P old, brood patches
refeathering; 9 juv., 38908, ovary inactive, 54 gm., in body molt;
Camp lI, July 6. Male, 38910, testis 6x 3 mm., 60 gm., 4 P old; 9,
322 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
38909, ova to 1 mm., 57 gm., 4 P old, brood patches refeathering;
Camp 1, July 8. Male, 38911, testis 2x 1 mm., 55 gm.; juv., 38912,
no weight or sex recorded; Camp 2, July 9.
Numenius americanus parvus Bishop: Long-billed Curlew.—
Lone individuals and groups of two to five were noted occasionally
along the beach each day. In total, some 30 to 50 birds were
counted, but some individuals may have been recorded more than
once on different days. Specimens (2): ¢, 38918, testis 4 mm.,
some fat, 459 gm., Camp 2, July 9; ¢, 38933, ova to 1 mm., no
weight recorded, Camp 2, July 8.
Our assumption that some or all individuals seen by us were non-
breeding, summering birds is supported by the fact that our speci-
mens are referable to the small, northwestern subspecies, N. a. par-
vus, rather than to N. a. americanus; the latter breeds south in the
eastern United States to south-central Texas (A.O.U. Check-list,
1957:181). Loetscher (1955:27) saw a flock of 39 curlews near
Tamdés on June 30, and he notes that nonbreeding birds are fairly
common at all seasons in Veracruz. Similarly, the species is present
throughout the year on the central Gulf coast of Texas (Hagar and
Packard, 1952:8). Authors of the Mexican Check-list (1950:94)
do not mention the possibility that birds of this species recorded
in México in July are summering rather than migrating. Twelve
supposed migrants seen along Laguna Chila (Cacalilao), Veracruz,
by Coffey (1960:291) on May 81, 1957, may have been summering
birds.
Limosa fedoa (Linnaeus): Marbled Godwit—Three were seen
in shallow waters of the laguna at Camp 2 on July 9. Specimen:
3 , 88919, testis 6 x 2 mm., fat, 805 gm., 6 P old, Camp 2, July 9.
Probably our records were of nonbreeding birds, which are known
to occur in summer elsewhere in México (Mexican Check-list, 1950:
94), sparingly in Texas (Hagar and Packard, 1952:8), and in South
Carolina (A.O.U. Check-list, 1957:205). Apparently the only record
for this species in Veracruz is one seen on May 11, 1954, east of
Cacalilao (Coffey, 1960:292).
Tringa melanoleuca (Gmelin): Greater Yellowlegs.—Three
birds were seen on alkaline flats at Camp 1 on July 7, and two were
noted at Camp 2 on July 9. There is one previous report of this
species in Tamaulipas, and, since it has been recorded as a migrant
and winter resident in México between July 26 and April 26 ( Mexi-
can Check-list, 1950:95), our records seem to pertain to unusually
early autumnal migrants or, possibly, to nonbreeding, summering
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 323
birds. Other mid-summer records are available from Tamds on
June 80 and July 1, and the species is “to be expected every month
of the year” in Veracruz (Loetscher, 1955:27). Sight records for
Veracruz in May (Coffey, 1960:291) may well pertain to summering
birds. There are northern-summer records for this species from
Texas (Hagar and Packard, 1952:8), Surinam (Haverschmidt, 1955:
367), and other areas within the winter range of this yellowlegs
(A.O.U. Check-list, 1957:190).
Catoptrophorus semipalmatus semipalmatus Gmelin: Willet—
The Willet was common on the island. We found evidence of breed-
ing and also saw large flocks of birds that were either nonbreeders
summering in the area or early, postbreeding migrants from more
northerly places. All along the beach and at the edge of the laguna
at both camps we found Willets in twos or threes, often accompanied
by one or two Black-bellied Plovers. On July 10 a small juvenile
was captured; two adults in breeding plumage evidenced obvious
concern at this action. On July 6 a flock of 30 birds flew east over
Camp 1, and a flock of 90 was seen flying south over Camp 1 on
July 7.
Specimens (7): ¢, 38922, testis 6 x 1 mm., 264 gm., breeding
plumage; 9° , 38923, ova to 2 mm., 269 gm., breeding plumage; ° ,
38924, ova to 1 mm., 280 gm., 3 P old; ¢ , 38925, ova to 1 mm., 319
gm.; ¢, 38921, testis 7x 2 mm., 211 gm., breeding plumage; Camp
1, July 7. Male, 38927, fat light, 231 gm., 4 P old, Camp 2, July 9.
Juvenile, sex not recorded, 38920, 43.0 gm., 1 mile south of Washing-
ton Beach, July 10. Two of our specimens, both males, are in worn
breeding plumage and evidence no molt; another specimen, a fe-
male, is also in breeding plumage but is molting on the breast. The
remaining two adult skins in our series are three-quarters through
the molt and are for the most part in fresh winter feather.
Dresser (1866:37) took an unspecified number of specimens of
the Willet at the “Boca Grande” in July and August, but actual
breeding in Tamaulipas was first established by C. R. Robins, who
found a “scattered colony of breeding Willets” and took a female
with an egg in the oviduct on May 9, 1949, near Tepehuaje (Sutton,
1950:1385). Sutton (op. cit.) has discussed the characters of this
specimen and of birds from Cameron County, Texas. The specimen
from Tepehuaje reportedly is closer to C. s. inornatus than to C. s.
semipalmatus both in size and color, and birds from Cameron
County are intermediate between the two subspecies in size but like
C. s. inornatus in color.
324 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
Measurements of our five adults from the barrier island are pre-
sented in Table 3 for comparison with those of C. s. semipalmatus
and C. s. inornatus given by Ridgway (1919:316-319). Like the
TABLE 3.—MEASUREMENTS IN MILLIMETERS OF SPECIMENS OF CATOPTROPHORUS
SEMIPALMATUS FROM THE BARRIER ISLAND OF TAMAULIPAS
SEX AND Full Weight
CaTALOGUE Wing Tail culmen Tarsus in
NUMBER grams
(apples 12774) al ae Oe 197 80.6 61.0 59.0 211
DOOD ese) Soh euros 198 74.4 61.9 57.9 264
SSeS Za ame 194 75.5 60.4 56.4 231
OMBOOZO Tas waa ct a6 201 71.0 59.0 55.4 269
Oi SSO24 he =. Socks. 199 71.0 61.3 59.0 280
* Specimens in worn breeding plumage.
specimens from Cameron County examined by Sutton (op. cit.), our
birds are intermediate in size between average-sized individuals of
the two named subspecies. In color and pattern, we find that our
specimens in breeding plumage fall within the range of variation
of C. s. semipalmatus as exemplified by five specimens in nearly
identical states of wear and fading in the Museum of Natural
History.
On the basis of the evidence presently available, we are reluctant
to follow Sutton (1950:186) in assigning breeding birds from the
Gulf coastal region to C. s. inornatus, a name otherwise applied to a
population of birds breeding inland, in northwestern North America
south to central Utah and Colorado and east to South Dakota (and
formerly to western and southeastern Minnesota and Iowa; see
A.O.U. Check-list, 1957:190). The intermediate characters of birds
breeding in coastal Texas and Tamaulipas probably represent not
the results of actual genetic intermixing of the two named popula-
tions but, rather, an adaptive response of the eastern coastal stock
(C. s. semipalmatus ) to environmental modalities distinct from those
operating elsewhere within the range of the eastern coastal popula-
tion or on the inland population. Accordingly, we tentatively use
the name C. s. semipalmatus for our Tamaulipan specimens, realiz-
ing that the patterns of geographic variation in the species do not
lend themselves well to taxonomic treatment by the trinomial no-
PLATE 5
BROWNSVILLE
MATAMOROS ae
Tamaulipas
Zamorina
Camp 2
Boca Santa Marla
(Fourth Pass)
Barra
Trinidad
La Carbonera
Boca Jesus Maria
(Eighth Pass)
Miramar
10) 10
Statute Miles
Map of coastal Tamaulipas, showing the barrier island and localities men-
tioned in text. Stippled areas are extensively marshy.
PLADEG
Fic. 1.—Croton and Fimbristylis on stabilized dunes; the Laguna Madre and
surrounding alkaline flats and clay dunes are visible in the background.
Habitat of Road-runner, Ord kangaroo rat, and keeled lizard.
-
¢
Fic. 2.—Active dune near Camp 1. Other active dunes can be seen in the
background, in the right foreground is a clump of Croton, and in the left
foreground is a small clump of Fimbristylis. Habitat of Road-runner, Ord
kangaroo rat, and keeled lizard.
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 325
menclatural system. The need for a comprehensive analysis of
geographic variation in this species, based, if possible, on proper
segregation of age classes along the lines followed by Pitelka (1950)
for Limnodromus, is obviously indicated.
Arenaria interpres morinella (Linnaeus): Turnstone.—Approxi-
mately 40 individuals were noted along the beach from July 6 to 10,
mostly in small groups; the largest flock included 15 individuals.
Specimens (5): ¢, 38931, testis 4 x 1 mm., moderately fat, 107
gm.,4 Pold; ¢, 38932, testis 3 x 1 mm., moderately fat, 103 gm.,
molting; 75 miles south of Washington Beach, July 8. Male, 38928,
testis 2 mm., 111 gm., 3 P old; ¢, 38929, testis 3 mm., moderately
fat, 106 gm., 6 P old; ¢ , 38930, testis 2.5 mm., moderately fat, 108
gm., 6 P old; Camp 2, July 9.
The only previous record of the Turnstone in Tamaulipas is an
observation of an unspecified number at Tepehuaje on May 9, 1949
(Robins, Martin, and Heed, 1951). The dates of our records suggest
that nonbreeding birds summer along the coast of Tamaulipas. The
species is present in small numbers in summer along the central!
Gulf coast of Texas (Hagar and Packard, 1952:8). Loetscher (1955:
26-27) does not report records for Veracruz in summer, but records
of the species in Yucatan on May 31, 1952 (Paynter, 1955:101), and
on June 16, 1900 ( Mexican Check-list, 1950:79), probably represent
summering nonbreeders. Probably also in the same class are sup-
posed “migrants” seen at Coatzacoalcos on May 17, 1954, and June
4, 1955 (Coffey, 1960:290).
Inasmuch as Haverschmidt (1955:368) reports that nonbreeding
birds summering in Surinam only occasionally assume breeding
plumage, it is noteworthy that our specimens were molting from
nuptial (summer) to winter plumage. None of the nonbreeding
northern shorebirds observed by Eisenmann (1951:183) in Panama
in summer were in nuptial plumage.
Crocethia alba (Pallas): Sanderling.—This sandpiper was noted
each day along the beach, occasionally singly but more frequently
in groups ranging from 10 to 50 individuals. Specimens (7): 7,
38936, testis 2 mm., light fat, 49 gm., 5 P old, Camp 1, July 7.
Female, 38937, ova to 1 mm., fat, 58 gm., 4 P old; 3g, 38939, fat,
no weight recorded, 6 P old, breeding plumage; 3 ¢ 3, 38940-
38942, fat, no weight recorded, 4-5 P old; Camp 2, July 9.
With one exception as noted, our specimens are in worn, non-
breeding plumage and are replacing their old feathers with new
ones fundamentally the same in color and pattern; the exceptional
2—3002
326 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
specimen is molting from worn breeding plumage into nonbreeding
plumage. Only one other individual in breeding feather was seen
on the island.
According to the Mexican Check-list (1950:99), the Sanderling
has been recorded in México from August to May 19. In Texas,
Peterson (1960:107) reports that it is a migrant, April to June and
July to November, and that it winters along the coast. We suspect
that many of the birds present in Texas in June and July, together
with those recorded by us in Tamaulipas in July, are nonbreeding,
summering individuals. Haverschmidt (1955:368) reports northern-
summer records from Surinam, and, according to the A.O.U. Check-
list (1957:208), nonbreeding birds occur in summer extensively
through winter range of the species, including the Gulf coast of the
United States.
Micropalama himantopus (Bonaparte): Stilt Sandpiper—Two
birds in worn winter plumage were taken as they foraged together
at the edge of the laguna near Camp 2 on July 9. Specimens (2):
gS , 38934, testis 2.5 mm., heavy fat, 116 gm., 4 P old; ¢, 38935,
testis 3 mm., fat, 111 gm., 4 P old.
Our specimens probably were nonbreeding birds summering be-
tween the breeding range in arctic America and the winter range
in northern South America. The A.O.U. Check-list (1957:202) does
not mention nonbreeding, summering records of this species. The
251 birds seen by Coffey (1960:292) at Cacalilao, Veracruz, on May
11, 1954, were probably migrants.
Recurvirostra americana Gmelin: American Avocet.—This spe-
cies was seen only in three large flocks flying south along the beach.
as follows: 56 birds 72 miles south of Washington Beach, July §;
38 birds 73 miles south of Washington Beach, July 8; 29 birds 72
miles south of Washington Beach, July 10. All birds were in winter
plumage.
All these birds were possibly autumnal migrants, but the dates
are early; the species has not previously been recorded on migration
in México before August (Mexican Check-list, 1950:101). The spe-
cies is known to breed in San Luis Potosi (Mexican Check-list, loc.
cit.) and along the lower coast of Texas (“rarely to Brownsville’;
A.O.U. Check-list, 1957:209); avocets thus may also breed in coastal
Tamaulipas.
Larus argentatus Pontoppidan: Herring Gull.—A first-year bird
was observed near Camp 2 on July 8, and two subadult individuals
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 327
were seen on the beach between the Third and Fourth passes on
July 8.
Larus atricilla Linnaeus: Laughing Gull.—This gull was common
all along the beach. Many individuals were in full breeding feather
and many subadult birds were also present. Specimens (6): 2
subadult, 38944, testis 5 x 1 mm., 325 gm., molting; 9° , 38945, ovary
small, 309 gm., in molt, brood patches refeathering; sex?, 38943, 315
gm., in molt; sex? subadult, 38946, 327 gm., in molt; Camp 1, July 7.
Female subadult (second-year), 38947, 305 gm., in molt, Camp 2,
July 8. Female, 38926, ova to 2.5 mm., 313 gm., 8 P old, Camp 2,
July 10.
The Mexican Check-list (1950:105) refers to the Laughing Gull
as a common winter resident on both coasts of México from August
7 to May 17, but Loetscher (1955:29) found it locally common
throughout the year on the coast of Veracruz, and he mentioned
seeing birds a short distance south of Tampico in June and July.
The status of this gull in Tamaulipas remains to be determined:
probably it will be found breeding locally, but many of the birds
summering in eastern México are most likely nonbreeders (A.O.U.
Check-list, 1957:226).
Chlidonias niger surinamensis (Gmelin): Black Tern.—On July
6, 7, 8, 9, and on the morning of July 10, we saw this species only
occasionally, recording in total not more than 50 individuals. But,
about noon on July 10, we observed at least 300 birds in compact
flocks of about 50 individuals each between Washington Beach and
a point about 9 miles south of that locality. Approximately one in
ten birds seen was in breeding plumage, the rest being in winter
or subadult plumages, which are indistinguishable in the field. Per-
haps some of the birds seen were nonbreeding, summering indi-
viduals, but we presume that the large groups were southbound
migrants, and we note that autumnal migrants appear in northern
Veracruz as early as July 1 (Loetscher, 1955:30). On the central
Gulf coast of Texas, Hagar and Packard (1952:9) indicate that an
influx of birds occurs in the last week of July, and small numbers
of birds, presumably nonbreeding individuals, are present along the
Gulf coast throughout June and July. Dresser (1866:45) found this
species to be “common at the Boca Grande during the summer.”
Specimens (2): ¢ , 38948, testis 6 mm., moderately fat, 68 gm.,
in breeding plumage, Camp 1, July 7. Female, 38949, ovary in-
active, 49 gm., molt into winter feather almost complete, Camp 2,
July 10.
328 UNIVERSITY OF KANSAS Pus.s., Mus. Nat. Hist.
Hydroprogne caspia (Pallas): Caspian Tern.—The only pub-
lished record of the Caspian Tern in Tamaulipas is a report of one
seen at Lomas del Real on November 20, 1956 (Coffey, 1960:260),
but we found it moderately common all along the beach and at the
margin of the laguna. It was frequently associated with the Royal
Tern, which outnumbered it better than three to one (see Table 2).
The species is resident and breeds along the coast of Texas, and it
probably has similar status in Tamaulipas. However, in Vera-
cruz it is known only as a winter visitant (Loetscher, 1955:30) and
as a spring migrant (Coffey, 1960:293). Specimen: ¢ , 38950, ova
to 2 mm., moderately fat, weight not recorded, 5 P old, Camp 2,
July 9.
Sterna hirundo hirundo Linnaeus: Common Tern.—We took a
specimen ( ¢ ?, 38951, no fat, 165 gm.), 49 miles south of Washing-
ton Beach on July 8, and saw two others over the laguna at Camp 2
on July 9. Our specimen had nearly finished with molt and feather
growth into adult winter plumage. The status of Common Terns
in Tamaulipas is uncertain; our record, and records from Tamés on
July 1, 1952, and June 12, 1953 (Loetscher, 1955:29), probably per-
tain to nonbreeding, summering birds. Yet, the species has bred
on the Texas Gulf coast (A.O.U. Check-list, 1957:235), and it rea-
sonably may be expected to nest in Tamaulipas. Coffey (1960:293 )
saw two individuals at Altamira on May 10, 1954.
Sterna forsteri Nuttall: Forster Tern.—Six were recorded near
Camp 1 on July 7, and two were seen on the beach on July 6 and 10.
The Mexican Check-list (1950:108) does not cite records for
Tamaulipas, but the A.O.U. Check-list (1957:234) includes northern
Tamaulipas within the breeding range. Evidence suggesting breed-
ing of the species in extreme northern Veracruz is reported by
Loetscher (1955:29) in the form of a female specimen with “ovary
greatly enlarged” taken seven miles west of Tampico on May 80,
1947. In the same area the species also seems to spend the summer
as a nonbreeder, for Loetscher (loc. cit.) saw 20, nearly all in non-
breeding plumage, on July 1, 1952.
Specimens (4): 2, 38952, testis 4.5 mm., 150 gm., 8 P old; ¢,
38955, testis 2 mm., 138 gm., 2 Pold; ¢ , 38953, testis 5 x 1 mm., 142
gm., 5 P old; 9° , 38954, ova to 1 mm., 148 gm., 2 P old; Camp 1,
July 7.
Sterna albifrons antillarum (Lesson): Least Tern.—The status
of this species in Tamaulipas is uncertain, but there is reason to
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 329
believe that it breeds, at least in small numbers. We found the
species moderately common and generally flying about in twos,
possibly mated pairs, near both camps and on the beach. Breeding
is suggested by the large sizes of the testes of the two males collected
and by the presence of brood patches on a female taken on July 6,
but we have no direct evidence of nesting in Tamaulipas, and it
should be noted that this species is known to spend the summer in
nonbreeding condition at many places (A.O.U. Check-list, 1957:
239). Loetscher (1955:30) suggests that the species may be found
breeding in Veracruz and mentions a record of 15 seen at Miramar,
Tamaulipas, on June 26, 1952. Dresser (1866:45) found it to be
“abundant” at the “Boca Grande” in summer.
On July 10, we saw flocks of 15 to 20 individuals flying along the
beach a few miles south of Washington Beach.
Specimens (4): 2, 38958, testis 11 x 4 mm. (right testis 5 x 4
mm. ), light fat, 45 gm., 6 P old; ¢ , 38959, testis 11 x 4 mm. (right
testis 7 x 4 mm.), light fat, 45 gm., 6 P old; @ , 38956, ova to 2.5
mm., 42.5 gm., 6 P old, brood patches refeathering; Camp 1, July 6.
Female, 38957, ova to 1 mm., 44 gm., Camp 1, July 7. This last
specimen had essentially completed the autumnal molt into winter
plumage, with only a few feathers remaining ensheathed basally.
Our specimens are referable to S. a. antillarum, being paler dor-
sally and slightly lighter gray on the hind-neck than specimens of
S. a. athalassos from Kansas, with which they were compared.
Thalasseus maximus maximus (Boddaert): Royal Tern.—This
species was common all along the beach, occurring for the most
part in flocks of from ten to 50 individuals in association with Cabot
Terns.. Data on gonadal condition and brood patches of some of
our specimens suggest that breeding occurs in coastal Tamaulipas, as
previously reported by the Mexican Check-list (1950:110). Robins,
Martin, and Heed (1951) report seeing one Royal Tern near Te-
pehuaje on May 9, 1949, and Dresser (1866:44) found the species
“common at the Boca del Rio Grande during the summer.”
Specimens (6): ¢, 38960, testis 9 x 4.5 mm., not fat, 484 gm., 6
P old, brood patches refeathering, 4 miles south of Washington
Beach, July 6. Male, 38961, testis 7 x 3 mm., 455 gm., no brood
patches, 8 miles south of Washington Beach, July 6. Male, 38962,
testis 10 x 5 mm., 387 gm., brood patches refeathering; 9 , 38963,
ova to 1 mm., 358 gm., 3 P old; 9? , 38964, ova to 3 mm., 389 gm.,
8 P old; Camp 1, July 7. Female, 38994, ova to 2 mm., 5386 gm.,
brood patches refeathering, Camp 2, July 10.
330 University oF KAnsaAs Pusts., Mus. Nar. Hist.
Thalasseus sandvicensis acuflavidus (Cabot): Cabot Tern.—This
tern was moderately common along the beach and margin of the
laguna, and it was seen frequently in company with Royal Terns.
Like the latter, this tern breeds in coastal Texas (A.O.U. Check-list,
1957:241), and it probably also nests in Tamaulipas, although direct
evidence is not available. The only previous record of this species
in Tamaulipas is a report (Robins, Martin, and Heed, 1951) of two
observed on the beach near Tepehuaje on May 9, 1949.
Specimens (4): ¢, 38965, testis 9 x 4.5 mm., 208 gm., 9 P old,
49 miles south of Washington Beach, July 8. Male, 38966, testis
8 x 3 mm., not fat, 192 gm., 8 P old; ¢ , 38967, ova to 3 mm., 193
gm., 7 P old, brood patches refeathering; @ , 38968, ova to 1 mm.,
186 gm., 8 P old, no brood patches; 52 miles south of Washington
Beach, July 8.
Rynchops nigra nigra Linnaeus: Black Skimmer.—We found this
species moderately common at the edge of the laguna at both camps
and occasionally saw it along the beach. Generally two birds, prob-
ably mated pairs, were seen together; twice birds were seen carrying
food in their bills, presumably intended for nestlings. The species
is known to nest in Tamaulipas from “Matamoros Lagoon” south to
Tampico (Mexican Check-list, 1950:112).
Specimens (2): ¢, 38970, testis 40 x 23 mm. (abnormally large,
possibly as a result of hemorrhage), 418 gm., brood patches re-
feathering; ¢ , 38969, testis 17 x 4 mm., fat light, 442 gm., brood
patches refeathering; Camp 1, July 7.
Zenaidura macroura Linnaeus: Mourning Dove.—Our only rec-
ord is a lone bird seen in a mesquite near Camp | on July 6. Possibly
the species breeds along the margin of the laguna, although Aldrich
and Duvall (1958:118, map) do not include coastal Tamaulipas in
the known breeding range. Loetscher (1955:30) suggests that the
Mourning Dove may be found breeding in the lowlands of northern
Veracruz and cites a record of one seen at Tamds on July 1, 1952.
Geococcyx californianus (Lesson): Road-runner.—At least four
individuals were seen in ]arge dunes at Camp 1 on July 7 and 8.
On several occasions we watched them pursue lizards (Holbrookia
propinqua) at the margins of clumps of Croton and Ipomoea.
Chordeiles minor aserriensis Cherrie: Nighthawk.—Nighthawks
of this species were seen regularly at Camp 1, where we flushed
them from alkaline flats in the day and heard them calling as they
foraged over the dunes in late afternoon.
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 831
Specimens (3): 2 , 38971, testis 5 mm., no fat, 62 gm., Camp 1,
July 6. Male, 38972, testis 7.5 mm., no fat, 58 gm.; ¢ , 38973, testis
?, no fat, 53 gm.; Camp 1, July 7. The gonads of these birds were
not in full breeding condition, but it is highly probable that the
birds were members of a population that had bred in the area.
Variation in Chordeiles minor in Tamaulipas has recently been
studied by Graber (1955). Two specimens taken by him on August
3, 1953, approximately 9 miles south of Carbonera, resemble birds
from Terrell County, Texas, and represent C. m. aserriensis, as do
our three birds from the barrier island. Two of Graber’s specimens
from Lomas del Real, in southeastern Tamaulipas, are distinctly
darker and probably represent C. m. néotropicalis, a subspecies sub-
sequently described from Chiapas (Selander and Alvarez del Toro,
1955).
Muscivora forficata (Gmelin): Scissor-tailed Flycatcher.—On
July 7 near Camp 1, two individuals were found in stands of mes-
quite. One was taken and proved to be an adult male (38974, testis
6 x 3 mm., not fat, 40 gm.) in postnuptial molt (6 P old).
We presume that the two birds recorded by us were members of
a population breeding on the barrier island, rather than autumnal
migrants. The Mexican Check-list (1957:69) records this species
in México only as a transient and winter visitant. But, on the basis
of records of birds seen along the highway between Matamoros and
Ciudad Victoria, Davis (1950) has suggested that the species breeds
in Tamaulipas, and this is supported by a report of one seen at the
north end of the Monterrey Airport on June 1, 1957 (Coffey, 1960:
294). Brown (1958) has recently established that the species breeds
in Nuevo Leon by finding a nest 33 kilometers (by road) north of
Sabinas, Hidalgo, on July 19, 1954.
Myiarchus cinerascens cinerascens (Lawrence): Ash-throated
Flycatcher.—A juvenal male (38975, testis 2 mm., no fat, 35.0 gm.)
taken in mesquite at Camp 1 constitutes our only record for this
species. Lanyon (1961:441, map) has shown that most of Tamauli-
pas is devoid of these flycatchers in the breeding season; the nearest
known breeding Ash-throated Flycatchers are slightly west of
Corpus Christi, Texas, about 200 miles north-northwest of Camp 1
on the barrier beach. Our specimen closely resembles eight speci-
mens from Coahuila, México, in general coloration and, especially,
in the pattern of colors on the outer rectrices. Probably No. 38975
was from southwestern Texas or Coahuila and had begun its south-
ward migration. Against this idea lies chiefly the fact that young-
332 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
of-the-year tend to move south later than adults of the same species;
so, this bird possibly had been reared in coastal Tamaulipas.
Eremophila alpestris giraudi (Henshaw): Horned Lark.—This
species occurred in moderate numbers on alkaline flats and almost
barren sand flats at both camps. At the time of our visit to the
island, the breeding season apparently was coming to an end, but
we noted no tendency in the birds to flock.
Specimens (7): 2, 38981, testis 6 mm., 21.0 gm.; ¢, 38977,
testis 7.5 x 4mm., not fat, 27.5 gm.; ¢ , 38979, testis 11 x 7 mm., 29.0
gm.; 9 , 38976, ova to 3 mm., brood patch vascular but regressing,
no fat, 24.4 gm.; sex? juv., 38987, no fat, 21.0 gm.; sex? juv., 38980,
24.0 gm.; Camp 1, July 7. Male, 38982, testis 9.5 x 6 mm., 27.5 gm.,
Camp 2, July 9.
The subspecies E. a. giraudi, which is endemic to the Gulf coastal
plain of Texas and Tamaulipas, has been reported in Tamaulipas
previously only from Bagdad, near Matamoros (Mexican Check-list,
1957:106). The fact that our specimens show characters totally
consistent with those of E. a. giraudi indicates that there is little
genetic interchange between the population we sampled and those
of E. a. diaphora, the closest of which reportedly breeds at Miqui-
hana, in southwestern Tamaulipas.
Corvus cryptoleucus Couch: White-necked Raven.—Several
groups of six to ten birds were present at Washington Beach on July
6 and 10; but, southward on the island, we recorded this species only
once, on July 9, when a lone individual flew near Camp 2, being
pursued and “buzzed” by two Least Terns. The Mexican Crow
(Corvus imparatus) reportedly is common in the coastal region of
Tamaulipas (Mexican Check-list, 1957:118) but was not seen by us.
Thryomanes bewickii cryptus Oberholser: Bewick Wren.—This
species seemingly breeds in small numbers in mesquite stands near
Camp 1, where we obtained a juvenile and saw another individual.
Specimen: @ juv., 38983, no fat, 10.0 gm., Camp 1, July 8. T. b.
cryptus is reported to intergrade with T. b. murinus of Veracruz in
southern Tamaulipas (Mexican Check-list, 1957:160-161 ).
Mimus polyglottos leucopterus (Vigors): Northern Mockingbird.
—We recorded this species only near Camp 1, where a few pairs
were breeding in stands of mesquite. Males were in full song and
territorial display.
Specimens (2): ¢ , 38985, testis 11 x 7 mm., not fat, 48 gm.; 9°,
38984, ova to 4.5 mm., vascular brood patch, 49.0 gm.; Camp 1,
July 7.
PLATE 7
Fic. 1.—Mesquite-cactus formation on clay dune at margin of the Laguna
Madre west of Camp 1. Habitat of Northern Mockingbird, Cardinal, Bob-
white, black-tailed jackrabbit, and Great Plains woodrat.
x
Fic. 2.—Batis-Monanthochloé formation on alkaline flats near the Laguna
Madre, with mesquite bordering stabilized dunes in the left background.
Salicornia, a classical dominant of salt marshes, is here relatively inconspicu-
ous. Habitat of Nighthawk and Horned Lark.
PLATE 8
“Fossilized” burrow of Texas Pocket Gopher in a sandy trough between active
dunes. A part of the cast has been broken away to show the general shape
of the old burrow. The diameter of the cast is about 3.5 inches.
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 333
Cassidix mexicanus prosopidicola Lowery: Great-tailed Grackle.
—Small, postbreeding flocks composed of both adult and juvenal
birds were seen moving along the edge of the laguna at Camp 1.
In the morning the flocks flew south, and in the afternoon groups
of similar size flew north, presumably to a roost at an undetermined
distance north of our camp. Occasionally, a few birds stopped to
rest or to forage on the dunes or in stands of mesquite. At Camp 2
on July 9, a postbreeding adult female and a well-grown, presumably
independent juvenile were taken as they perched in a clump of
mesquite in which we found three old nests of Cassidix; two of the
nests were about four feet apart in one tree, and the third was in
another tree 100 feet from the first.
Specimens (4): ¢ adult, 38988, testis 6 mm., no fat, 209 gm.,
6 P old, Camp 1, July 7. Female, 38989, ova to 3 mm., fat, 115 gm.,
old brood patch, Camp 1, July 8. Female, 38990, ova to 1 mm.,
moderate fat, 107 gm., 7 P old, brood patch refeathering; ¢ juv.,
38991, testis 3 x 1 mm., not fat, 172 gm., 6 P old; Camp 2, July 9
Specimens from the barrier island are clearly referable to C. m.
prosopidicola, showing no approach to the larger and, in the female,
TABLE 4.—MEASUREMENTS IN MILLIMETERS OF ADULT MALES OF
CASSIDIX MEXICANUS
; Weight
Locality No. Wing Tail Tarsus in
grams
Austins Wexasis 2s... -|) Iv= 184.3 203.8 46.38 225.6 June
137! | (173-200) | (178-232) | (41.8-50.0) | (204-253)
202.2 July
(195-207)
San Patricio Co.,
MeEXASeM eet arr 5 185.2 204.2 46.74 237 .6
| (182-188) | (190-219) | (45.1-50.2) | (228-245)
Barrier Is., Tamps... . 1 178 185 47.1 209
Victoria, Tamps.?....| 4 192.2 224.2 47.77 254.3
(186-200) | (215-232) | (46.0-49.1) | (239-276)
Tampico, Tamps.‘....| 1 197 214 48.3 260
Catemaco, Veracruz ®.. 1 193 216 48.2 257
Data from Selander (1958: 370, 373). Sample sizes, as follows: wing, 137; tail,
119; ‘pill length, 20 (June and July); tarsus, 133; weight, 17 ‘for June, 3 for July.
2. June 13, 1961; breeding condition.
3. May 6, 1961; breeding condition.
4. May 7, 1961; breeding condition.
5. November 28, 1959.
3—3002
334 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
darker C. m. mexicanus of Veracruz and San Luis Potosi. In Table
4, measurements of the adult male from the barrier island may be
compared with those of specimens of C. m. prosopidicola from Texas
and a specimen of C. m. mexicanus from Veracruz; it is apparent
that our specimen is assignable to the former.
Evidence of intergradation between the two subspecies is shown
in a series of birds collected near Ciudad Victoria, Tamaulipas, in
May, 1961. The females in the series are highly variable in color
individually, but are on the average paler than C. m. mexicanus from
Veracruz; the males are distinctly larger than C. m. prosopidicola
from Texas. At Miramar, near Tampico, Tamaulipas, a decided
approach to C. m. mexicanus is also evident in the dark color of
females and in the large size of both males (Table 4) and females,
Agelaius phoeniceus megapotamus Oberholser: Red-winged
Blackbird—This species was recorded only at Camp 1 on July 7,
when we saw two males, one of which was flying south along the
edge of the dunes in a flock of five Great-tailed Grackles. Specimen:
2 , 38992, testis 10 x 7 mm., fat, 54 gm., Camp 1, July 7. The large
size of the testes of this individual indicates breeding condition.
Sturnella magna hoopesi Stone: Eastern Meadowlark.—Meadow-
larks were found in small numbers along the margins of the alkaline
flats at both camps. Breeding was still in progress, for males were
singing and a female shot on July 9 had only recently laid eggs.
Specimens (2): ¢, 38986, testis 13 x 8 mm., not fat, 102 gm.; 9,
38987, ova to 6 mm., 3 collapsed follicles, not fat, 88 gm.; Camp 2,
July 9.
Richmondena cardinalis canicaudus Chapman: Cardinal.—This
species was recorded only in stands of mesquite near Camp 1, as
follows: July 7, two pairs seen, from which a breeding female was
taken; July 8, three birds seen. Specimen: 92 , 38933, edematous
brood patch, 36.5 gm., Camp 1, July 7. Intergrades between the
present subspecies and R. c. coccinea of Veracruz are reported from
Altamira, Tamaulipas (Mexican Check-list, 1957:329).
Mammals
Dasypus novemcinctus mexicanus Peters: Nine-banded Arma-
dillo—Remains of an armadillo (89017) were found in a mesquite
thicket in the dunes near Camp 1 on July 7. The bones are not badly
weathered and were not embedded in sand.
This species has not been recorded previously on the barrier
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 835
island of Tamaulipas, nor, for that matter, on any of the barrier
islands on the western shore of the Gulf of Mexico.
Lepus californicus merriami Mearns: Black-tailed Jackrabbit.—
From two to four individuals were recorded daily in dunes and on
alkaline flats in the vicinity of stands of mesquite and cactus.
Specimens (2): @ adult, 89018, pregnant (two embryos, 28 mm.
in crown-rump length), Camp 1, July 6. Male immature, 89019,
Camp 1, July 7. Our specimens have been compared with two skins
of L. c. curti from the type locality at Eighth Pass, with which they
agree reasonably well in color. The size of the adult female is about
that characteristic of other specimens of adult L. c. curti, but char-
acters of the skull are consistent with those of L. c. merriami.
A specimen of this species from Matamoros and several from
Brownsville, Texas, have been assigned by Hall (1951:48) to L. c.
merriami. Specimens from Padre Island, Texas, reportedly resemble
L. c. curti in smallness of the tympanic bullae but are in other char-
acters referable to L. c. merriami (Hall, 1951:44).
Spermophilus spilosoma annectens (Merriam): Spotted Ground
Squirrel_—These squirrels were moderately common in dunes at
both camps. They were heard calling, and many tracks and holes
were seen. On July 7, at Camp 1, a lactating, adult female (89020)
and two dependent juveniles (89021, skull only, 89022, skin and
skull) were shot at the entrance of a burrow; the uterus of the adult
showed six placental scars.
Our adult specimen has been compared with ten specimens ob-
tained by Hall and von Wedel at Eighth Pass in March, 1950; ours
differs from the ten in being paler and slightly larger. The pallor
is perhaps attributable to seasonal variation, and the size (246-79-
38-7; weight, 133 gm.) is within limits that would be expected in a
larger series of the population sampled by Hall and von Wedel.
Hall (1951:38) referred specimens of this squirrel from Eighth
Pass to S. s. annectens.
Geomys personatus personatus True: Texas Pocket Gopher,—
This pocket gopher was abundant on low, stabilized dunes on the
barrier island from four to 73 miles south of Washington Beach.
One of us (Wilks) made a trip down the beach on May 20 and 21,
1961, and collected specimens at localities four miles south and 33
miles south of Washington Beach; additional specimens were taken
at both Camp 1 and Camp 2 from July 6 to 10. At these localities
the gophers seemed to maintain population densities approximating
336 University OF Kansas Pusts., Mus. Nat. Hist.
those of G. personatus on Padre and Mustang islands on the Texan
coast.
There is but one other record of the Texas Pocket Gopher from
México. Goldman (1915) described G. p. tropicalis from Altamira
on the basis of specimens collected in 1898. Since that time, the
species has not been reported as occurring south of Cameron County,
Texas (Kennerly, 1954), some 50 miles northwest of the closest
station of occurrence of the gophers on the barrier beach of
Tamaulipas.
Our specimens are slightly smaller than G. p. personatus and
slightly larger than G. p. megapotamus, the subspecies of nearest
geographic occurrence to the barrier island. The degree to which
our specimens differ in other respects, such as configuration of the
pterygoid, is being studied further by Wilks. For the present,
reference of our material to the nominate subspecies best expresses
the relationships of these coastal gophers.
The fact that pocket gophers from the Tamaulipan barrier island
occupy a position geographically intermediate between present
Texan populations and the isolated population in southern Tamauli-
pas (G. p. tropicalis) helps explain the origin of the latter. It is
likely that G. p. tropicalis represents the southern remnant of a once
continuously-distributed population of pocket gophers living in
coastal Tamaulipas in mid-Wisconsin to late Wisconsin time. At
that time, sea level is thought to have been considerably lower than
at present, exposing a sandy strip 80 to 100 miles wide off the present
coastline. Presumably this would have been an area suitable for
gophers and for southward dispersal of individuals from Texas.
The only conceivable barrier to dispersal, and thus to a panmictic
population, would have been the Rio Grande, but over the wide,
low and sandy coastal plain the river channel almost certainly shifted
regularly, thus decreasing its effectiveness as a barrier to movement.
With subsequent rise in sea level, the gophers at Altamira became
isolated and have presumably remained so for a considerable time.
To judge by the marked morphologic differentiation of G. p. trop-
icalis, its degree of isolation from other populations has been much
greater than those of populations inhabiting the Tamaulipan barrier
island and the barrier islands of the coast of Texas. Contact be-
tween the latter two populations was probably fairly regular before
man’s stabilization of the channel of the lowermost reaches of the
Rio Grande.
At Camp 1 we found evidence of the former occurrence of gophers
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 337
in an area now largely covered by active beach dunes. Numerous
skeletal parts of gophers and “fossilized” burrows (Plate 8) were
found on the surface where troughs between active dunes reached
down to an older, darker, and more tightly cemented layer of sand
underlying the present dunes. It is clear that these gophers were
not transported there, because the bones were not damaged, some
of the skeletons were almost complete, and many of the bones were
found near the “fossilized” burrows. Weathered but well preserved
skeletal remains of at least 12 gophers were picked up at this site.
Specimens (17): @ , 89023, Camp 1, May 20. 4 92 9, 89024-
26, 89029; 3 3 g, 89027, 89028, 89030; Camp 1, May 21. Male,
89031, Camp 1, July 6. Three g g, 89032, 89035, 89038; 4 9 9,
89033, 89034, 89036, 89037; Camp 2, July 9. Female, §9039, Camp
2, July 10.
Perognathus merriami merriami Allen: Merriam Pocket Mouse.
—An individual taken in a trap in the dunes near Camp 2 constitutes
the first record of this species from the barrier island of Tamaulipas.
This pocket mouse seems to be uncommon on other barrier islands
of the western Gulf of Mexico, for there is only one published report
of its occurrence on Padre Island, Texas (Bailey, 1905:141). Other
nearby stations of occurrence are Altamira, Tamaulipas (Hall and
Kelson, 1960:477), Brownsville, Texas (Bailey, loc. cit.), and 17
miles northwest of Edinburg, Texas (Blair, 1952:240).
Specimen: sex?, 89040, skull only, Camp 2, July 10.
Dipodomys ordii parvabullatus Hall: Ord Kangaroo Rat.—We
found this species uncommon and confined in distribution to dunes,
in which it was recorded as follows: an adult female was shot and
two other individuals were seen at night on July 6 at Camp 1; three
were trapped near Camp 1 on July 7; two were trapped at Camp 2
on July 10.
Specimens (5): @, 89041, 2 placental scars, 46 gm., Camp 1,
July 6. Male, 89042, testes scrotal, 47 gm.; g , 89044, 60 gm.; 2,
89043, 44 gm.; Camp 1, July 7. Sex?, 89045, skel. only, Camp 2,
July 10.
Our material does not differ significantly from specimens obtained
by Hall and von Wedel at Boca Jésus Maria in March, 1950, which
formed the basis for Hall’s description (1951:41) of D. o. parva-
bullatus. This subspecies is presumably confined in distribution
to the barrier island of Tamaulipas. Two immature specimens from
Bagdad, Tamaulipas, were tentatively assigned by Hall (1951:41)
338 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
to D. 0. compactus, a subspecies known otherwise only from Padre
Island, Texas.
Neotoma micropus micropus Baird: Southern Plains Woodrat.—
This species was noted only near Camp 1, where numerous houses
were seen in stands of mesquite and prickly-pear cactus and an adult
male (89046, 330 gm.) was taken on July 6. This species has not
been reported previously from the barrier island of Tamaulipas. Our
specimen is referable to the nominate subspecies and shows no
approach to N. m. littoralis, a subspecies known only from the type
locality at Altamira, Tamaulipas (see map, Hall and Kelson, 1960:
684).
Procyon lotor (Linnaeus): Raccoon.—A weathered skull and a
broken humerus were found at Camp 2. The skull is being studied
by Dr. E. L. Lundelius, who informs us that it matches a number
of raccoon skulls found in archaeological sites along the Balcones
Escarpment of Texas. Such skulls are larger than skulls of raccoons
occurring today in Texas (P. I. fuscipes) and closely resemble skulls
of raccoons (P. I. excelsus) presently confined in distribution to
Idaho, eastern Oregon, and eastern Washington. Further details of
this situation are to be reported elsewhere by Lundelius.
Taxidea taxus (Schreber): Badger.—Two burrows were found
in the stabilized dunes near Camp I, tracks were noted on the alka-
line flats, and a weathered skull (89047) was found on the flats west
of Camp 1 on July 7. The skull appears to be of an immature animal,
for the sutures are not well closed and the teeth show little wear.
Our records require an extension of known range of this species
southeasterly by approximately 50 miles. The only previous record
in coastal Tamaulipas is based on two skulls from Matamoros
(Schantz, 1949:301). The skull from the barrier island cannot be
determined to subspecies but on geographic grounds is referable
to T. t. littoralis, with type locality at Corpus Christi, Texas.
Canis sp.—Numerous tracks made either by Coyotes (C. latrans
Say) or by domestic dogs were seen in dunes and on the beach at
both camps. A weathered, posterior part of a canid skull was found
in dunes at Camp 2 on July 10, and a partial left mandible was taken
on the beach at Camp 1 on July 6. Unfortunately, specific identifi-
cation of the skull fragments is not possible, but the few reasonably
good characters that we can use suggest that our material is of
domestic dogs rather than of Coyotes. Hall (1951:37) found tracks
and other signs of Coyotes at Eighth Pass but did not take specimens.
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 339
Most of the canid scats examined by us contained remains of crabs
and fishes.
Odocoileus virginianus (Boddaert): White-tailed Deer—A
weathered Recent fragment of a mandible (89048) and part of a
femur (89049) of this species were found near Camp 1 on July 7,
and a metapodal was picked up in the dunes at Camp 2 on July 9.
This species has not been reported previously on the barrier island
of Tamaulipas and it probably no longer occurs there, for we saw
no tracks or other signs of it. Hall (1951) did not find it at Eighth
Pass.
Our specimens probably pertain to O. v. texanus but are possibly
of O. v. veraecrucis, which has been reported from Soto la Marina
(Goldman and Kellogg, 1940:89).
The only species of mammal known from the barrier island of
Tamaulipas that we did not find is the Hispid Cotton Rat (Sigmodon
hispidus). Two specimens of this species trapped near Eighth Pass
in March, 1950, formed the basis for the description of S. h. solus
(Hall, 1951:42), a subspecies known only from the type locality.
Discussion
The known vertebrate fauna of the barrier island of Tamaulipas
consists of one species of tortoise, two species of lizards, at least one
(unidentified) species of snake, 49 species of birds (48 recorded
by us and the Semipalmated Sandpiper), and 12 species of mam-
mals. This is clearly a depauperate fauna, such as is characteristic
of islands generally, and indicates that the peninsular nature of the
northern part of the barrier island is of relatively small consequence
in determining presence or absence of species. It is likely that
the restricted environmental spectrum is much more important in
this regard than is the fact of semi-isolation.
Of the 49 species of birds, 10 are known to breed on the island
and an additional 21 are suspected of breeding either on the island
or on small islets in the adjacent Laguna Madre de Tamaulipas.
Eleven species occur on the island as nonbreeding summer residents,
about which we will have more to say below. Four species have
been recorded on the island in summer but breed elsewhere, that
is to say, they only wander over the island (Man-o’-war Bird, Turkey
Vulture, etc.). Two species are known only as migrants, and the
status of one, the Sora Rail, is uncertain. The number of migrant
species doubtless will be greatly increased by field work at those
times when birds migrate.
340 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
The avifauna is not depauperate owing to the exclusion of any
one of the three major zoogeographic stocks thought to be im-
portant in the development of the present North American avifauna
(Mayr, 1946). If we examine the breeding passerine birds of the
barrier island and the breeding passerine assemblage at the same
latitude in lowland Sonora (Mayr, loc. cit.) as to their ultimate
evolutionary sources, we find that for both places somewhat more
than half the birds have developed from indigenous, North American
stocks, about one-third have been derived from South American
stocks, and one-fifth to one-eighth are from Eurasian stocks. It is
most unlikely that such close correspondence in relative composition
of the two avifaunas would occur by chance. Thus, we can only
conclude that each of the historical avian stocks is proportionately
restricted in numbers on the barrier island.
Faunistically, the barrier island resembles Padre and Mustang
islands and the adjacent mainland of Tamaulipas and southern
Texas, reflecting the relative uniformity of environment in this
region. It is apparent that there is a faunal “break” or region of
transition in the vicinity of Tampico, in extreme southeastern
Tamaulipas. On the coastal plain, many tropical species and sub-
species occurring in Veracruz are found north to Tampico but fail
to extend farther northward to the barrier island of northeastern
Tamaulipas. Axtell and Wasserman (1953:4-5), have already com-
mented on this situation, mentioning a number of snakes and lizards
that have differentiated subspecifically on opposing sides of the
Tampican region. They also note that large numbers of the lowland
Neotropical floral and faunal elements reach their northern limits of
distribution within the zone of transition around Tampico, and, also,
many Nearctic elements find their southern distributional limits
there.
Our small samples of birds and reptiles from the island show no
detectable morphological differentiation from adjacent populations.
However, several of the mammals are moderately-well differentiated,
but the patterns and degrees of geographic variation are such that
we can only speculate on the historical derivation of the insular
populations. Lepus californicus curti is presently known only from
the barrier island of Tamaulipas, but Hall (1951:43) has suggested
that it may also occur on the adjacent mainland. A resemblance
between individuals of this subspecies and specimens of L. c. merri-
ami from Padre Island in smallness of the tympanic bullae is re-
garded, probably correctly, by Hall (1951:44) as independent
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 341
development—that is, parallel adaptation to similar environmental
conditions reaching fullest expression on the barrier island of
Tamaulipas. As is also true with Geomys personatus and Neotoma
micropus, the barrier island population of Lepus californicus shows
relationships with animals from Texas and northern Tamaulipas
(L. c. merriami) and no connection with (resemblance to) animals
from the south (L. c. altamirae, known only from the type locality
at Altamira, near Tampico).
In color and cranial proportions, Dipodomys ordii parvabullatus
of the barrier island is closer to D. 0. compactus of Padre Island than
to D. o. sennetti of southern Texas and the Tamaulipan mainland.
But, D. 0. parvabullatus resembles D. o. sennetti in external meas-
urements (Hall, 1951:39). Possibly D. 0. parvabullatus and D. o.
compactus are phylogentically closer to one another than is either
to D. o. sennetti. It is also possible that each evolved independently
from a mainland stock represented today by D. o. sennetti; the re-
semblance of the two insular populations would thus be a matter of
convergence in response to like environmental conditions.
Sigmodon hispidus solus is an insular differentiate that probably
reached the barrier island from the adjacent mainland of Tamauli-
pas, where its apparent closest relative, as judged by morphological
similarity, now occurs.
Nonbreeding shorebirds in summer south of breeding ranges.—
Certain aspects of this subject have already been discussed by
Eisenmann (1951). As he notes, the phenomenon is more regular
and widespread than generally has been appreciated. The old idea,
that such oversummering individuals were “abnormal” or “senile,”
is totally inadequate, especially in view of the frequently large num-
bers of individuals involved.
Eisenmann’s suggestion that nonbreeders are immature is prob-
ably valid, and it is supported by Pitelka’s examination of dowitchers
(1950:28, 51). For gulls, which can be aged by characters of
plumage, there is no question that most nonbreeders are immature.
Unfortunately, there are few criteria for determination of age in
charadriiform birds.
With the possible exception of a specimen of Limosa fedoa, none
of the presumed nonbreeding, oversummering shorebirds collected
by us showed gonadal enlargement above expected minimal sizes
for the species. Even so, the season was late at the time when we
were on the island and most of the birds were molting; it is possible
their gonads had been enlarged earlier in the season. Behle and
342 UNIVERSITY OF KansAS Pusts., Mus. Nat. Hist.
Selander (1953) and Johnston (1956) have shown that nonbreeding
first-, second-, and third-year California Gulls (Larus californicus)
undergo gonadal enlargement in summer. Additionally, nonbreed-
ing first-year males of certain passerine species (for example, the
Brown Jay, Psilorhinus morio; Selander, 1959) are known to ex-
perience partial gonadal recrudescence in summer. It would be
useful, and would facilitate discussion, to have data on gonadal con-
dition of oversummering birds; any functional enlargement would
be worth documenting.
Some species, notably the Semipalmated Sandpiper, Semipalmated
Plover, and Black Tern, oversummer as nonbreeders in such large
numbers that it is obvious that a significant fraction of the total
population of the species does not breed in any one year. This
raises questions concerning the possible ecologic situations that
would select for delay in time of recruitment of young birds into
the breeding segment of the population, assuming that nonbreeders
are immature birds. Delay in maturation, or slow rates of matura-
tion, may show general relationship to paucity of sites of breeding,
as Orians (1961:308) suggests, but the shorebirds with which we
are dealing breed in regions or in habitat-types not characteristically
imposing general restriction on sites of nesting; more than one
answer is necessary for the question even at this level. Data on age
and numbers of nonbreeders, as well as on the ecology of breeding
populations, are critical and are badly needed for most species.
In any event, species for which we have data demonstrating that
they regularly oversummer south of their breeding ranges are prob-
ably adapted to having a part of their populations refrain from
breeding each year. Whether this phenomenon can be explained
solely in terms of selection at the level of individual birds (Lack,
1954) or involves selection of an adaptive response of the popula-
tion as a whole (Wynne-Edwards, 1955; see also Taylor, 1961, con-
cerning Rattus) is a problem that cannot be resolved at this time.
We may note that the species involved ordinarily breed in arctic
and subarctic regions, and it would seem advantageous (as set forth
below) for nonbreeders to remain well south of such high latitudes.
The numbers of oversummering individuals may fluctuate with
over-all population density, possibly as a result of crude density, but
possibly also as a result of emigration of individuals in excess of
optimal density on breeding grounds (see Wynne-Edwards, 1959).
One aspect of this phenomenon not explicitly discussed by Wynne-
Edwards is the possibility that some individuals never move north
to breeding grounds at all, perhaps as a result of a behavioral char-
VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 343
acter genetically-grounded and mediated by delayed maturation of
the neurohumoral “clock.” This certainly would be an economical
means by which population numbers could be regulated, for there
would be a saving of energy in that some individuals not only would
not move north, but also would not participate in the behavioral
interactions involved in territorial spacing. Occurrence of these
birds throughout southern North America, Middle America, and
northern South America may thus reasonably be understood.
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VERTEBRATES FROM BARRIER ISLAND, TAMAULIPAS 345
Ortrans, G. H.
1961. The ecology of blackbird ( Agelaius) social systems. Ecol. Monogr.,
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1955. The ornithogeography of the Yucatin peninsula. Peabody Mus.
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1960. A field guide to the birds of Texas. Houghton Mifflin Co., Boston.
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1951. Frigate-bird, oystercatcher, upland plover and various terns on the
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SELANDER, R. K.
1958. Age determination and molt in the boat-tailed grackle. Condor,
60:355-376.
1959. Polymorphism in Mexican brown jays. Auk, 76:385-417,
SELANDER, R. K., and ALVAREZ DEL Toro, M.
1955. A new race of booming nighthawk from southern Mexico. Condor,
57:144-147.
SHANTzZ, V. S.
1949. Three new races of badgers (Taxidea) from southwestern United
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1946. Handbook of lizards. Comstock Publ. Co., Ithaca, New York.
557 pp.
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1950. The southern limits of the willet’s continental breeding range. Con-
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TayLor, J. M.
1961. Reproductive biology of the Australian bush rat Rattus assimilis.
Univ. California Publ. Zool., 60:1-66.
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1958. Semipalmated sandpiper from Tamaulipas. Wilson Bull., 70:288.
WyYNNE-Epwaprps, V. C.
1955. Low reproductive rates in birds, especially sea-birds. Acta XI
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Transmitted March 15, 1962.
O
29-3002
aN igh nt .
nieen mn
Hv ip
me is
Vol. 12. 1.
2.
3.
4,
(Continued from inside of back cover)
Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry
A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. July 8, 1959
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.
The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49
figures in text. February 19, 1960.
A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By
Theodore H, Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in
text. May 2, 1960.
Natural history of the bell vireo. By Jon C. Barlow. Pp. 241-296, 6 figures
in text. March 7, 1962.
Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C.
: Fox. Pp. 297-307, 6 figures in text. May 21, 1962.
Vol. 18.
fy
10.
Vertebrates from the barrier island of 'Tamaulipas, México. By Robert K.
Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. . Pp. 309-
345, pls. 5-8.- June 18, 1962.
More numbers will appear in volume 12.
Five natural hybrid combinations in minnows (Cyprinidae). By Frank B.
Cross and W. L. Minckley. Pp. 1-18. June 1, 1960.
A distributional study of the amphibians of the Isthmus of Tehuantepec,
México. By William E. Duellman. Pp. 19-72, pls. 1-8, 3 figures in text.
August 16, 1960.
A new subspecies of the slider turtle (Pseudemys scripta) from Coahuila,
oe By John M. Legler. Pp. 73-84, pls. 9-12, 3 figures in text. August
Autecology of the copperhead. By Henry S. Fitch, Pp. 85-288, pls. 13-20,
26 figures in text. November 30, 1960. :
Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and
Rocky Mountains. By Henry S. Fitch and T. Paul Maslin. Pp. 289-308,
4 figures in text. February 10, 1961.
Fishes of the Wakarusa river in Kansas. By James E. Deacon and Artie L.
Metcalf. Pp. 309-322, 1 figure in text. February 10, 1961.
Geographic variation in the North American cyprinid fish, Hybopsis_ gracilis.
By Leonard J. Olund and Frank B. Cross.. Pp. 323-348, pls. 21-24, 2 figures
in text. February 10, 1961.
Descriptions of two species of frogs, genus Ptychohyla; studies of Ameri-
can hylid frogs, V. By William E. Duellman. Pp. 349-357, pl. 25, 2
figures in text. April 27, 1961.
Fish populations, following a drought, in the Neosho and Marais des Cygnes
rivers of Kansas. By James Everett Deacon. Pp. 359-427, pls. 26-80, 8 figs.
August 11, 1961.
Recent soft-shelled turtles of North America (family Trionychidae). ~~ By
ae Webb. Pp. 429-611, pls. 31-54, 24 figures in text. February
Index in press.
Vol. 14, 1.
2.
3.
4,
Vol. 15. 1.
2.
3.
Neotropical bats from western México. By Sydney Anderson. Pp, 1-8.
October 24, 1960.
Geographic variation in the harvest mouse, Reithrodontomys megalotis, on
the central Great Plains and in adjacent regions. By J. Knox Jones, Jr.,
and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961.
Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson.
Pp. 29-67, pls. 1 and 2, 3 figures in text. July 24, 1961. :
A new subspecies of the black myotis (bat) from eastern Mexico. By E.
Rencnd Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December
North American yellow bats, “Dasypterus,” and a list of the named kinds
of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr.
Pp. 73-98, 4 figures in text. December 29, 1961.
Natural history of the brush mouse (Peromyscus boylii) in Kansas with
description of a new subspecies. By Charles A. Long. Pp. 99-111, 1 figure
in text. December 29, 1961.
Taxonomic status of some mice of the Peromyscus boylii group in eastern
Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 118-
120, 1 figure in text. December 29, 1961.
A new subspecies of ground squirrel (Spermophilus spilosoma) from Ta-
maulipas, Mexico. By Tieul Alvarez. Pp. 121-124. March 7, 1962.
Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J.
Knox Jones, Jr., and Ticul Alvarez. Pp. 125-133, 1 figure in text. March 7,
1962.
A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene,
of Texas. By E. Raymond Hall and Walter W. Dalquest. Pp. 135-138,
2 figures in text. April 30, 1962.
A new subspecies of wood rat (Neotoma) from northeastern Mexico. By
Ticul Alvarez. Pp. 139-143. April 30, 1962.
Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul
Alvarez, and M. Raymond Lee. _Pp. 145-159, 1 figure in text. May 18,
1962.
A new bat (Myotis) from Mexico, By E, Raymond Hall. Pp. 161-164,
1 figure in text. May 21, 1962.
More numbers will appear in volume 14.
The amphibians and reptiles of Michoacdn, México. By William E. Duell-
man. Pp. 1-148, pls. 1-6, 11 figures in text. December 20, 1961.
Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox
Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962...
A new species of frog (Genus Tomodactylus) from western México. By
Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962.
More numbers will appear in volume 15.
7.
(Continued from inside of front cover)
Additional remains of the multituberculate genus Eucosmodon. By Robert
W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.
Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-885, 75 figures
in text. June 15, 1956.
Comments on the taxonomic status of Apodemus peninsulae, with description
of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 837-346,
1 figure in text, 1 table. August 15, 1956.
Extensions of known ranges of Mexican bats. By Sydney Anderson. — Pp.
847-351. August.15, 1956.
A new bat (Genus Leptonycteris). from Coahuila: By Howard J. Stains.
Pp. 353-356. January 21, 1957.
A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico.
By Robert J. Russell. Pp. 357-361. January 21, 1957.
Geographic variation in the pocket gopher, Thomomys bottae, in Colorado.
By Phillip M. Youngman. Pp. 368-387, 7 figures in text. February 21, 1958.
New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones,
Jr. Pp. 385-388. May 12, 1958.
Pleistocene bats from San Josecito Cave, Nuevo Leén, México. By J. Knox
Jones, Jr. . Pp. 389-396. December 19, 1958.
New subspecies of the rodent Baiomys from Central America. By Robert
L. Packard. Pp. 397-404. December 19, 1958.
Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405-
414, 1 figure in text, May 20, 1959.
Distribution, variation, and relationships of the montane vole, Microtus mon-
tanus. By Sydney Anderson. Pp. 415-511, 12 figures in text, 2 tables.
August 1, 1959.
Conspecificity of two pocket mice, Perognathus goldmani and P, artus. By
E. ep hee ica and Marilyn Bailey Ogilvie. Pp. 513-518, 1 map: Janu-
ary 14, b
Records of harvest mice, Reithrodontomys, from Central America, with de-
scription of a new subspecies from Nicaragua. By Sydney Anderson and
J. Knox Jones, Jr. Pp, 519-529. January 14, 1960.
Small carnivores from San Josecito Cave (Pleistocene), Nuevo Leén, México.
By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, 1960.
Pleistocene’ pocket gophers from San Josecito Cave, Nuevo Leén, México.
By Robert J. Russell. Pp. 539-548, 1 figure in text. January 14, 1960.
. Review of the insectivores of Korea. ‘By ae Knox Jones, Jr., and David H.
238.
Johnson. Pp. 549-578, February 23, 196 d
Speciation and evolution of the pygmy mice, genus Baiomys. By Robert L.
Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960.
Index. Pp. 671-690.
Vol. 10. 1.
2.
8.
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.
Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.
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. j
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.
Birds found on the Arctic slope of northern Alaska. By. James W. Bee.
Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958.
The wood rats of Colorado: distribution and ecology. By Robert B. Finley,
Jr. Pp. 213-552, 34 plates, 8 figures in text, 85 tables. November 7, 1958.
Home ranges and movements of the eastern cottontail in Kansas. By Donald
W. Janes., Pp. 553-572, 4 plates, 8 figures in text.. May 4, 1959.
Natural history of the salamander, Aneides hardyi. By Richard F. Johnston
and Gerhard A. Schad. Pp. 573-585. - October 8, 1959.
A new subspecies of lizard, Cnemidophorus sacki, from Michoac4in, México.
By William E. Duellman, Pp. 587-598, 2 figures in text. May 2, 1960.
A taxonomic study. of the middle American snake, Pituophis deppei. By
William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960.
Index. Pp. 611-626.
Vol. 11. 1.
DOO AU AN Oe
10.
The systematic status of the colubrid snake, Leptodeira discolor Giinther.
By William E. Duellman. Pp. 1-9, 4 figures. July 14, 1958.
Natural history of the six-lined racerunnet, Cnemidophorus sexlineatus. By”
Henry S. Fitch. Pp. 11-62, 9 figures, 9 tables, September 19, 1958.
Home ranges, territories, and seasonal movements of vertebrates of the
Natural History Reservation. Bv Henry S. Fitch. Pp. 63-326, 6 plates, 24
figures in text, 3 tables. December 12, 1958.
A new snake of the genus Geophis from» Chihuahua, Mexico. By John M.
Legler. Pp. 327-334, 2 figures in text. January 28, 1959.
A new tortoise, genus Gopherus, from north-central Mexico. By John M.
Legler. Pp. 835-343. April 24, 1959.
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.
Fishes of the Big Blue river basin, Kansas. By W. +L. Minckley. Pp. 401-
4492, 2 plates, 4 figures in text, 5 tables. May 8, 1959.
Fas from Coahuila, México. By Emil K. Urban. Pp. 448-516. August I,
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.
Natural history of the ornate box turtle, Terrapene ornata ormata Agassiz. By
John M. Legler. Pp. 527-669, 16 pls., 29 figures in text. March 7, 1960.
Index Pp. 671-703.
(Continued on outside of back cover)
UNIVERSITY OF KANSAS PUBLICATIONS... =
MusEuM OF NATURAL HIsToORyY
Volume 12, No. 8, pp. 347-362, 10 figs.
October 1, 1962
Teeth of Edestid Sharks
BY
THEODORE H. EATON, JR.
UNIVERSITY OF KANSAS
LAWRENCE
1962
UNIvERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL History
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 12, No. 8, pp. 347-362, 10 figs.
Published October 1, 1962
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY
JEAN M. NEIBARGER, STATE PRINTER
TOPEKA, KANSAS
Teeth of Edestid Sharks
BY
THEODORE H. EATON, JR.
The Edestidae are a family of Paleozoic sharks, known from rocks
of Mississippian to Late Permian age, and characterized by a series
of median, symphysial teeth of specialized structure. In Edestus
such teeth occur in both the upper and the lower jaw, but in other
genera it is not yet certain whether those of the two jaws are alike.
Several genera have been described from single sets of symphysial
teeth, presumably belonging to the lower jaw. Sets of lateral teeth
of the lower jaw are known in Campodus (Agassizodus), associated
with the symphysial series. In this genus the structure and arrange-
ment of the symphysial series are not radically different from those
of the lateral teeth, but in other genera the symphysials are in-
creasingly modified until, in the Permian Helicoprion, an extraor-
dinary spiral band of teeth is formed in the symphysis, the func-
tion and position of which have been difficult to determine.
Moy-Thomas (1939) divided the Chondrichthyes into two orders,
presumed to represent divergent evolutionary lines, on the basis of
tooth structures. One, the Bradyodonti (proposed on different
grounds by Woodward, 1921), comprised sharks in which the outer
layer of the tooth was of hard dentine containing vertical tubules,
but without enamel. Moy-Thomas included in the Bradyodonti
the Edestidae and Orodontidae, as well as the Petalodontidae, Co-
chliodontidae, Psammodontidae and Copodontidae (following
Woodward), and he added also the Holocephali. Radinsky (1961)
investigated tooth histology in Chondrichthyes, and concluded that
the order Bradyodonti is probably artificial, that it cannot be de-
fined by the character of “tubular dentine,” and that it should not
include Edestidae and Orodontidae. He suggests that the name
Bradyodonti be retained, however, for the four families designated
by Woodward, “on the basis of slowness of tooth replacement,”
and that the Bradyodonti be “included with chimaeroids under the
term Holocephali . . . Until further evidence is found, it is
suggested that the edestids be kept as a separate group, related to
hybodonts and heterodonts.”
Accepting this idea, we nevertheless meet other difficulties in
attempting to understand the Edestidae. Little material other than
teeth has been found, but probably certain fin-spines and denticles
known under other names will eventually be associated with Edestid
(349)
850 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
teeth. Lateral and symphysial teeth are seldom found in associ-
ation with each other, and this circumstance has resulted in con-
fusion of some generic names, as described on pages 350 and 351. A
new species of Fadenia, based on symphysial teeth, is named and
described farther on. The arrangement of the symphysial teeth
in this family has been the subject of prolonged controversy, now
diminished as a result of the general agreement of authors that
these teeth belong in the median line of the lower jaw, and that
the upper jaw also bears a series of symphysial teeth. But the di-
rection in which the series are oriented, and the manner in which
the Helicoprion spiral evolved from the simpler patterns seen in
other genera, have not been demonstrated satisfactorily. A solution
to these problems is proposed in the last part of this paper. The
illustrations were prepared by Merton C. Bowman.
Obruchev (1953), in a monograph devoted primarily to the work
and discoveries of A. P. Karpinsky, reviewed much of the history of
investigations of the Edestidae. Without adding conclusions or
data of his own, he compiled most of the information so far pub-
lished, with excerpts from unpublished correspondence of Karpin-
sky and others, in a useful summary of the subject. As this publi-
cation, in Russian, may not be conveniently available to many
students in the United States, several figures have been redrawn
from it for the present paper. The value of Obruchev’s work seems
to be as a historical source rather than a contribution of new evi-
dence or interpretations.
Status of Campodus and Agassizodus
DeKoninck (1844:618) described the genus Campodus on the
basis of scattered teeth (C. agassizianus) found in “calcareous
nodules in the black shale of Chokier, underneath the coal forma-
tion” (translated), in the Lower Westphalian or Namurian beds of
Belgium, early Pennsylvanian in age. The teeth are weakly arched,
oblong, up to about 15 mm. in length, and surmounted by a small
series of hard, shiny tubercles, each of which is also oblong but
with its axis transverse to that of the tooth. The tubercles them-
selves bear minute ridges, and similar ridges are seen also in the
depressions between the tubercles. Other, much larger ridged
teeth were intermingled with these, but were referred by DeKoninck
to Orodus ramosus Agassiz.
Teeth apparently congeneric with the latter were described from
the Mississippian of Illinois by Newberry and Worthen (1870:358),
as O. corrugatus. The same authors also described, but with some
TEETH OF EpDESTID SHARKS 351
hesitation regarding its generic distinctness, a series of teeth from
the Pennsylvanian of Illinois under the name Lophodus variabilis
(1870:360). In 1875, however, St. John and Worthen, finding that
the name Lophodus was preoccupied, proposed the genus Agassi-
zodus for L. variabilis, and included in the same genus O. corru-
gatus and two other species. These authors illustrated a large man-
dible of A. variabilis bearing numerous rows of teeth (St. John and
Worthen, 1875, pl. 8, fig. 1). The middle row contains the largest
teeth, but towards both ends of the mandible the teeth become
much reduced and show a form much like those named Campodus
by DeKoninck; the specimen was found in Upper Pennsylvanian
beds near Osage, Kansas.
Lohest (1884) examined DeKoninck’s specimens of Campodus
agassizianus, obtained more material from the beds at Chokier, and
was authorized by DeKoninck to continue the description of the
genus. His figures show teeth closely comparable to the American
Agassizodus, and an almost complete intergradation between these
and the specimens of DeKoninck’s Campodus. There can be little
doubt that the teeth figured by Lohest all belong to one species, it
not to one individual. Lohest concluded that the name Campodus
should apply both to DeKoninck’s C. agassizianus and to the Amer-
ican species of Agassizodus. After study of the figures published
by the authors mentioned, and by Eastman (1902, 1903) and other
more recent writers, as well as of the material in the collection of
the Museum of Natural History, University of Kansas, I am con-
vinced that Lohest was correct, and shall therefore refer to species
that have been described under the name Agassizodus as Campodus.
Most authors subsequent to Lohest (except Eastman) have con-
tinued to recognize Agassizodus, however. Probably this is be-
cause the smallest teeth in the jaws, when present, have not been
compared in detail with DeKoninck’s series, or have not been illus-
trated with sufficient care to enable others to notice the resemblance.
Nielsen (1932), in a review of the literature on Edestid teeth,
justified the name Agassizodus on the ground that symphysial teeth
show generic characters better than lateral teeth, and that whereas
symphysials were known for the American Agassizodus, none were
as yet available for Campodus. This view disregards Lohest’s evi-
dence, which appears satisfactory, that there was no basis for sepa-
rating the two genera in the first place, inasmuch as the broad range
of variation of the lateral teeth in both encompasses the same char-
acters. Neither Campodus nor Agassizodus was founded upon
symphysial teeth.
352 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
Symphysial teeth of Campodus variabilis have been figured by
Eastman (1902, pl. 1, 2, 3; 1903, pl. 1). Isolated symphysial teeth
of the same species are in the KU collection (Fig. 1). St. John
and Worthen (1875, pl. 8, fig. 24) illustrated an imperfect sym-
physial tooth of more massive form, with a thicker median knob,
Fic. 1.—Campodus variabilis, prety aa tooth, posterior
aspect. KU 1056, x 1.5
as “A.” corrugatus. Nielsen (1932:37) doubted that this could be-
long to the same genus as the lateral teeth so named, but I see no
reason to question it.
A New Species of Fadenia
Among the Lower Permian fishes described by Nielsen (1932)
from East Greenland were two species of Edestidae, both repre-
sented by an abundance of teeth. One he named Agassizodus gron-
landicus (here regarded as Campodus), and the other Fadenia
TEETH OF EpESTID SHARKS 353
crenulata. In the latter the symphysial teeth are much more mas-
sive, both in the crown and in the steep lateral flanges, than those
of Campodus. The median peaks are blunt, the anterior and poste-
rior edges of the flanges of each tooth are crenulated, the anterior
most strongly, and the lateral surfaces of the flanges are flattened
and bear horizontal wrinkles (Nielsen, 1932, pl. 4, figs. 1, 2, 9-12).
Nielsen’s characterization of Fadenia is quoted: “Symphysial
teeth (at least of one jaw) disposed in an unpaired row, not fused
with each other, and of a bilaterally symmetrical shape. Crown of
the symphysial teeth as normally in the Edestids developed in such
a way that its right and left halves meet in an acute angle forming a
pronounced rostro-caudal edge. . . . Crown of the symphysial
teeth at the median plane broader than one half of the length, with
the labial margin much and the lingual margin only slightly folded,
and with a sculpture of plicae, which, possibly on account of wear,
are much less distinct on the highest median than on the lowest
lateral parts. ” He describes and figures both the sym-
physial and the lateral teeth, but inasmuch as no lateral teeth ac-
company the specimens to be described here, only the symphysials
are pertinent. Nielsen also remarks that there is less difference be-
tween the lateral and symphysial teeth of Fadenia than between
those of any other edestid, and this genus must therefore be the
most primitive one known in the family. The type species, F. crenu-
lata, was found in Pennsylvanian limestone, Cape Stosch, East
Greenland.
Two specimens in the Museum of Natural History, University of
Kansas, have characters that place them in Fadenia rather than
Campodus, and are described here as
Fadenia gigas new species
Type: Two symphysial teeth in place on a block of osteodentine (Fig. 2);
No. 1023, Museum of Natural History, The University of Kansas; found 4 feet
below top of Cherokee shale (Lexington coal), Lower Pennsylvanian, at Lucas,
Henry County, Missouri. There is no information as to the collector or date.
Diagnosis: The teeth resemble those of F. crenulata in proportions and
shape, in having a rostro-caudal edge in the median line, and in a sculpturing
of plicae on the lateral surfaces. But they differ in having both the labial
and the lingual margins much folded, and in being of far greater size (gigas,
Greek, giant). The height of the more complete tooth is 79 mm., not in-
cluding a small part of the tip that is missing; its anteroposterior breadth in
the median plane is 45 mm. These measurements contrast with 23 and 15.5
mm., respectively, as determined from Nielsen’s fig. 12 on plate 4.
854 University OF Kansas Pusts., Mus. Nat. Hist.
Fic. 2.—Fadenia gigas, new species, symphysial teeth,
right side. KU 1028, x 0.65.
Referred specimen: KU 865, consisting of two symphysial teeth, attached
to a block of osteodentine, that are smaller than those of the type. The folds of
the anterior and posterior margins are farther apart and less numerous than
in the type, but the general shape and the pattern of plicae on the surface
closely resemble it. There is no information on the source of this specimen,
but its form, color, and mode of preservation strongly suggest that it came
from the same horizon as the type. Probably the specimens pertain to different
regions of the symphysial series, but there is no evidence to show whether the
series formed a simple arch or a spiral; the former, to judge from Nielsen’s
F. crenulata, is more probable. The shark from which they came must have
been large, perhaps four or five meters in length, to accommodate a series
of such massive teeth and their counterparts in the opposing jaw.
Position and Evolution of the Symphysial Teeth
Neither the orientation of the Helicoprion spiral in the animal
that carried it, nor the relationship of this curious device to the
symphysial series in other edestid sharks seems to have been stated
convincingly. In the more generalized members of the family,
Campodus and Fadenia, it is clear that in the lower jaw, at least,
a median row of teeth curved forward and outward, that they repre-
sented a growth series, and that the more lateral teeth were also
arranged in series that curved up and out over the occlusal surface
of the mandible. Whether these also were growth series is not en-
tirely clear, but the manner of their replacement may not have been
TEETH OF EDESTID SHARKS 355
the same as in most sharks. The symphysial and lateral teeth in
Campodus and Fadenia are not greatly different in structure. A
growth series in a modern
shark, Lamna (Fig. 3), shows
the usual method of growth
and replacement, the largest
teeth being those that have be-
come functional and that will
presently be lost.
In Edestus mirus (Fig. 4)
Hay (1912) was able to show
that both the lower jaw and
the upper carried symphysial
teeth in long arched series;
Hay also made it clear that the
upper series was single, like the
lower, and not paired (as erro-
neously stated by Nielsen,
1932:26). The lower ends of
the serrated symphysial crowns
in Edestus turn back, each be-
ing overlapped by the tooth
behind it, and the osteoden-
tinal bases are partly or com-
Bie tely piece pea ENE bar Fic. 3.—Vertical section through de-
that supports the series. Fig. 5 veloping tooth-series in Lamna. Ante-
( Hay, 1909) shows the sym- Onachee ee (After Owen, from
physial series of E. crenulatus,
and Fig. 6 (Woodward, 1917) shows that of E. newtoni. These
figures demonstrate the varying curvature of the arch in Edestus;
E. newtoni seems most nearly to approach the spiral form.
Having established that the lower ends of the crowns, on each
side of the bar, turn posteriorly beneath the following teeth, it
seems that in the more extended series in Helicoprion (Fig. 7) the
same characteristic could be used, in the absence of any associated
parts of the head, to decide which way is posterior in the functional
(largest) teeth of the series, provided that any close relationship
exists between these two genera. Hay (1912) decided that this
procedure was not acceptable because it led “to the absurd con-
clusion that the very small teeth of the innermost coil are the
tS
=e
AS
ns
os
fe
356 University OF Kansas Pusts., Mus. Nat. Hist.
ones that were last formed,” and some other, more recent, writers
have agreed. If, however, the teeth belong in the mouth and are
homologous with those of Edestus and other sharks (which cannot
be doubted), there are only two possible ways to place them in
the symphysis. Either the large teeth are posterior to the sym-
Fic. 4.—Edestus mirus, upper and lower symphysial teeth,
right side. (After Hay, 1912.) x 0.75.
Fic. 5.—Edestus crenulatus, symphysial foeth, right side. (After Hay, 1909.)
x 0.65.
TEETH OF EpEesTID SHARKS 857
Fic. 6.—Edestus newtoni, symphysial teeth, left side. (After Woodward,
1917.) x 0.4.
physis, and the size decreases forward, as teeth that were formerly
in use are pushed down, under and inward (the smallest teeth
being oldest of all), or the large teeth are anterior, and are fol-
lowed from behind by a long coiled series of replacing teeth which
are still in the earlier stages of growth; in this case the smallest
teeth are, indeed, those most recently formed, as is true in all
known replacement series in sharks. The teeth come up and for-
ward over the surface of the jaw from behind, and as the older,
larger teeth are broken off anteriorly, those behind take their place.
This concept is fully in agreement with our information on the
teeth of Edestus, it accounts for the existence of the spiral as a
growth series, and it places Helicoprion at the peak of specializa-
tion in the family. There is, however, some difficulty in under-
standing how an arched replacement series evolved into a spiral,
and how it is that the youngest tooth-buds come to lie inside of
two or more whorls of older teeth.
The key to both problems may lie in the fusion of the bases
of the symphysial teeth into a continuous curved rod. The crowns
fit against one another, even in early stages of growth, and the
rod, at first small, slender, and merely arched, becomes extended
as the teeth grow. A given number of teeth occupying a certain
distance on the rod at an earlier stage would necessarily occupy
a greater space as they grew. Pressure is therefore exerted to make
the spiral grow in length, a pressure corresponding to that which
358 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
causes the movement of tooth-buds upward in the growth series
of such a shark as Lamna. In Helicoprion, however, the rigidity
of the growing spiral rod, formed by fusion of the basal dentine
of the teeth, probably compelled an actual movement in two di-
rections during the ontogeny of the shark (Fig. 8). Presuming
‘VY
Fic. 7.—Helicoprion ferrieri, symphysial teeth, right side. (After Hay, 1909.)
Approximately x 1.
that the first stage of formation of the symphysial tooth-row took
place on the lingual, or posterior, aspect of the symphysis, as is
most probable, and that the rate of growth is greatest in the
TEETH OF Eprestip SHARKS 359
youngest individuals, the tendency of the uppermost teeth in the
series to move up and forward would be equalled or exceeded by
the tendency of the lower part of the series to be pressed down.
Since the curvature of the symphysial surface in the youngest
individuals is much greater than in older, and since the spiral rod
is present upon this surface at an early age, any downward move-
ment compels the series of younger tooth-buds to retreat inward
and forward beneath the symphysis, curling eventually up and
back, below the functioning row of larger teeth that stand on the
occlusal surface. It appears more likely that a row of embryonic
tooth-buds would retreat into the tissues of the jaw, forming in a
sense a spiral pocket for their development, than that old, used
Fic. 8.—Helicoprion. Diagram of growth of spiral band of symphysial teeth.
teeth would be forced down into the flesh in a spiral from the
anterior end of the functioning tooth-row. If growth, loss and
replacement in the symphysial series were rapid, the largest teeth,
here considered to have been those at the anterior margin of the
jaw, would be replaced frequently as they broke off and the large
end of the spiral moved outward.
Evidently in the family Edestidae the food was something that
required crushing; there is little difference in the lateral teeth
between one genus and another, so far as known. But evolutionary
advance in the family was associated with the increasing specializa-
tion of the median tooth-series, probably in both upper and lower
jaws. This implies that the feeding action involved a different
use of the symphysial teeth from that of the lateral series, and in
Edestus the form suggests a scissorlike motion of the powerful
symphysial series in a vertical plane. Presumably the teeth could
then be used to cut off or pull away objects growing on a surface
(such as mussels, corals, hydroid clusters, stalked barnacles, and
360 Universiry OF KANnsAs Pusts., Mus. Nat. Hist.
crinoids), or, alternatively, to dig and pull out burrowing clams
from mud or sand. It seems more likely that the first of these is
correct, and moreover that the teeth did not have a defensive or
predatory function.
Fic. 9.—Hypothetical reconstruction of Helicoprion, showing symphysial
teeth.
Fic. 10.—Van Den Berg’s Sar aeay of Helicoprion. (From Obruchev,
1953.
TEETH OF EprEsTID SHARKS 361
If Helicoprion is the end-form of the family in a morphological
sense, and if it resembled Edestus in having a symphysial series in
the upper as well as the lower jaw, then the diagrammatic restora-
tion in Fig. 9 suggests the possible arrangement of the teeth, the
head being viewed as a transparent object. Although direct evi-
dence is lacking, there can be little doubt that in support of a
mechanism primarily for crushing shells the upper jaw must have
been fused with the cranium, as in Holocephali. The only previous
figure known to the writer, in which the spiral is shown with the
larger teeth forward, is a sketch sent by Van Den Berg to Karpinsky
in a letter dated November 21, 1899, reproduced by Obruchev,
1953, and here shown as Fig. 10. In this drawing, however, Van
Den Berg appears to have regarded the tooth series as resting upon
a median lower jaw, and he did not know that there were also
lateral teeth. He compared the spiral to the radula of a gastropod
mollusk.
LITERATURE CITED
DerKoninck, LAuRENT G.
1842-44, Description des animaux fossiles qui se trouvent dans le terrain
carbonifére de Belgique. 2 vols. Liége.
EAsTMAN, C. R.
1902. Some Carboniferous Cestraciont and Acanthodian Sharks. Bull.
Mus. Comp. Zool., 39(2): 55-99, 14 figs., 7 pls.
1903. Carboniferous fishes from the central western states. Bull. Mus.
Comp. Zool., 39(7): 163-226, 17 figs., 5 pls.
HaAvaO iP:
1909. On the nature of Edestus and related genera, with descriptions of
one new genus and three new species. Proc. U. S. Nat. Mus., 37:
43-61, 7 figs., 4 pls.
1912. On an important specimen of Edestus; with description of a new
species, Edestus mirus. Proc. U. S. Nat. Mus., 42: 81-38, 2 pls.
LouweEst, MAXIMIN.
1884. Recherches sur les poissons des terrains paléozoiques de Belgique.
Annales Soc. Geol. Belgique, 11: 295-325, 4 text figs., 3 pls.
Moy-THoMas, J. A.
1939. The early evolution and relationships of the elasmobranchs. Biol.
Reviews, 14: 1-26, 12 figs.
Newesenrry, J. S., and A. H. WorTHEN.
1870. Descriptions of fossil vertebrates. In: Paleontology of Illinois.
Geol. Surv. Illinois, 4(2): 345-874.
NIELSEN, Ec.
1932. Permo-Carboniferous fishes from East Greenland. Meddelelser
om Gr¢gnland, 86(3): 1-63, 7 figs., 16 pls.
OxsRUCHEV, Dimitry.
1958. Study of Edestids and the works of A. P. Karpinsky. (In Russian)
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Ravinsky, L.
1961. Tooth histology as a taxonomic criterion for cartilaginous fishes.
Jour. Morph., 109(1): 73-92, 2 text figs., 10 pls.
Sr. Joun, Orestes, and A. H. WoRTHEN.
1875. Descriptions of fossil fishes. In: Paleontology of Illinois. Geol.
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Woopwapp, A. S.
1917. A new species of Edestus. Quart. Jour. Geol. Soc. London, 72
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Soc. London, sess. 183: 20-89, 4 figs.
Transmitted June 18, 1962.
O
29-4226
UNIVERSITY OF KANSAS PUBLICATIONS Se elec
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October 25, 1963
Variation in the Muscles and Nerves
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(Tympanuchus and Pedioecetes)
BY
E. BRUCE HOLMES
UNIVERSITY OF KANSAS
LAWRENCE
1963
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16. Mammals of ithe 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 Sydney Anderson. Pp. 415-511, 12 figures in text, 2 tables.
August 1, 1959.
(Continued on inside of back cover)
UNIVERSITY OF KANSAS PUBLICATIONS
MvuSsEUM OF NATURAL HISTORY
Vol. 12, No. 9, pp. 363-474, 20 figs.
October 25, 1963
Variation in the Muscles and Nerves
of the Leg in Two Genera of Grouse
(Tympanuchus and Pedioecetes)
BY
E. BRUCE HOLMES
UNIVERSITY OF KANSAS
LAWRENCE
1963
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 12, No. 9, pp. 363-474, 20 figs.
Published October 25, 1963
UNIVERSITY OF KANSAS
Lawrence, Kansas
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PRINTED BY
JEAN M. NEIBARGER, STATE PRINTER
TOPEKA, KANSAS
1963
29-5835
Terminology
Variation in the Muscles and Nerves
of the Leg in Two Genera of Grouse
(Tympanuchus and Pedioecetes )
BY
E. BRUCE HOLMES
CONTENTS
Acknowledgments ............. Zip ftte bRet skPs a
Skeleton
Nerves
Sciatic Nerve
Peroneal Nerve
Tibial Nerve
Muscles
. Extensor Iliotibialis Lateralis
. Extensor Iliotibialis Anticus
. Flexor Ischiofemoralis
. Adductor Superficialis
. Adductor Profundus
SSSSSSESSSSSSSSSESS
PATTIES Dats Ie. Maio AOS LA eta dye oo Oe
Extensor Jliofibularis
Piriformis
Psoas
Flexor Cruris Medialis
Caudofemoralis
866 UnIverSITY OF Kansas Pusts., Mus. Nat. Hist.
PAGE
NiO btuirator (0 Pa: aedet ahs yeti Eyes Oe ae 422
M: -Femorocruvalis’*{ r= yet eo ae eee SOWIE ree Bere Bs 425
M. Gastrocnemius 71): 295.0) CEL! Rie Se Oe eee 426
M. Flexor Perforans et Perforatus Digiti II ............... 427
M. Flexor Perforans et Perforatus Digiti II] .............. 429
Ma Hlexor Perioratus igttiwiy We ae ee eee 430
M; Flexor Pertoratus Dictate e..28 32> ea nt eee 432
IM. tlexom Pertoratus Digit Mie eo ae cre tena eee 433
MM sblexor, THallucis! Longus a2) 4 vino nbeees ct eee 435
INL AMtATIS 721, Bde ets aes ke ca 435
Ne Hlexor Digitorum: (z0N@US 6. eda se ea ee ee 436
NMP ODMLCUS a, tect siamese anu cnn. 1 5 oe eerie 438
Wi spPEVONEUS: MOMOUS Scum nied Mec mnvien wor eae ae teen eR ates 438
Nicol btaltG *AmtCUSs oat, yee e siete daha nts dyavees Oe Baie Meare 439
M-., Extensor: Digitorumy LOnMeuUs 2 5... ..ces.nue sae eenes 440
IVE; ABEFONEUS <DIeVAS <.4-3 o. saci» ee ie beens deneee eae 441
Nie extensor, Malluciswicongus Qo. Al... ....ca2 eee at 442
IME vAbductor Digit 4) wage oe ete i oan eee eee 443
M.. Hextensor brevis Digit (le 0) 2 si. fan « nc cls harehee See 444
Mi. textensor Proprius Jorn 20). 2c hs Scie oees gee 444
NESE xtensOr BrevissD1entt RY. eect 46h echoces oe eee gee 445
NV iepiGUMDEICAlis” = funn tee nk ake eysacdes weaves a Soke se aero aoe 445
Nie -Abductor Droit “We eee Fe omg nuele enn es ee 446
Ve Flexor “Halhicis Brevis 2575.2. oy «sys eee get or eee 446
Wiscussion and Conclusions. 37.40.42. sche on ss eee ee 446
Analysis, Gf Individual Variation... 4.0... . sue. so ee eee ee 446
MAAS CLES epee. oe Aen esa abe TENS suet dio oh on Ont vin sn SuSE a ote Oe 447
INGEVES SGA eiaa bk a edeee. win ote ttl mc on ate gece eae 449
Analysis of Variation Between Species ................... 451
Comparison with Other Studies of Innervation ............ 452
Summary. «5-3 fee <a ee ee ee Tene! Oe Pet eh ree ee 457
Literature Cited . oy Ee phen eee be Oe eS Ee eo ee 473
INTRODUCTION
The purposes of this study were: (1) to obtain information on
individual variation in the anatomy of the muscles and nerves of
the leg of Tympanuchus cupido pinnatus (Greater Prairie Chicken),
T. c. attwateri (Attwater’s Prairie Chicken), T. pallidicinctus
(Lesser Prairie Chicken), and Pedioecetes phasianellus jamesi
(Sharp-tailed Grouse); (2) to determine whether or not the two
species of the genus Tympanuchus differ constantly in the myology
of the leg; and (3) to determine what constant differences in the
myology of the leg exist between the two closely related genera
Tympanuchus and Pedioecetes.
These particular birds were chosen because they are closely re-
lated, and closely resemble one another in habitats occupied and in
patterns of behavior. It was desired to study examples that showed
as few adaptive differences as possible among the grouse. Series
of each of the three species of grouse were readily obtainable, mak-
ing it possible to draw comparisons at the level of individuals, sub-
species, species, and genera.
The study here reported on was begun in the spring of 1957 and
was completed in the autumn of 1961.
Prior work on the muscles of the leg of birds has been reviewed
by Hudson (1937) and Hudson, et al. (1959). Only papers deal-
ing with the innervation of the leg in birds are reviewed below.
DeMan (1873) treated the nerves of Paradisea papuana, Corvus monedula,
and the chicken; he also commented briefly on a few other species. Jhering
(Ihering, 1873) briefly described the lumbosacral plexus in approximately a
dozen birds, but illustrated only two. Gadow (1880) described the nerves in
Struthio, Rhea, and Casuarius; his paper contains some excellent illustrations of
nerves. Unfortunately, the text is marred by numerous confusing typographical
errors. Carlsson (1884) described the nerves of Eudyptes chrysolopha, Alca
torda, Mergulus alle, and Mormon arcticus. Gadow (1891) described the
nerves in a study that included a large variety of birds, but published few
illustrations. DuToit (1913) described the lumbosacral plexus of the chicken.
Romer (1927) gave the innervation of the hip and thigh muscles in the chicken,
but did not cover the lumbosacral plexus. Appleton (1928) gave the inner-
vation, in various birds, only of those muscles of the hip and thigh that are
supplied by the tibial and peroneal nerves; he did not include the lumbosacral
plexus. Sudilovskaya (1931) described the nerves of Struthio, Rhea, and
Dromaeus (Dromiceius). Unfortunately, his illustrations are almost useless as
far as the nerves are concerned. Boas (1933) described the lumbosacral
plexus in a large number of birds. His extensive account includes numerous
good illustrations. Howell (1938) listed the innervation of the hip and thigh
muscles in the chicken; he did not include the lumbosacral plexus. Fisher
(1946) listed the innervation of the muscles of vultures, but did not include
the lumbosacral plexus. Wilcox (1948) gave the innervation of the muscles
(367)
368 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
of Gavia immer, but did not include the lumbosacral plexus. Fisher and Good-
man (1955) described the nerves in the Whooping Crane. Papers by Chomiak
(1950) and Yasuda, et al. (1959), both dealing with the chicken, were not
examined.
MATERIALS AND METHODS
Complete dissections of the muscles and nerves were made in eight legs
(of five specimens) of the Lesser Prairie Chicken (Tympanuchus pallidicinctus ),
six legs (of four specimens) of the Greater Prairie Chicken (T. cupido pin-
natus ), three legs (of two specimens) of Attwater’s Prairie Chicken (T. cupido
attwateri), and six legs (of four specimens) of the Sharp-tailed Grouse
( Pedioecetes phasianellus jamesi).
For convenience and simplicity of reference, each specimen has been
designated by a symbol consisting of the first letter of the genus and of the
species (and also of the subspecies in T. cupido) plus a number. The letter
“L” or “R” is added to indicate the left or right leg. Thus the symbol T.p.
1L refers to the left leg of specimen number one of T. pallidicinctus.
All specimens are in the University of Kansas Museum of Natural History.
The catalogue number of each specimen, and the legs of it that were dissected,
are listed below.
4120 ama Ont Sergi Seek ea KU38520 Repeal ogee. it. eee KU38518
ep OR ee ee ne KU88521 Teas AR goed oe ee KU36617
pc hs sent ese KU38522 SECO ate a eer ae KU36618
SWAT, es Oy, Rae RED KU38523 Pips ERs Ay sees, eee KU38526
MO er nS bot fas KU38524 1 OA Dee RARER eh Dee So KU38527
Depry Jee KU88515 Ben Slea s o.t senator KU38528
SLRS ys kA bre] A Re en KU38516 Pip salsa tik eet. Ma ae KU38529
AROS Oks 6: fiche eke aac KU38517
The specimens were injected in the field either with formalin (10%) or
embalming fluid, except for those of T. c. attwateri, which were frozen; the
latter were later injected with embalming fluid. Injection in all the birds was
by hypodermic syringe into all major muscle masses, into the body cavities,
and subcutaneously in the neck, wings, and feet. In those specimens injected
with embalming fluid, the body cavities were injected with formalin. The
embalming fluid consisted of 70 per cent alcohol, glycerin (or propylene
glycol), and formalin (full strength) in the approximate ratio of 78:20:2, re-
spectively. This fluid gave good preservation; these specimens had the ad-
vantages of lacking almost entirely the irritating odor of formalin and of hav-
ing pliable tissues. The skin of those specimens originally injected with
formalin was slit in several places and they were transferred to crocks con-
taining embalming fluid (without the formalin). After a period of many
weeks, with two changes of fluid, most of the formalin odor was eliminated
and the muscles were sufficiently pliable to be easily dissected. All specimens
were kept in containers filled with embalming fluid. No mold ever appeared,
even though no phenol or other chemical was added.
To facilitate comparison, two or three specimens were frequently dissected
simultaneously. The nerves and smaller muscles were dissected with the aid
of a stereoscopic microscope mounted on a long movable arm. In order satis-
factorily to expose the lumbosacral plexus the posterior half of the sternum and
pectoral muscles, as well as the abdominal viscera, were removed.
To insure more nearly accurate proportions, drawings of the pelvis and of
some of the muscles were made with the aid of photographs of the several
specimens listed above.
MuscLes AND NERVES OF LEG OF GROUSE 869
TERMINOLOGY
Skeleton
The majority of the osteological terms used in the present paper
are those used by Howard (1929); however, many skeletal features
are not named by Howard. Since names for most of these parts
were not found in the other literature examined, it was necessary
for me to propose terms for them. Most of this new terminology
pertains to the pelvis. All of the osteological terms used in the
present paper, whether used by Howard or not, are briefly defined
below. Those of the pelvis are illustrated in fig. 1. Most of the re-
maining terms are illustrated by Howard (1929).
PELVIS
The median dorsal ridge is the blunt ridge in the midline of the anterior
part of the synsacrum formed by the neural spines of the vertebrae. The anti-
trochanter, on the posterodorsal rim of the acetabulum, is a pyramid-shaped
projection that articulates with the proximal end of the femur. The anterior
iliac crest is a ridge along the dorsomedial border of the ilium, beginning al-
most at the anterior end of that bone; the crest curves laterally as it extends
posteriorly and (for purposes of the present definition) ends at the level of
the posterior edge of the antitrochanter, where the crest is continuous with the
lateral iliac process. The lateral iliac process is a pronounced, laterally or
ventrolaterally, projecting ridge on the ventrolateral surface of the ilium pos-
terior to the level of the antitrochanter; the process does not extend as far as
the posterior end of the ilium. The lateral ischiatic ridge is a relatively slight
ridge continuous with the posterior end of the lateral iliac process and curves
posteroventrally across the lateral surface of the posterior part of the ischium;
the ridge extends to the ventral edge of the ischium in some individuals and
not in others. The dorsolateral iliac ridge begins at the lateral edge of the
ilium near the posterior end of the lateral iliac process and curves postero-
medially and somewhat dorsally, extending to the posterior edge of the ilium.
The lateral iliac fossa is the concavity below the overhanging lateral iliac
process. The ilio-ischiatic fenestra is a large oblong opening behind the
acetabulum between the ilium and the ischium. The obturator foramen is a
small oval opening posteroventral to the acetabulum between the ischium and
the pubis. The ventral ischiatic tubercle is the angle formed by the ventrally
projecting ischium at the point (near its midlength) where the ischium over-
laps and lies lateral to (and fused to) the pubis. The pectineal process is an
anterolaterally directed projection of the ventrolateral edge of the ilium antero-
ventral to the acetabulum. The femoral notch of the ilium is a shallow notch
in the ventrolateral edge of the ilium approximately halfway between the last
rib and the pectineal process. The oblique iliac crest is a pronounced blunt
ridge on the ventral surface of the ilium and extends from the posterolateral
corner of the last synsacro-thoraco-lumbar vertebra to near the anteroventral
border of the ilio-ischiatic fenestra. The internal ilio-ischiatic crest is more or
less continuous with the oblique iliac crest and extends posteriorly along the
dorsal border of the ischium (forming the ventral border of the ilio-ischiatic
fenestra), and then curves sharply dorsomedially onto the ventral surface of
the ilium. The iliac recess is a concavity dorsolateral to the sharply curving
posterior end of the internal ilio-ischiatic crest.
370 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
anterior iliac crest
median dorsal
acetabulum
antitrochanter
ilio-ischiatic fenestra
ilium
vertebra
ilium
vertebra erie
pectineal process :
obturator foramen lateral
lateral iliac fossa ischiatic
lateral iliac process eS ridge
ventral ischiatic tubercle
synsacro-thoracic
vertebra lium
synsacro-thoraco- ie — ase a
lumbar vertebrae SS ayy
sce parapophysis
femoral notch
synsacro-lumbar G oblique iliac crest
x
vertebrae <a) pa se
synsacro-sacral me | aS
vertebrae =<" fes o
4 diapophysis
pectineal process
Pe
obturator foramen
antitrochanter
internal ilio-
ischiatic
crest
iliac recess
lateral iliac
process
lateral
iliac fossa
Be >
‘
synsacro-
caudal
vertebrae<}
ischium
ilium
pubis
Fic. 1. Pelvis of Tympanuchus pallidicinctus. A. Lateral view. X 1.
B. Ventral view. X 1%.
Muscles AND NERVES OF LEG OF GROUSE 871
The terminology applied to the synsacral vertebrae by different authors
varies. The terminology proposed by DuToit (1913) is employed in the pres-
ent account. See my fig. 1B. This terminology differs considerably from that
used by Howard (1929). DuToit divides the fused synsacral vertebrae into
the following five groups, listed in anteroposterior sequence: (1) synsacro-
thoracic, which bear movable ribs; (2) synsacro-thoraco-lumbar, which lack
movable ribs but possess well developed laterally directed parapophyses, in
addition to the more dorsally directed diapophyses; (3) synsacro-lumbar, which
lack parapophyses, although possessing inconspicuous diapophyses; these
vertebrae are shortened anteroposteriorly and are so firmly fused together that
often the number present can be determined only by counting the inter-
vertebral foramina; (4) synsacro-sacral, which have much more pronounced
transverse processes than do the synsacro-lumbar vertebrae; these transverse
processes are expanded distally where they fuse with the ilium and represent
both parapophyses and diapophyses partly or completely fused together plus
sacral ribs (detectable only in the embryo); there are considered to be two
of these vertebrae; they are situated at approximately the level of the
acetabulum; (5) synsacro-caudal, which include the remainder of the fused
vertebrae; no marked gross morphological features differentiate the synsacro-
sacral and the synsacro-caudal groups of vertebrae. The boundaries between
all but the last two groups of vertebrae are usually, but not always, easily de-
termined. It may be difficult to determine whether a vertebra with rudi-
mentary parapophyses belongs to the synsacro-thoraco-lumbar or the synsacro-
lumbar group. Sometimes a parapophysis will be better developed on one side
of a vertebra than on the other.
FEMUR
The trochanter is a large squarish tuberosity on the lateral surface of the
proximal end of the femur. The trochanteric ridge is a sharp, longitudinal
(relative to the femur) ridge forming the anterior edge of the trochanter.
The obturator ridge is a short, blunt, longitudinal ridge forming the posterior
edge of the trochanter. The anterior intermuscular line is a slight ridge
extending distally from the trochanteric ridge. The posterolateral inter-
muscular line is a slight ridge extending distally from the obturator ridge.
The posterior intermuscular line is a slight, longitudinal ridge on the mid-
posterior surface of the femur. The internal condyle is a large rounded
articular prominence on the medial side of the distal end of the femur. On
the lateral side of the distal end of the femur are two articular prominences—
the lateralmost, smaller one is the fibular condyle, separated by the fibular
groove (visible from posterior aspect only) from the larger and more medial
external condyle. The popliteal area is a depression on the posterior surface
of the distal part of the femur immediately proximal to the condyles.
TIBIOTARSUS AND FIBULA
The inner cnemial crest is pronounced and directed anteriorly on the
anterior surface of the proximal end of the tibiotarsus. The outer cnemial
crest is pronounced and directed anterolaterally on the anterolateral surface
of the proximal end of the tibiotarsus. The rotular crest is transverse and forms
the anterior border of the proximal end of the tibiotarsus; the crest extends
between the dorsal ends of the two cnemial crests and also extends medial
872 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
to the inner cnemial crest. The fibular crest is longitudinal on the lateral
surface of the tibiotarsus and fuses with the middle part of the fibula. The
fibular tubercle is small and on the lateral surface of the fibula near the
level of the middle of the fibular crest. The anteromedial intermuscular line
is a slight ridge extending from the inner cnemial crest down the anteromedial
surface of the tibiotarsus. The anterolateral intermuscular line is a slight
ridge extending from the fibular crest down the anterolateral surface of the
tibiotarsus. The supratendinal bridge is a transverse bony arch over a longi-
tudinal groove near the distal end of the anterior surface of the tibiotarsus.
TARSOMETATARSUS
The hypotarsus is a large, pronounced, squarish protuberance on the
posterior surface of the proximal end of the tarsometatarsus and contains
grooves and canals for the passage of the flexor tendons. The longitudinal
ridges forming the lateral and medial edges of the posterior surface of the
hypotarsus are termed calcaneal ridges. The posterior metatarsal crest is long
and sharp; it is continuous with the medial calcaneal ridge that extends
most of the way down the posterior surface of the tarsometatarsus medial
to the midline; there is an opening between this crest and the tarsometatarsus
immediately distal to the hypotarsus. The medial metatarsal depression is
large; it is on the medial surface of the proximal end of the tarsometatarsus.
The anterior metatarsal groove is a longitudinal groove in the midline of
the proximal part of the anterior surface of the tarsometatarsus. The three
trochleae are large rounded articular prominences at the distal end of the
tarsometatarsus; there is one at the base of each of the digits II, III, and IV.
The term distal foramen (as used by Howard) refers to a short, anteropos-
teriorly directed canal that perforates the tarsometarsus a short distance
proximal to the intertrochlear notch between the trochleae for digits III and
IV. Beginning at the middle of this canal and extending distally at a right
angle to it is the intertrochlear canal, which opens via the terminal foramen
into the intertrochlear notch between the trochleae for digits ITI and IV.
Nerves
For ease of description I have coined terms for the major divi-
sions of the femoral and sciatic nerves.
Muscles
My terminology follows that of Fisher (1946) and Fisher and
Goodman (1955) except for Mm. femoritibialis externus, flexor
cruris lateralis (accessory head), and obturator internus et ex-
ternus. Fisher (1946:547) states that most of his names for the
hip and thigh muscles are those of Howell (1938) and the names
for the shank and foot muscles are those of Hudson (1987). Fisher
deviates, without explanation, from Howell’s terminology in respect
to Mm. vastus medialis and femoritibialis internus, M. caudo-
femoralis, M. flexor cruris lateralis, and Mm. obturator internus
and obturator externus. Fisher’s synonymy of these muscles (1946:
Muscies AND NERVES OF LEG OF GROUSE 373
table 42) is in error. Fisher understandably deviates from Hudson
in respect to Mm. extensor brevis digiti III and extensor proprius
digiti III (see Holmes, 1962), although Fisher’s synonymy is in
error here. See my table 1.
I am not using Fisher and Goodman’s term femoritibialis externus; this
muscle is here considered as a part of M. vastus lateralis. A great deal of
confusion surrounds the terminology of the muscle complex here termed Mm.
vastus lateralis and vastus medialis. Hudson (1937), Hudson, e¢ al. (1959),
Fisher (1946), and Fisher and Goodman (1955) have used different ter-
minology for this complex. Most of the confusion stems from Gadow’s
(1891) unclear description of this complex, which he subdivided into two
units termed Mm. femori-tibialis externus and femori-tibialis medius. Many
birds have three parts to this complex. It is difficult to determine how to
apply Gadow’s two terms to these three parts. As nearly as I can determine,
the correct method is that of Hudson, et al. (1959); but because Gadow’s
terms have been used in different ways (even by the same worker), it seems best
to abandon these terms. Berger (1956:272) believes that the muscle unit
that Fisher and Goodman term M. femoritibialis externus represents a head
of M. vastus lateralis; I am accepting his opinion. For the three parts
of the complex under discussion, I am using the terms M. vastus medialis
and M. vastus lateralis pars lateralis and pars postica.
Fisher (Fisher, 1946; Fisher and Goodman, 1955) considers the muscle
here termed M. femorocruralis as an accessory head of M. flexor cruris
lateralis. The two muscle units in question are closely associated; they
insert broadly on opposite sides of a common tendinous raphe. Howell
(1988:73) considers this to be a secondary fusion of unrelated muscles.
Romer (1927:366) states that in the chick embryo M. femorocruralis is in
reality a shank muscle that migrates into the thigh during development.
Therefore, Fisher’s usage of a single name for these two unrelated muscles
is unsatisfactory. I am using Howell’s terminology in which the name flexor
cruris lateralis represents the main head only of Fishers M. flexor cruris
lateralis and the name femorocruralis represents Fisher's accessory head.
Gadow (1891) divides the obturator complex into two muscles (or muscle
groups), which he terms M. obturator and Mm. accessorii M. obturatoris.
He states that the former is homologous with the mammalian obturator
internus and the latter with the obturator externus. Hudson (1937), accept-
ing Gadow’s homologies, renamed these muscles M. obturator internus and
M. obturator externus. Nearly all subsequent workers have followed Hud-
son’s terminology, with its implication that these muscles are homologous
with the mammalian muscles of the same name. Howell (1938) is an
exception. He points out (pp. 78, 79) that the obturator internus of Hudson
is homologous with the obturator externus of mammals. His evidence is
convincing: “In origin the obturator is somewhat suggestive of the mam-
malian obturator internus, for which it has uniformly been mistaken. That
the latter interpretation is incorrect, however, is attested by the facts that
it receives twigs of n. obturatorius within the pelvis, passes through the
obturator foramen rather than dorsal to the border of the ischium, and it
is segregated from any muscle with tibial innervation. Insertion has shifted
only to a slight and unimportant degree as compared with that of the
874 UNIvERSITY OF Kansas Pusts., Mus. Nat. Hist.
mammalian obturator externus, and beyond question it is the equivalent of
that muscle. The stimulus for a longer muscle, has been the same, resulting
in the extension of origin to within the pelvis of the externus in birds
and the internus in mammals, but the obturator internus is an extension
of a part of the gemellus mass and this does not occur in any vertebrate
class but Mammalia.” Howell applies the term M. obturator to the entire
obturator complex.
Romer (1927), studying the development of the thigh musculature in
chick embryos, concluded that the entire obturator complex is homologous
with the mammalian obturator externus plus quadratus femoris, He con-
sidered the avian M. flexor ischiofemoralis to be the homologue of the
mammalian obturator internus.
Gadow, in his work on the ratites (1880:34), states that M. obturator
(obturator internus of Hudson) cannot be homologous to the mammalian
obturator internus, but must represent the obturator externus. His reasoning
is as follows: “Als M. pectineus kann man diesen Muskel nicht auffassen,
da er auf der Aussenfliche des Trochanter major inserirt, ferner auch nicht
als M. obturator internus der menschlichen Anatomie, da er nicht vom Plexus
ischiadicus, sondern vom Plexus cruralis aus innervirt wird. Seiner Inner-
vation und Insertion nach ware er nur mit dem M. obturator externus zu
vergleichen, wobei er seinen Ursprung im Verhiltniss zum Menschen nur
bedeutend weiter auf das Os ischii und Os pubis distalwiirts ausgedehnt
hatte und so allerdings der Lage nach mit Ausnahme seines Insertionsdrittels
ein ‘internus’ geworden wire.”
Since Gadow gives different homologues for M. obturator in two of his
works (1880 and 1891), one would suspect that he had changed his opinion
in the interim; however, there is no evidence that he did so. In 1880 he
gives supporting evidence (quoted above) for his view; in 1891 he does
not. After describing (1891:173) how the origin of M. obturator in bird
ancestors presumably migrated from a location outside the pelvis to a
position inside the pelvis prior to the meeting of the pubis and ischium
external to the muscle, he states: “Eine ahnliche Entwicklung ist fiir den
Obturator internus der Saugethiere anzunehmen, welchem der M. obturator
der Vo6gel entspricht.” A similar development in mammals is impossible,
owing to the different relationship of the muscle to the pelvic bones in
this class. Gadow says nothing more about the mammalian homologue of
M. obturator. In view of this discrepancy, Gadow can hardly be considered
as a supporter of the idea that the avian M. obturator is homologous with
the mammalian obturator internus.
The evidence is conclusive, it seems to me, that the obturator internus of
Hudson is not homologous with the mammalian obturator internus. Therefore,
the term obturator internus is inappropriate for the avian muscle and must
be abandoned. I shall follow Howell (1938) in naming the entire obturator
complex M. obturator. This term, of course, is not used in the sense in
which it is used by Gadow. The use of the term obturator externus for
the entire complex is avoided because it may not correspond exactly to the
mammalian obturator externus. As mentioned previously, Romer considers
the avian muscle to be homologous not only with the mammalian obturator
externus but also with the quadratus femoris.
MuscLes AND NERVES OF LEG OF GROUSE 375
I am following the policy of Wilcox (1948) and Berger (1952) in latin-
izing the term anterior, changing it to anticus. When preceded by the
feminine word pars, the feminine ending is used (antica).
In table 1 my terminology is compared with that of Fisher and Goodman
(1955), Howell (1938), Hudson (1937), and Gadow (1891). The termi-
nology of Fisher (1946) is identical with that of Fisher and Goodman (1955)
except that in his earlier work Fisher did not describe or name M. femori-
tibialis externus, and M. lumbricales of his earlier work is not mentioned in
his later work. The terminology of Hudson, et al. (1959) is identical with
that of Hudson (1937) except that the manner in which the femoritibialis
complex is subdivided is identical with that of Gadow (1891) and different
from that in Hudson’s earlier work; also the abbreviations p. ext. and p. int.
are substituted in his later paper for pars anterior and pars posterior,
respectively, of M. adductor longus et brevis.
ACKNOWLEDGMENTS
I gratefully acknowledge the generous help of Professor A. Byron Leonard,
under whose guidance this study was conducted and thank Professor E.
Raymond Hall, Professor Howard A. Matzke, and Dr. Irwin Baird for
numerous helpful suggestions and criticisms.
For help in collecting specimens I thank J. R. Alcorn, W. C. Glazener
(through the courtesy of the Texas Game and Fish Commission), Dr. Harrison
B. Tordoff, Jerry Tash, William Brecheisen, and Louis Brecheisen. I thank
also Edwin Gebhard of the Kansas Forestry, Fish and Game Commission
for help in locating the Lesser Prairie Chickens.
I am grateful for the assistance of Mrs. Chester Alexander and Dr, L. C.
Dahl in translating a Russian and a Dutch reference, and thank George
Young and James Bee for making equipment used in my study.
All of the original drawings except fig. 1 were made by me, although the
final inking of figs. 12 through 19 was done by Bret Waller. Fig. 1 was
drawn by Kay Swearingen.
I was aided in this study during the summer of 1960 by a research grant
from the University of Kansas.
SKELETON
Although no special study was made of the skeleton, certain
conspicuous variations are discussed here,
There are a few pronounced differences between the pelvis of
Tympanuchus and that of Pedioecetes. Whereas in the former the
thick lateral iliac process has a pronounced overhang with the
ventral edge lateral to the ischium (fig. 1), in Pedioecetes there is
no overhang at all and the edge of this process is much thinner.
The ischium in Pedioecetes is wider (in dorsoventral extent), es-
pecially posteriorly, than in Tympanuchus. In Tympanuchus the
posteroventral margin of the ischium is rounded and is free from
the pubis, whereas in Pedioecetes it is pointed and fused with the
pubis.
376 University OF Kansas Pusts., Mus. Nat. Hist.
In Tympanuchus cupido (both subspecies) the lateral iliac
process extends farther ventrally than in T. pallidicinctus, approach-
ing or extending ventral to the level of the pubis in the former
species; also the edge of this process is thicker in T. cupido.
All specimens studied have a single synsacro-thoracic vertebra.
The number of combined synsacro-thoraco-lumbar and synsacro-
lumbar vertebrae is eight in each specimen of Tympanuchus and
in one specimen of Pedioecetes phasianellus jamesi and is seven in
three specimens of the latter. In most specimens of Tympanuchus
there are three synsacro-thoraco-lumbar and five synsacro-lumbar
vertebrae, although in two specimens (T. pallidicinctus) there are
four of each group; in one of these latter two specimens the
parapophysis on one side of the fourth synsacro-thoraco-lumbar
vertebra is small. The first (of five) synsacro-lumbar vertebra
has a rudimentary parapophysis on one side in one specimen of
Tympanuchus and on both sides in another specimen. One speci-
men of Pedioecetes phasianellus jamesi has five synsacro-lumbar
vertebrae and the others have four; all have three synsacro-thoraco-
lumbar vertebrae.
NERVES
For each nerve (or plexus) the condition found in most speci-
mens of the Lesser Prairie Chicken (T. pallidicinctus) is described
first. Following this, variations from the typical T. pallidicinctus
condition are given for T. pallidicinctus, then for T. cupido (both
subspecies considered together ), and finally for P. p. jamesi.
Lumbosacral Plexus, Figs. 2, 3
T. pallidicinctus
Descrirtion.—Eight spinal nerves contribute to the lumbosacral plexus.
These are the second through the ninth synsacral spinal nerves (S2 to S9).
The entire ventral ramus of each of these nerves, excepting $2 and S9, con-
tributes to this plexus. The ventral ramus of S2 divides into two branches,
only the posterior of which contributes to the plexus; the anterior branch
directly innervates muscles of the abdominal wall (as does the entire ventral
ramus of S1). The ventral ramus of S9 divides into two branches, only the
anterior of which contributes to this plexus; the posterior branch contributes
to the more posteriorly situated pudendal] plexus.
Each root of the plexus corresponds to a single spinal nerve except one
spinal nerve (S5—the furcal) that contributes a root to both the femoral
nerve and the sciatic nerve; thus typically the plexus has nine roots (but see
below). The four anteriormost roots (S2 to S5) contribute to the femoral
nerve, although the contribution from S2 is small. S38 and S4 contribute
to the obturator nerve. The five posteriormost roots (S5 to $9) contribute
to the sciatic nerve, although the contribution from $9 is relatively small.
MUSCLES AND NERVES OF LEG OF GROUSE 877
InNpiIvipvAL VaRIATION.—In all specimens (of all species) examined, the
right and left sides of the plexus in any one individual were practically
identical. In T.p. 2 (fig. 2B), there appear to be two furcal nerves; $5 is
typical, but a small branch of $4 apparently also contributes to the sciatic
nerve. In T.p. 5, S9 is unique in dividing into three branches; the anterior
two join the sciatic nerve separately; the posterior one joins the pudendal
plexus as usual,
T. cupido
INDIVIDUAL VARIATION.—S2 or S5, or both, may contribute to a limited
extent to the obturator nerve. In T.c.p. 3 (fig. 3A) and T.c.a. 1 and 2,
much of the plexus has shifted one segment anteriorly, relative to the synsacral
vertebrae (the so-called prefixed condition); the roots of the femoral nerve
are S2, S3, and S4 (all large); the furcal nerve is $4 (in T.c.a. 1, S5 gives
an extremely small root to the femoral nerve, thus making two furcal nerves);
six roots (S4 to S9) contribute to the sciatic nerve; S3 and $4 remain as
the main contributors to the obturator nerve except in T.c.a. 2 in which only
S2 and S$ contribute to it.
P. p. jamesi
INDIVIDUAL VaRIATION.—In P.p. 1, the plexus resembles the typical con-
dition in T. pallidicinctus. In P.p. 2, 3, and 4, the plexus is prefixed. P.p. 2
resmbles T.c.p. 3. In P.p. 3 and 4 (fig. 3B) there are two furcal nerves
(S4 and $5); S2 to $4 are the main contributors to the femoral nerve; only
S2 and S3 contribute to the obturator nerve; $4 to S9 contribute to the sciatic
nerve (the anteriormost and posteriormost roots are small).
Femoral Nerve, Figs. 4, 5
T. pallidicinctus
DescripTion.—The femoral nerve is short, dividing inside the pelvis into
six major divisions—anterior, middle, posterior, anterodorsal, dorsal, and
posterodorsal. The anterodorsal and posterodorsal divisions are short, failing
to extend so far laterally as the inguinal ligament; the posterodorsal division
is also small and is usually covered by other divisions and is not visible
when viewed from the ventral side.
The anterior division passes ventral to Mm. iliotrochantericus medius and
iliacus and dorsal to the anterior end of the inguinal ligament. The division
branches into two parts, The anterior part extends around the posterior
border of M. extensor iliotibialis anticus and sends several twigs to the
lateral surface of this muscle. The posterior part passes between the proximal
parts of Mm. extensor iliotibialis anticus and extensor iliotibialis lateralis and
supplies the skin,
The middle division passes ventral to Mm. iliotrochantericus medius and
iliacus and dorsal to the inguinal ligament. The division branches into a
large but variable number of parts. A variable number of branches (usually
two) pass posterior to M. extensor iliotibialis anticus and penetrate the
medial surface of M. extensor iliotibialis lateralis. Several branches supply
the fused Mm. vastus lateralis and vastus medialis. The posteriormost
branch of this division passes between Mm. ambiens and vastus medialis,
378 Universiry OF Kansas Pusts., Mus. Nat. Hist.
giving twigs to the lateral surface of M. ambiens, and sometimes also to the
medial surface of M. vastus medialis, and terminates in M. femoritibialis
internus.
The posterior division, which does not subdivide, spirals completely
around M. psoas (passing in turn anterior, dorsal, posterior, and ventral to it)
and gives twigs into this muscle. This nerve then extends distally into the
proximal part of the shank and there has a nonmuscular termination.
The short, thick anterodorsal division, partly covered by the anterior divi-
sion, turns dorsally and passes through the femoral notch of the ilium and
penetrates the deep surface of M. gluteus profundus,
The slender dorsal division passes ventral to M. iliotrochantericus medius
and dorsal to the inguinal ligament and penetrates the ventral surface of
M. iliacus.
The small, short posterodorsal division penetrates the ventral surface of
M. iliotrochantericus medius.
INDIVIDUAL VARIATION.—In two legs the anterior division gives a twig
or two twigs to M. extensor iliotibialis lateralis, The dorsal division may
fuse proximally with either the anterior or middle division, thus appearing
to be a branch of one of these divisions, In one leg (fig. 5A), there are two
separate branches (both fused with the middle division) to M. iliacus, On
both sides of one specimen (fig. 5A), the anteriormost branch of the middle
division, which supplies M. extensor iliotibialis lateralis, gives off a twig
that anastomoses with the branch of the anterior division that supplies M.
extensor iliotibialis anticus. On both sides of another specimen, the an-
terodorsal division passes lateral to the anterior end of M. iliotrochantericus
medius instead of through the femoral notch, which is lacking.
T. cupido
INDIVIDUAL VaARIATION.—In three legs, the anterior division gives twigs
into M. extensor iliotibialis lateralis. The dorsal division is fused proximally
with the middle division in one instance. In three cases, a twig from the
middle division anastomoses with the branch of the anterior division supply-
ing M. extensor iliotibialis anticus. In the example shown in fig, 5B, a
twig comes off the cutaneous branch of the anterior division, perforates the
ventral part of M. iliacus, and rejoins the cutaneous branch, In both legs
of one specimen, the cutaneous branch of the anterior division perforates the
anterior edge of M. extensor iliotibialis lateralis instead of passing between
the latter and M. extensor iliotibialis anticus. The posteriormost branch of
the middle division, which terminates in M. femoritibialis intemus, perforates
the medial part of M. vastus medialis in one leg. In another leg, one of
the branches to the fused Mm. vastus lateralis and vastus medialis sends a
twig into M. extensor iliotibialis lateralis.
P. p. jamesi
INprvipvaL VARIATION.—In three legs, the anterior branch of the anterior
division is cutaneous and the posterior branch supplies M. extensor iliotibialis
anticus. The dorsal division may fuse proximally with either the anterior or
middle division. In one leg (fig. 4B), there are two branches to M. iliacus,
one associated with the anterior division and one with the middle division.
Musc.Les AND NERVES OF LEG OF GROUSE 879
Obturator Nerve
T. pallidicinctus
Description.—The long slender obturator nerve passes along the oblique
iliac crest and divides into several branches immediately before reaching the
obturator foramen. One or two branches, which do not pass through the
foramen, penetrate the superficial surface of M. obturator pars postica.
Several small branches (variable in number and arrangement) pass through
the obturator foramen and supply pars ventralis, pars dorsalis, and pars antica
of M. obturator, When pars ventralis and pars dorsalis are fused, one branch
perforates the proximal end of this mass and reaches pars antica. One large
branch passes through the obturator foramen dorsal to the tendon of M.
obturator pars postica, then turns ventrally, passing lateral to the latter; the
branch passes between Mm. adductor superficialis and adductor profundus
and gives twigs to each of these two muscles,
InpivipvaL VariaTion.—None of significance in any of the three species.
Sciatic Nerve, Figs. 6, 7, 8, 9
T. pallidicinctus
DescripTIOoN.—The sciatic nerve passes through the anterior part of the
ilio-ischiatic fenestra. Several branches diverge from the nerve immediately
after it emerges from the fenestra. The main trunk of the nerve then extends
distally through the thigh deep to M. extensor iliofibularis and superficial
(lateral) to Mm. flexor ischiofemoralis, caudofemoralis, adductor superficialis,
and femorocruralis. The main trunk subdivides into two large nerves—
peroneal and tibial—that are adjacent and bound to each other throughout
the thigh; the peroneal nerve lies anterior to the tibial. At the distal end of
the thigh the main trunk splits grossly into two large branches that diverge
and enter the shank. This division does not represent the separation between
peroneal and tibial nerves, as is sometimes assumed; the anterior branch
includes a part of the tibial nerve as well as the entire peroneal nerve.
A longitudinal groove is visible grossly on the lateral surface of the main
trunk, except at the proximal end; distally a second groove is visible posterior
to the first one (fig. 6). The long anterior groove indicates the boundary
between the peroneal and tibial nerves; this groove may disappear distally,
although the posterior groove is always visible distally. The posterior groove,
which is continuous with the division of the sciatic nerve into anterior and
posterior branches, represents the boundary between two divisions of the
tibial nerve. (This is discussed in detail below.) In the middle of the
thigh the peroneal and tibial nerves are enclosed in separate connective
tissue sheaths, although the two sheaths are fused together; the point of
fusion is marked by the anterior groove. If the two sheaths are slit open, the
two nerves can be removed and can be seen to be entirely separate. In the
proximal part of the main trunk the peroneal and tibial components are
enclosed in a single sheath and appear as an undivided trunk; but if the
sheath is removed, the two components can be pulled apart rather easily,
although there may be some intermingling of a few fibers. This separation
can be extended to a point proximal to the origin of all the branches of the
sciatic nerve; thus it can be determined which branches arise from the
2—5835
880 UNIvErsITY OF Kansas Pusts.; Mus. Nat. Hist.
peroneal component and which from the tibial. (These -branches arise
from the sciatic nerve as, or immmediately before, the nerve passes through
the ilio-ischiatic fenestra; since this level of the intact nerve could not be
adequately observed, it was necessary to cut the nerve inside the pelvis
and pull the intrapelvic part of the nerve out through the ilio-ischiatic
fenestra. In doing this, care had to be taken to avoid damaging the
most proximal branches.)
Three main branches arise from the peroneal component (apart from
the main trunk) and two from the tibial. Including the peroneal and tibial
components of the main trunk, the sciatic nerve can be divided into seven
major divisions—anterior peroneal, middle peroneal, dorsal peroneal,: posterior
or main peroneal (contributes to main trunk), anterior or main tibial (con-
tributes to main trunk), middle tibial, and posterior tibial. Farther distally,
the posterior peroneal division becomes the peroneal nerve and the anterior
tibial division becomes the tibial nerve. For descriptive purposes, the
term peroneal (or tibial) nerve will be applied only where the nerve is
enclosed in its own sheath, but regardless of whether or not the sheath is
fused with another; proximal to this, where the separation may not be precise,
the terms peroneal (or tibial) division or component will be used.
_The small anterior peroneal division arises from the anterior edge of the
sciatic nerve. Immediately after emerging from the ilio-ischiatic fenestra,
the division turns anteriorly and passes deep to M. piriformis, to which the
division gives a twig (in some cases more than one twig), then continues
forward to supply the posterior part of M. gluteus profundus.
The middle peroneal division branches into two parts. One part pene-
trates the deep surface of the anteroproximal part of M. extensor iliofibularis.
The other part emerges between the proximal ends of Mm. extensor ilio-
fibularis and vastus lateralis and penetrates the deep surface of M. extensor
iliotibialis lateralis.
The dorsal peroneal division arises from the posterodorsal part of the
peroneal component, then angles posteriorly, crossing the dorsal surface of
the anterior tibial division and superficially appears to arise from the tibial
component. The dorsaJ peroneal division usually subdivides into two unequal
branches, both of which penetrate the deep surface of the proximal end of
M. extensor iliofibularis.
The large middle tibial division soon subdivides into two branches that
pass posterodistally lateral to M. flexor ischiofemoralis. One branch (usually
the anterior one) passes lateral to M. caudofemoralis (both heads) and
emerges between Mm. extensor iliofibularis and flexor cruris lateralis and
enters the skin. The other branch passes deep to M. caudofemoralis pars
iliofemoralis, and divides into several branches. Several tiny branches pene-
trate the deep surface of M. caudofemoralis pars iliofemoralis. Another
branch also enters the substance of the latter and emerges from the ventral
edge of it, giving a twig to pars caudifemoralis, then passes lateral to pars
caudifemoralis and enters M. flexor cruris lateralis. Still another branch
passes deep to both heads of M. caudofemoralis and enters the anterior part
of M. flexor cruris medialis.
The small posterior tibial division arises from the posterior edge of the
sciatic nerve. The division diverges from the remainder of the nerve, as the
MuscLes AND NERVES OF LEG OF GROUSE 881
latter passes through the ilio-ischiatic fenestra, and penetrates the dorsal
surface of M. flexor ischiofemoralis.
Below the middle of the main trunk a bundle of fibers of moderate size
separates from the anterior edge of the tibial nerve, leaves the tibial sheath,
and enters its own sheath, lying superficially between the tibial and pero-
neal sheaths (fig. 6). At the distal end of the thigh the sheath enclosing
this bundle of fibers remains fused with the posterior edge of the peroneal
nerve and passes with the latter (diverging from the remainder of the tibial
nerve) through the tendinous guide loop for M. extensor iliofibularis, and
then diverges from the peroneal nerve. Since this bundle of fibers is dis-
tributed with the peroneal nerve, and since the origin of the bundle may be
easily overlooked, it has sometimes been misinterpreted as a branch of the
peroneal nerve, whereas it almost certainly is a branch of the tibial nerve;
this bundle will here be termed the paraperoneal branch of the tibial nerve.
A small but long branch separates from the posterior edge of the proximal
end of the tibial nerve or from the tibial component proximal to this and
extends distally for some distance adjacent to the tibial nerve, then passes
posterodistally between Mm. extensor iliofibularis and flexor cruris lateralis
and supplies the skin.
A small branch separates from the anterior edge of the peroneal nerve
a short distance above the distal end of the main trunk and passes disto-
laterally between Mm. extensor iliotibialis lateralis and extensor iliofibularis
and supplies the skin.
A twig comes off the medial surface of the tibial nerve near the distal
end of the main trunk, passes anteriorly deep to the peroneal nerve, and
penetrates the lateral surface of M. femorocruralis; in some cases two twigs
enter this muscle.
InprvipuaAL VartATIon.—In one leg (fig. 7), the twig to M. caudofemoralis
pars caudifemoralis arises more proximally than usual and perforates pars
iliofemoralis independently of the branch to M. flexor cruris lateralis. The
nerve supplying M. flexor cruris lateralis does not perforate M. caudofemoralis
pars iliofemoralis, but passes deep to it in three legs. In half the legs, the
paraperoneal branch of the tibial nerve, after extending a short distance in
its own sheath, enters the sheath of the peroneal nerve and appears grossly
to unite with it; if, however, the sheath is slit open, the paraperoneal branch
can be easily pulled apart from the posterior edge of the peroneal nerve;
the paraperoneal branch is again enclosed in its own sheath at the distal
end of the thigh. In one leg, the cutaneous branch of the peroneal nerve
perforates the posteroproximal part of M. gastrocnemius pars externa; in
three others, this branch is absent. In one of these last three legs (fig. 7),
the distal cutaneous branch of the tibial nerve is also absent. In three legs
(of different specimens), a minute twig from the middle tibial division passes
posteriorly deep to M. caudofemoralis pars caudifemoralis toward the tail
(fig. 7); this twig joins the pudendal plexus in one leg; in the other two the
twig could not be traced to its termination. Minute twigs come off the
peroneal nerve near the middle of the thigh and enter M. extensor iliofibu-
laris in some legs. In a few cases, a minute nonmuscular twig arises from
the peroneal nerve near the distal end of the main trunk and passes anteriorly
deep to M. vastus lateralis pars pestica (fig. 7).
382 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist.
T. cupido
INDIVIDUAL VARIATION.—In several legs, the nerve supplying M. flexor
cruris lateralis does not perforate M. caudofemoralis pars iliofemoralis, but
passes deep to it. The branch to M. flexor cruris medialis arises from the
posterior (rather than the middle) tibial division in one instance (fig. 8). In
one leg, a minute twig from the middle tibial division passes posteriorly and
joins the pudendal plexus; in another, a similar twig is present but could not
be traced to its termination. In some specimens, minute twigs come off the
peroneal nerve near the middle of the thigh and enter M. extensor iliofibularis.
In one leg, a nonmuscular twig arises from the base of the cutaneous branch
of the peroneal nerve and passes anteriorly deep to M. vastus lateralis pars
postica. In another leg (fig. 8), a tiny additional twig arises from the
posterior edge of the tibial nerve and subdivides, one branch joining the
cutaneous branch of the middle tibial division and the other joining the distal
cutaneous branch of the tibial nerve.
P. p. jamesi
INDIvibUAL VaARIATION.—In both legs of one specimen, the branch to M.
flexor cruris medialis arises from the posterior (rather than the middle) tibial
division; in three legs, this branch arises as an independent division of the
tibial nerve (fig. 6). (Only in one leg does this branch arise as in T. pallidi-
cinctus.) The branch to M. flexor cruris medialis perforates the lateral part
of M. flexor ischiofemoralis in one instance, In all legs except one (nerve
possibly destroyed), a second twig to M. flexor ischiofemoralis arises from the
branch to M. flexor cruris medialis (fig. 6). In one leg (fig. 9), an addi-
tional branch, arising as an independent division of the sciatic nerve, enters
M. extensor iliofibularis distal to the point of entrance of the dorsal peroneal
division; this extra branch arises posterior (adjacent) to the dorsal peroneal
division, but it could not be determined with certainty whether it arises from
the peroneal or tibial component. A minute twig from the branch to M.
flexor cruris medialis passes posteriorly and joins the pudendal plexus in one
leg (fig. 6); in another, a similar twig is present but could not be traced to
its termination. In nearly all the legs, minute twigs come off the peroneal
nerve near the middle of the thigh and enter M. extensor iliofibularis (fig. 6).
In both legs of one specimen, the paraperoneal branch enters the peroneal
sheath (although separable from the peroneal nerve), The distal branch to
M. femorocruralis gives off a long twig to M. gastrocnemius pars media in one
instance (fig. 6).
Peroneal Nerve, Fig. 10
T. pallidicinctus
DescrirTion.—The branch that is given off in the thigh has been dis-
cussed above. The peroneal nerve passes, with the paraperoneal branch of
the tibial nerve, through the guide loop for M. extensor iliofibularis. The
peroneal nerve diverges from the paraperoneal branch and passes along the
anterior (proximal) edge of the tendon of M. extensor iliofibularis medial to
the common tendon of the lateral heads of Mm. flexor perforatus digiti IV
MusSCLES AND NERVES OF LEG OF GROUSE 883
and flexor perforatus digiti II and lateral to the common tendon of the
anterolateral heads of Mm. flexor perforatus digiti IV, flexor perforatus digiti
II, and flexor perforatus digiti III.
The peroneal nerve soon gives off a spray of branches that supplies the
following: femoral head of M. tibialis anticus, tibial head of M. tibialis
anticus (branch passes deep to femoral head), M. extensor digitorum longus
(branch passes deep to tibial head of M. tibialis anticus), and M. peroneus
longus. A part of the nerve may or may not pass through a notch in the
proximal end of the lateral head of M. flexor digitorum longus. The nerve
then extends distally along the anterolateral edge of the latter muscle and
subdivides into two long branches. Gadow (1891) termed these branches the
superficial peroneal and the deep peroneal; his terminology will be used here.
The superficial peroneal branch, after giving off, near its proximal end,
one or two twigs into M. peroneus brevis, passes lateral to the retinaculum
for the tendon of M. tibialis anticus, then across the intratarsal joint lateral
to the latter, then lateral to the insertion of M. tibialis anticus, where the
branch subdivides. One of the two resulting branches gives one or two
twigs into M. extensor brevis digiti IV, then terminates nonmuscularly in
the digits. The other branch passes between the main and accessory inser-
tions of M. tibialis anticus and joins the branch of the deep peroneal which
supplies M. abductor digiti II. (See next paragraph.)
The deep peroneal branch passes through the retinaculum for the tendon
of M. tibialis anticus, lying lateral, then deep, then medial to the latter; it
crosses the intratarsal joint medial to the latter. Immediately above the
insertion of M. tibialis anticus, the deep peroneal branch divides, one branch
passing on each side of the main insertion. The branch passing lateral
to the main insertion passes between the latter and the accessory insertion
(medial to the latter) and is joined by a branch of the superficial peroneal
nerve. This fused branch extends distally between Mm. extensor hallucis
longus and extensor brevis digiti IV and medial to M. extensor brevis digiti
III, giving twigs into the latter and into M. abductor digiti II before terminat-
ing nonmuscularly in the digits. The branch of the deep peroneal nerve
that passes medial to the main insertion of M. tibialis anticus gives one or
two twigs into the proximal head of M. extensor hallucis longus, then
terminates nonmuscularly in the digits.
InpiIvipuAL VARIATION.—In four legs, the branch of the superficial peroneal
nerve that usually joins the lateral branch of the deep peroneal nerve is
lacking (fig. 10B). In these legs it can be seen that Mm. extensor brevis
digiti III and abductor digiti II are supplied by the deep peroneal nerve.
T. cupido
INDIVIDUAL VARIATION.—In two legs, the same branch that gives twigs
into the proximal head of M. extensor hallucis longus also sends a twig into
the distal head of this muscle (fig. 10C).
P. p. jamesi
InDIvmUAL VaRIATION.—None of significance.
384 UNIVERSITY OF KANSAS PusLts., Mus. Nar. Hist.
Tibial Nerve, Fig. 11
T. pallidicinctus
DescripTion.—The branches given off in the thigh have been discussed
in the account of the sciatic nerve. At the distal end of the thigh the
peroneal nerve and the paraperoneal branch of the tibial nerve diverge from
the remainder of the tibial nerve and pass through the tendinous guide loop
for M. extensor iliofibularis whereas the remainder of the tibial nerve does
not. This main part of the tibial nerve immediately divides into three main
divisions—lateral, posterior, and medial.
The lateral division passes between Mm. flexor perforatus digiti IV and
gastrocnemius pars externa and subdivides into two branches, one of which
penetrates the medial surface of M. gastrocnemius pars externa. The other
branch passes deep to the latter and sends twigs into the posterior head
of M. flexor perforans et perforatus digiti II, then passes deep to the
latter and enters M. flexor perforans et perforatus digiti ITI.
The posterior division sends a branch into the medial head of M. flexor
perforatus digiti IV, then passes between the latter and the medial head
of M. flexor perforatus digiti III, and extends distally giving off twigs to
each of the three heads of M. flexor perforatus digiti IV, to each of the
two heads of M. flexor perforatus digiti III, and to each of the three heads
of M. flexor perforatus digiti II. The number and arrangement of these
twigs is variable.
The medial division passes medial to the medial head of M. flexor perforatus
digiti III, sends a twig to the lateral surface of M. gastrocnemius pars media,
then passes into the shank musculature between Mm. plantaris and flexor
hallucis longus, and sends a branch along the medial edge of M. flexor
hallucis longus that gives several twigs into this muscle before terminating
nonmuscularly. A small branch extends to M. popliteus, another to M.
plantaris, and another to the posterior head of M. flexor digitorum longus.
A nonmuscular branch passes between the medial and posterior heads of
M. flexor digitorum longus and extends distally deep to this muscle. A
long branch gives off near its proximal end a variable number of twigs that
pass deep to M. plantaris and enter M. gastrocnemius pars interna; the
branch then extends distally along the lateral edge of M. plantaris and
terminates nonmuscularly.
The paraperoneal branch diverges from the peroneal nerve, passing
medial and then distal to the insertion of M. extensor iliofibularis, whereas
the peroneal nerve passes proximal and then lateral to this insertion. The
paraperoneal branch passes deep to the lateral heads of Mm. flexor perforatus
digiti IV and flexor perforatus digiti II and superficial to the tendon of the
anterolateral head of M. flexor perforatus digiti IV and then passes distally
along the anterolateral borders of the latter and the lateral head of M.
flexor perforatus digiti III and the posterolateral border of M. flexor digitorum
longus. This branch is thus separated from the peroneal nerve by M. flexor
digitorum Jongus and by the fibula; the branch passes along the lateral
surface of the tibial cartilage, continues lateral to the hypotarsus, then turns
medially before extending distally between Mm. abductor digiti IV and
flexor hallucis brevis, sending twigs into each of these muscles and a long
twig into M. lumbricalis before terminating nonmuscularly.
MuscLES AND NERVES OF LEG OF GROUSE 385
INDrIviDbUAL VARIATION.—In T.p. 3L,R (fig. 11B), an extra branch arises
from the tibial nerve as a separate (fourth) division; it enters the medial
head of M. flexor perforatus digiti IV and also gives off a twig that anas-
tomoses with the posterior division (left leg) or with the first branch of the
posterior division (right leg). In T.p. 3R (fig. 11B), a large extra branch
arises from the proximal part of the medial division and passes medial and
then deep to the medial head of M. flexor perforatus digiti III, perforates
the tendinous part of the medial head of M. flexor perforatus digiti II, and
joins the posterior division (lateral to the medial head of M. flexor perforatus
digiti III). A similar branch is found in T.p. 3L except that it arises from
the proximal part of the posterior (rather than the medial) division, In T.p.
8R (fig. 11B), the branch to M. gastrocnemius pars externa arises so far
proximally that it appears as a separate (fifth) division of the tibial nerve.
In two legs, the branch of the medial division that supplies M. gastrocnemius
pars media sends a twig into the distal end of M. femorocruralis (fig. 11A).
T. cupido
INpIvipvAL VARIATION.—!In one leg, an extra branch of the medial division
arises immediately distal to the branch to M. gastrocnemius pars media and
enters the proximal end of the medial head of M. flexor perforatus digiti III.
In one instance, the branch to M. gastrocnemius pars interna passes through
a gap in the origin of M. plantaris rather than distal to the origin of the
latter,
P. p. jamesi
InpivipvaL VaRIaATION.—The branch to M. gastrocnemius pars interna
gives a minute twig to the deep surface of the free belly of M. plantaris
in one leg.
886 Unrversiry OF Kansas Pusts., Mus. Nar. Hist.
t
last movable rib
to abdominal wall muscles
i
I.
1!
|
vertebral column =
> parrapoph ysis
eorece®
oe?
*.
.
.
.
.
.
«
2 = to abdominal wall muscles
3 |
Ugo
Gs
4|
[0 = femoral nerve
furcal nerve——=— 5 I to abdominal wall muscles
6 tn
7h: obturator nerve
8
sciatic nerve
to pudendal plexus
to abdominal wall muscles
femoral nerve
to abdominal wall muscles
obturator nerve
sciatic nerve
AS to pudendal plexus
{| =
Fic. 2. Ventral views of the lumbosacral plexus of Tympanuchus palli-
dicinctus. Sympathetic ganglionated chain removed. Numbers indicate
synsacral spinal nerves. X 2. A. T.p.1L. B. T.p. 2L.
MuscLEs AND NERVES OF LEG OF GROUSE 387
to abdominal wall muscles
to abdominal wall muscles
femoral nerve
furcal nerve
obturator nerve
ee sciatic nerve
to pudendal plexus
~~
to abdominal wall muscles
bw
o
femoral nerve
to abdominal wall muscles
obturator nerve
sciatic nerve
to pudendal plexus
Fic. 3. Ventral views of the lumbosacral plexus. Sympathetic gan-
glionated chain removed. Numbers indicate synsacral spinal nerves.
x 2. A. Tympanuchus cupido pinnatus 3L. B. Pedioecetes phasianellus
jamesi 4L.
388
UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
anterodorsal division I 4 2
dorsal division ee 3
anterior division
posterodorsal division
middle division
posterior division
\
level of inguinal er EE)
\ 17
Fic. 4. Semidiagrammatic ventral views of the femoral
nerve, showing the distribution of the branches. x 8. 1,2,
M. extensor iliotibialis anticus; 8, cutaneous; 4-6, M. ex-
tensor iliotibialis lateralis; 7,8, M. iliacus; 9, M. gluteus
profundus; 10-12, fused Mm. vastus lateralis and vastus
medialis; 13,14, M. vastus medialis; 15, M. ambiens;
16, M. femoritibialis internus; 17, nonmuscular; 18, M.
psoas; 19, M. iliotrochantericus medius. A. Tympanu-
chus cupido pinnatus 8L. B. Pedioecetes phasianellus
jamesi 3L.
MuscLES AND NERVES OF LEG OF GROUSE 889
Fic. 5. Semidiagrammatic ventral views
of the femoral nerve, showing the distri-
bution of the branches. X38. 1,2, M.
extensor iliotibialis anticus; 8, cutaneous;
5,6, M. extensor iliotibialis lateralis; 7,8,
M. iliacus; 9, M. gluteus profundus;
10,11, fused Mm. vastus lateralis and vas-
tus medialis; 13, M. vastus medialis; 15,
M. ambiens; 16, M. femoritibialis in-
ternus; 17, nonmuscular; 18, M. psoas;
19, M. iliotrochantericus medius. A. Tym-
panuchus pallidicinctus 2L. B. Tym-
panuchus cupido attwateri 1R.
390
UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
8
Se: peroneal nerve
tibial nerve
j
Fic. 6. Semidiagrammatic dorsolateral view of the sci-
atic nerve of Pedioecetes phasianellus jamesi 8R, show-
ing the distribution of the branches. X 2%. 1, M. glu-
teus profundus; 2, M. piriformis; 38, M. extensor
iliotibialis lateralis; 4-7, M. extensor iliofibularis; 8, M.
flexor cruris medialis; 9, cutaneous; 10, to pudendal
plexus; 11, M. flexor cruris lateralis; 12, M. caudo-
femoralis pars caudifemoralis; 18-15, M. caudofemoralis
pars iliofemoralis; 16,17, M. flexor ischiofemoralis;
18,19, M. femorocruralis (branch of tibial nerve); 20,
cutaneous; 21, M. gastrocnemius pars media (branch of
tibial nerve); 22, cutaneous.
paraperoneal branch
of tibial nerve
21 20
MUSCLES AND NERVES OF LEG OF GROUSE 391
V7 last root
posterior tibial division
level of emergence from
eae ilio-ischiatic fenestra
17
middle tibial division
NN anterior tibial division
dorsal peroneal division
anterior peroneal division
1
2
‘middle peroneal division
posterior peroneal division Al
7
10
3 13 12
11
9
\
8
peroneal nerve tibial nerve
8
|
: !
paraperoneal branch |
of tibial nerve ;
\
Fic. 7. Semidiagrammatic dorsolateral view of the sciatic nerve of
Tympanuchus pallidicinctus 2L, showing the distribution of the branches.
< 2%. 1, M. gluteus profundus; 2, M. piriformis; 3, M. extensor ilio-
tibialis lateralis; 4,7, M. extensor iliofibularis; 8, M. flexor cruris
medialis; 9, cutaneous; 10, to pudendal plexus; 11, M. flexor cruris
lateralis; 12, M. caudofemoralis pars caudifemoralis; 138-15, M. caudo-
femoralis pars iliofemoralis; 17, M. flexor ischiofemoralis; 18, M. fem-
orocruralis (branch of tibial nerve); 22, cutaneous; 23, nonmuscular
(branch of peroneal nerve).
892 University OF Kansas Pusts., Mus. Nat. Hist.
tibial nerve
paraperoneal branch
e of tibial nerve
18
Fic. 8. Semidiagrammatic dorsolateral view of the
sciatic nerve of Tympanuchus cupido pinnatus 8L,
showing the distribution of the branches. > 2%.
1, M. gluteus profundus; 2, M. piriformis; 3, M.
extensor iliotibialis lateralis; 4,7, M. extensor ilio-
fibularis; 8, M. flexor cruris medialis; 9, cuta-
neous; 11, M. flexor cruris lateralis; 12, M. caudo-
femoralis pars caudifemoralis; 13, M. caudofemo-
ralis pars iliofemoralis; 17, M. flexor ischiofem-
oralis; 18, M. femorocruralis (branch of tibial
nerve); 20, cutaneous; 22, cutaneous.
- Muscies AND Nerves OF LEG OF GROUSE
peroneal nerve
tibial nerve
+————— paraperoneal branch
ee |
of tibial nerve
Fic. 9. Semidiagrammatic dorsolateral view of the
sciatic nerve of Pedioecetes phasianellus jamesi 3L,
showing the distribution of the branches. x 2%.
1, M. gluteus profundus; 2, M. piriformis; 38, M.
extensor iliotibialis lateralis; 4,5,7, M. extensor
iliofibularis; 8, M. flexor cruris medialis; 9, cu-
taneous; 11, M. flexor cruris lateralis; 13,14, M.
caudofemoralis pars iliofemoralis; 16,17, M. flexor
ischiofemoralis; 18,19, M. femorocruralis (branch
of tibial nerve); 20, cutaneous; 22, cutaneous.
893
394 UNIVERSITY OF Kansas Pusxs., Mus. Nat. Hist.
deep peroneal superficial
branch peroneal
5 ff 0 J) }
Fi i]
2 Ti
6
~
superficial
peroneal deep peroneal
branch branch
18 1
0
9
deep peroneal =a superficial
| peroneal branch
branch
A
2
1
17
ie
15 14 13
Cc
Fic. 10. A,B. Semidiagrammatic drawings of the peroneal nerve
of Tympanuchus pallidicinctus 1L, showing the distribution of the
branches. X 2. C. Semidiagrammatic drawing of the distal part
of the peroneal nerve of Tympanuchus cupido attwateri 1R, show-
ing the distribution of the branches. xX 2. 1,2, M. tibialis anticus
(tibial head); 98,4, M. tibialis anticus (femoral head); 5, M.
extensor digitorum longus; 6, nonmuscular; 7,8, M. peroneus
longus; 9, M. peroneus brevis; 10,11, M. extensor hallucis longus
(proximal head); 12, M. extensor hallucis longus (distal head);
18-15, nonmuscular (to toes); 16, M. abductor digiti II; 17, M.
extensor brevis digiti III; 18, M. extensor brevis digiti IV.
MuscLes AND NERVES OF LEG OF GROUSE 395
lateral division
posterior division
KK
29
35 25 B
Fic. 11. A,B. Semidiagrammatic drawings of the tibial nerve (excluding the
paraperoneal branch) of Tympanuchus pallidicinctus, showing the distribution
of the branches. x 2. A. T.p. 1L. B. T.p. 3R. C. Semidiagrammatic draw-
ing of the distal part of the paraperoneal branch of the tibial nerve of Pedio-
ecetes phasianellus jamesi 2L, showing the distribution of the branches. x 2.
1, M. femorvocruralis; 2, M. gastrocnemius pars media; 3, M. popliteus; 4, M.
plantaris; 5, M. flexor digitorum longus; 6-8, nonmuscular; 9-11, M. gastroc-
nemius pars interna; 12,13, M. flexor hallucis longus; 14-16, M. flexor per-
foratus digiti IV (medial head); 17, M. flexor perforatus digiti III (medial
head); 18-20, M. flexor perforatus digiti II; 21, M. flexor perforatus digiti IV
(lateral head); 22-24, M. flexor perforatus digiti IV (anterolateral head); 25,
M. flexor perforatus digiti III (anterolateral head); 26, M. flexor perforans et
perforatus digiti III; 27,28, M. flexor perforans et perforatus digiti II; 29, M.
gastrocnemius pars externa; 30,31, M. abductor digiti IV; 32,33, M. flexor
hallucis brevis; 34,85, nonmuscular (to toes).
38—5835
896 UNIVERSITY OF Kansas Pupsts., Mus. Nat. Hist.
MUSCLES
In the accounts of the muscles the name used by Hudson, et al.
(1959) for each muscle is given in parentheses after the name
used by me if the two differ.
In the account of each muscle, the description of the condition
found in most specimens of the Lesser Prairie Chicken (T. palli-
dicinctus ) is given first. This is hereafter referrred to as the typical
condition for T. pallidicinctus. Then any individual variations
found within this species are given. Under the heading T. cupido
any constant differences between this species and typical T. palli-
dicinctus are given first, and any individual variations found within
the species T. cupido (both subspecies considered together) are
given second. Under the heading P. p. jamesi any constant differ-
ences between this subspecies and the typical condition for T.
pallidicinctus (thus these differences are not necessarily constant
between the two genera) are given first, and any individual varia-
tions found within the subspecies P. p. jamesi are given second.
In the bird embryo, according to the studies of Romer (1927)
and Wortham (1948), the muscles within each segment of the
leg differentiate from distinct dorsal or ventral mesenchymal
masses. Presumably these represent the primitive dorsal extensor
and ventral flexor muscle masses. The list below indicates the
ontogenetic origin of the avian leg muscles, according to the
studies of Romer and Wortham. The individual muscles are
discussed in the order in which they are listed below.
Dorsal muscles of thigh
M. extensor iliotibialis lateralis M. extensor iliofibularis
M. extensor iliotibialis anticus M. piriformis
M. ambiens M. gluteus profundus
M. vastus lateralis M. iliacus
M. vastus medialis M. iliotrochantericus medius
M. femoritibialis internus M. psoas
Ventral muscles of thigh
M. flexor cruris lateralis M. adductor superficialis
M. flexor cruris medialis M. adductor profundus
M. caudofemoralis M. obturator
M. flexor ischiofemoralis M. femorocruralis
Ventral muscles of shank
M. gastrocnemius M. flexor perforatus digiti III
M. flexor perforans et perforatus M. flexor perforatus digiti II
digiti II M. flexor hallucis longus
M. flexor perforans et perforatus M. plantaris
digiti III M. flexor digitorum longus
M. flexor perforatus digiti IV M. popliteus
MuscLes AND NeErvES oF LEG oF GROUSE 397
Dorsal muscles of shank .
M. peroneus longus M. extensor digitorum longus
M. tibialis anticus M. peroneus brevis
Dorsal muscles of foot
M. extensor hallucis longus M. extensor proprius digiti III
M. abductor digiti II M. extensor brevis digiti IV
M. extensor brevis digiti III
Ventral muscles of foot
M. lumbricalis (M. adductor digiti II—not present)
M. abductor digiti IV (M. adductor digiti IV—not pres-
M. flexor hallucis brevis ent)
extensor iliotibialis anticus
extensor iliotibialis
lateralis
flexor cruris
lateralis
pp caudofemoralis
pars caudifemoralis
extensor iliofibularis
flexor cruris medialis
gastrocnemius pars interna
gastrocnemius pars externa
flexor perforans et perforatus digiti HI
flexor perforans et perforatus digiti I
peroneus longus
Fic. 12. Tympanuchus pallidicinctus 2L. Lateral view of the superficial
muscles of the left leg. x 1.
898 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist.
M. Extensor Iliotibialis Lateralis (M. iliotibialis), Figs. 12, 18, 20F, G
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Most superficial muscle on lateral
surface of thigh; broad, flat, and triangular; bounded anteriorly by M. ex-
tensor iliotibialis anticus and posteriorly by M. flexor cruris lateralis; posterior
part considerably thicker than anterior part; anteroproximal and centrodistal
parts aponeurotic; extreme posteroproximal corner also aponeurotic (could be
considered tough sheet of connective tissue intimately fused with M. ex-
tensor iliotibialis lateralis, rather than part of muscle itself; see fig. 20F);
latter aponeurosis, as well as adjacent fleshy fibers, overlapped by M. flexor
cruris lateralis; this aponeurosis fused with posterior end of underlying M.
caudofemoralis pars iliofemoralis; centrodistal aponeurosis tightly fused to
underlying Mm. vastus lateralis and vastus medialis; fleshy fibers posterior to
this aponeurosis also fused with M. vastus lateralis, although posterior third of
extensor iliotibiclis anticus
jlictcus
extensor iliotibialis lateralis
vastus medialis
psoas
ambiens
femorotibialis internus
adductor profundus
caudofemoralis
flexor cruris medialis
gastrocnemius pars media:
gastrocnemius pars externa
flexor perforatus digiti III (medial head)
‘gastrocnemius pars interna
peroneus longus
Fic. 18. Tympanuchus pallidicinctus 2L.. Medial view of the super-
ficial muscles of the left leg. 1. Articular capsule shown by concen-
trically arranged dashes.
Musc.LEs AND NERVES OF LEG OF GROUSE 899
muscle free; fleshy part anterior to this aponeurosis bound by tough connective
tissue to underlying M. vastus medialis, although no fusion of fibers; an-
terior edge tightly bound by strong connective tissue to M. extensor iliotib-
ialis anticus, with some fusion of fibers (proximally); posteroproximal corner
bound by tough connective tissue to adjacent muscles; anteroproximal
aponeurosis fused with aponeurotic anteroproximal part of underlying M.
extensor iliofibularis. Continuous proximal aponeurosis of M. extensor il-
iotibialis anticus and of M. extensor iliotibialis lateralis underlain by tough
fascial sheet overlying M. gluteus profundus; anterior part of this fascia tightly
fused to latter muscle but free from overlying aponeurosis; posterior part of
this fascia tightly fused to overlying aponeurosis but free from M. gluteus
profundus; middie part of fascia fused to both aponeurosis and M. gluteus
profundus.
gluteus profundus
iliacus
vastus medialis
> vastus lateralis pars lateralis
adductor superficialis
ze Siiextensor iliofibularis
{ LAA
E9117 4
ny YY y U.S femorocruralis
WGi74,Z EY vastus lateralis pars postica
Wye Mj
QQ Sy flexor cruris medialis
i guide loop for extensor iliofibularis
SSN AS medial head pire
SS ES Gichall ase } flexor perforatus digiti IV
\ USS -flexor perforans et perforatus digiti III
WS posterior head } flexor perforans et
anterior head perforatus digiti II
tibialis anticus
medial head \ flexor perforatus
anterolateral head digiti ITI
flexor digitorum longus
flexor perforans et perforatus
KEN digiti III tendon
peroneus brevis
Fic. 14. Tympanuchus pallidicinctus 21. Lateral view of the muscles of
the left leg. The following muscles have been removed: extensor ilio-
tibialis lateralis, extensor iliotibialis anticus, gastrocnemius pars externa
and pars interna, and peroneus longus. X l.
400 UnIvERSITY OF Kansas Pusts., Mus. Nat. Hust.
Oricin.—Approximately the anterior half attaches by an extensive aponeu-
rosis, which is continuous anteriorly with that of M. extensor iliotibialis anticus,
to the anterior iliac crest, ending posteriorly at the anterior end of the lateral
iliac process; the posterior part attaches fleshily to the edge of the entire
lateral iliac process and (posterior few mm.) aponeurotically to the entire
lateral ischiate ridge. The proximal part of the belly is much thicker than
the fleshy origin. Two accessory aponeuroses associate with the anterior
part of the muscle; the proximal one of these comes off the deep surface several
mm. distal to the proximal end of the fleshy belly and passes medially be-
tween Mm. gluteus profundus and iliacus, fusing to both these muscles,
and attaches to the lateral edge of M. iliotrochantericus medius and to the
lateral edge of the ilium anterior to the latter; the aponeurosis actually splits
into two sheets at the edge of M. iliotrochantericus medius; these sheets fuse
to the dorsal and ventral surfaces of the latter muscle, enclosing it; the
part of this aponeurosis between Mm. iliacus and iliotrochantericus medius is
strongly fused with the underlying body wall. The distal accessory aponeu-
iliacus
psoas
vastus medialis
adductor profundus
flexor cruris medialis
caudofemoralis
pars ies NN : y
femorotibialis intemus
femorocruralis
flexor cruris lateralis
gastrocnemius pars media
flexor cruris medialis tendon
plantaris
flexor perforatus digiti III (medial head)
tibialis anticus
flexor hallucis longus
extensor digitorum longus
flexor perforatus digiti II
Fic. 15. Tympanuchus pallidicinctus 21. Medial view of the muscles
of the left leg. The following muscles have been removed: extensor
iliotibialis lateralis, extensor iliotibialis anticus, ambiens, flexor cruris
lateralis (in part), flexor cruris medialis (in part), gastrocnemius pars
externa and pars interna, and peroneus longus. X 1
MuscLes AND NERVES OF LEG OF GROUSE 401
rosis (sometimes weak) comes off the deep surface several mm. distal to the
proximal one and passes medially along the ventral surface of M. iliacus,
fusing with the latter, then joining the proximal accessory aponeurosis medial
to M. iliacus.
INsERTION.—The muscle inserts by a broad aponeurosis strongly fused to
the underlying Mm. vastus lateralis and vastus medialis; the aponeurosis
contributes superficially to the patellar tendon, attaching to the lateral half of
the rotular crest,
INNERVATION.—A variable number of branches (usually two) of the middle
division of the femoral nerve pass ventral to M. iliacus and between Mm.
extensor iliotibialis anticus and vastus medialis and enter the deep surface of
the anteroproximal part of the muscle. The branch of the middle peroneal
gluteus profundus
iliacus
piriformis
bturator pars postica
aed . :
ek oa Sec ischiofemoralis
“sadductor superficialis
caudofemoralis
pars iliofemoralis
pars caudifemoralis
vastus lateralis pars postica
flexor cruris medialis
femorocruralis
flexor cruris lateralis
guide loop for extensor iliofibularis
ambiens tendon
femoral head
tibial head
medial head
lateral head | flexor perforatus digiti IV
anterolateral head
tibialis anticus
flexor digitorum longus
medial head
flexor perforatus
anterolateral head eee
digiti III
peroneus brevis
Fic. 16. Tympanuchus pallidicinctus 2L. Lateral view of the muscles of the
left leg. The following muscles, in addition to those listed for Fig. 14, have
been removed: ambiens, vastus lateralis pars lateralis, vastus medialis (except
for part of patellar tendon), extensor iliofibularis, flexor cruris lateralis (in eon
flexor perforans et perforatus sigs ae and flexor perforans et perforatus digiti
Fula i
402 University OF Kansas Pusts., Mus. Nat. Hist.
division of the sciatic nerve emerges between the proximal ends of Mm. exten-
sor iliofibularis and vastus lateralis and sends twigs into the deep surface of
M. extensor iliotibialis lateralis.
InpivipvAL VaRIATION.—In two legs, the nerve supplying M. extensor
iliotibialis anticus gives twigs into M. extensor iliotibialis lateralis.
T. cupido
DIFFERENCES From T. pallidicinctus—The fleshy origin from the lateral
iliac process is considerably thicker (reflected in a thicker lateral iliac process).
INDIVIDUAL VARIATION.—In three legs the nerve supplying M. extensor
iliotibialis anticus gives twigs into M. extensor iliotibialis lateralis. In another
leg one of the branches to the fused Mm. vastus lateralis and vastus medialis
sends a twig into M. extensor iliotibialis lateralis.
iliacus
iliotrochantericus medius
obturator pars postica
flexor
ischiofemoralis
adductor
superticialis
adductor profundus
flexor cruris medialis
femorocruralis
flexor cruris lateralis
guide loop for extensor iliofibularis
ambiens tendon
medial head naib:
lateral head | flexor perforatus digiti I
anterolateral head
flexor digitorum longus
extensor digitorum longus
medial head } a
SS SAcnterolateral head flexor perforatus digi
\
NY
\\\ NN
Ae : peroneus brevis
Fic. 17. Tympanuchus pallidicinctus 2L. Lateral view of the muscles of
the left leg. The following muscles, in addition to those listed for Fig. 16,
have been removed: vastus lateralis pars postica, gluteus profundus, flexor
cruris medialis (in part), caudofemoralis, flexor perforatus digiti IV, and
tibialis anticus. * 1.
MuscLes AND NERVES OF LEG OF GROUSE 403
P. p. jamesi
DiFFERENCES FrRoM TypicaL T._ pallidicinctus——The posteroproximal
aponeurosis is more extensive, resulting in a narrower proximal fleshy end
(fig. 20G); the fleshy fibers adjacent to this aponeurosis are not overlapped
by M. flexor cruris lateralis. There is a fusion of fibers between the an-
terodistal fleshy part of M. extensor iliotibialis lateralis and the underlying M.
vastus medialis, but there is no fusion of fibers between the anterior edge of
M. extensor iliotibialis lateralis and M. extensor iliotibialis anticus. The con-
nective tissue binding the posteroproximal corner to adjacent muscles is
stronger. The fleshy part of the origin is narrower, partly tendinous, and
psoas
adductor profundus
peroneus brevis
Fic. 18. Tympanuchus pallidicinctus 21.. Lateral view of the muscles
of the left leg. The following muscles, in addition to those listed for
Fig. 17, have been removed: patellar tendon, iliacus, iliotrochantericus
medius, flexor cruris lateralis, flexor cruris medialis, flexor ischiofemoralis,
adductor superficialis, femorocruralis, gastrocnemius pars media, flexor
perforatus digiti III, flexor perforatus digiti II, flexor hallucis longus,
plantaris, flexor digitorum longus, popliteus, and extensor digitorum
longus. X 1.
404 UNIVERSITY OF Kansas Pusts.,; Mus. Nat. Hist.
{ |
longus
~obturator
A posterior head
NAMI lateral head & pars antica
i
\\
\\ i
plantaris Cc
flexor hallucis longus
SS
obturator
pars postica
insertion of tibialis anticus
flexor hallucis
brevis
abductor digiti IV
extensor brevis digiti IV
extensor brevis
digiti III
extensor brevis digiti III
abductor digiti II
lumbricalis
flexor hallucis brevis
tendon
extensor brevis digiti IV subarticular
tendon cartilage
Fic. 19. Tympanuchus pallidicinctus 21. A. Posterior view of the muscles of the left
shank. The following shank muscles, in addition to those listed for Fig. 17, have been
removed: gastrocnemius pars media, flexor perforatus digiti III, and flexor perforatus
digiti II. Xx 1. B. Posterior view of the proximal end of the shank, showing the most
deeply situated muscle. 1. C. Lateral view of the head of the left femur and the
middle part of the pelvis, showing the deepest part of M. obturator. 1. D. Medial
view of the posteroventral part of the left side of the pelvis, showing the intrapelvic part
of M. obturator. 1. E. Anterior view of the left tarsometatarsus, showing the dorsal
intrinsic muscles of the foot. 1%. F. Posterior view of the left tarsometatarsus, show-
ing the ventral intrinsic muscles of the foot. x 14.
MuscLEs AND NERVES OF LEG OF GROUSE 405
much thinner (reflected in a thin lateral iliac process). The proximal border
is much more nearly straight, owing to a less pronounced lateral iliac process.
The distal accessory aponeurosis is absent.
INDIvmUAL VARIATION.—The muscle is usually somewhat fused to the
posteroproximal and anteroproximal fleshy corners of the underlying M. ex-
tensor iliofibularis.
M. Extensor Iliotibialis Anticus (M. sartorius), Figs. 12, 13
T. pallidicinctus
GENERAL DescripTION AND RELATIONS.—Anteriormost muscle of thigh;
long and strap-shaped; proximal part entirely anterior (adjacent) to M.
extensor iliotibialis lateralis; posterior edge of middle part medial to latter
muscle; distal part mostly medial to Mm. extensor iliotibialis lateralis and
vastus medialis; proximal part aponeurotic, continuous posteriorly with
anteroproximal aponeurosis of M. extensor iliotibialis lateralis; anterior edge
of M. extensor iliotibialis lateralis bound by strong connective tissue to
adjacent part of M. extensor iliotibialis anticus; some fusion of fibers (proxi-
mally) between these two muscles; anteroproximal corner of fleshy part of
muscle sometimes fused to underlying anterior edge of ilium and fascia
covering body wall musculature adjacent (anterior) to ilium.
OriciIn.—The muscle arises aponeurotically from the anterior part of
the anterior iliac crest and (anteroproximal corner) from the anterior end
of the median dorsal ridge.
INSERTION.—The flat tendon, continuous posteriorly with the superficial
tendon of M. femoritibialis internus, fuses to the tendon of M. vastus medialis,
contributing superficially to the medial part of the patellar tendon, which
attaches to the medial half of the rotular crest; most of the tendon is over-
lapped by the edge of M. gastrocnemius pars interna.
INNERVATION.—A branch of the anterior division of the femoral nerve
gives twigs into the lateral surface of the posterior part.
INDIVIDUAL VARIATION.—In two legs, a twig from the anteriormost branch
of the middle division of the femoral nerve anastomoses with the typical
branch to M. extensor iliotibialis anticus.
T. cupido
InprivipvaL VarrATion.—In several legs, the anterior edge of origin
extends forward onto the neural spine of the last free thoracic vertebra. A
twig from the middle division of the femoral nerve anastomoses with the
typical branch to M. extensor iliotibialis anticus in three legs.
P. p. jamesi
DIFFERENCES FRoM TypicaL T. pallidicinctus—There is no fusion of
fibers between M. extensor iliotibialis anticus and M. extensor _iliotibialis
lateralis.
INDIVIDUAL VaRIATION.—The anterior edge of origin extends forward onto
the neural spine of the last free thoracic vertebra in some legs.
406 UniversITY OF KANSAS Pusts., Mus. Nat. Hist.
pars dorsalis
pars postica
pars ventralis
pars antica
origin of pars dorsalis
origin of pars antica
L
area covered
by pars
postica
extensor brevis digiti IV
insertion of tibialis anticus
extensor hallucis longus
proximal head
distal head
extensor proprius digiti IIT
abductor digiti II
extensor brevis digiti III
Ficure 20. Explanation on opposite page.
Musc.Les AND NERVES OF LEG OF GROUSE 407
EXPLANATION OF FIGURE 20
A-D. Dorsal views of M. iliotrochantericus medius, showing its relation-
ship to femoral notch. X 1. In D, note absence of femoral notch and loca-
tion of branch of femoral nerve. A. Tympanuchus pallidicinctus 2L. B.
T. cupido pinnatus 4L. C. Pedioecetes phasianellus jamesi 1L. D. T. palli-
dicinctus 3L.
E. Medial view of distal end of M. flexor cruris medialis of P. p. jamesi 4L.
<1. Part of insertion is covered by medial collateral ligament.
F,G. Lateral views of posteroproximal corner of M. extensor iliotibialis
lateralis (removed from specimen), x1. F. T. pallidicinctus 2L. G. P. p.
jamesi 3L.
H,I. Dorsolateral views of M. piriformis. x1. H. P. p. jamesi 1L. I.
T. cupido attwateri 1L.
J. Lateral view of M. caudofemoralis pars caudifemoralis (removed from
specimen) of T. c. pinnatus4L. xX 1.
K. Lateral view of extrapelvic part of M. obturator of T. pallidicinctus 3L
(bones not shown). X 2.
L,M. Region surrounding obturator foramen of T. pallidicinctus 3L, show-
ing points of attachment of three parts of M. obturator (muscles removed).
x 3. L. Lateral view. M. Medial view.
N. Anterior view of left tarsometatarsus of P. p. jamesi 4L, showing dorsal
intrinsic muscles of foot. >< 1%. Tendon of M. extensor digitorum longus has
been removed.
408 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
M. Ambiens, Figs. 13, 16, 17
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Thin and elongate; on medial
surface of thigh; broadest above middle of belly; belly narrowed distally,
forming long slender tendon passing lateral to distal part of M. extensor
iliotibialis anticus; bounded anterolaterally by M. vastus medialis and postero-
laterally by Mm. femoritibialis internus and psoas (proximally).
Oricin.—The muscle arises by a short flat tendon from the pectineal
process.
INSERTION.—The long slender tendon enters an elongate channel within
the patellar tendon; the point of entrance is at the proximal end of the latter
tendon just medial to the patella; the tendon passes distolaterally (within the
channel) below the patella and emerges from the distolateral edge of the
patellar tendon and then extends distally along the anterolateral surface of
the head of the fibula, superficial to the fibular arm of the guide loop for
M. extensor iliofibularis, and joins the anterolateral surface of the common
tendon of origin of the anterolateral heads of Mm. flexor perforatus digiti
III, flexor perforatus digiti IV, and flexor perforatus digiti II; the point of
junction is usually immediately proximal to the proximal end of the lateral
head of M. flexor digitorum longus.
INNERVATION.—The branch of the middle division of the femoral nerve
that supplies M. femoritibialis internus gives off a tiny twig or twigs that
penetrate the lateral surtace of the proximal part of M. ambiens.
INDIvipUAL VARIATION.—None of significance in T. pallidicinctus or in
P. p. jamesi; in T. cupido the origin is partly fleshy in one leg.
M. Vastus Lateralis (M. femoritibialis externus + part of M. femoritibialis
medius), Figs. 14, 16
Fisher and Goodman (1955) apply the name femoritibialis externus to
the muscle unit that I here term the pars postica of M. vastus lateralis. The
reasons for this change are discussed in the section on terminology.
T. pallidicinctus
GENERAL DeEscriPTION AND RELATIONS.—Thick; on lateral surface of
femur deep to M. extensor iliotibialis lateralis; anterior to M. extensor ilio-
fibularis and lateral to M. vastus medialis; much of lateral surface, except
proximal part, fused with overlying M. extensor iliotibialis lateralis; deep
surface of anterior half fused with M. vastus medialis; proximal part over-
lapping, but usually not fusing with, insertions of Mm. iliacus and caudo-
femoralis; partially separable into two parts—pars lateralis and pars postica,
former constituting main part of muscle; latter considerably smaller and
situated deep to posterodistal part of pars lateralis, except for posterodistal
part extending posterior to edge of pars lateralis; proximal part of pars
postica strongly fused with pars lateralis; posterodistal tendinous edge of
pars lateralis fused or not fused with lateral surface of pars postica; proximal
end (narrow) of pars postica tendinous and variable in length.
Oricin.—Pars lateralis: This arises fleshily from most of the lateral surface
and (distally) from the anterior surface of the femur, extending anteriorly to
MuscLes AND NERVES OF LEG OF GROUSE 409
the anterior intermuscular line, fusing with M. vastus medialis, and extending
posteriorly to the posterolateral intermuscular line (proximally) and the origin
of pars postica (distally); the proximal end begins at the level of the distal
edge of the insertion of M. iliotrochantericus medius, contacting the insertions
of Mm. iliotrochantericus medius, piriformis, and flexor ischiofemoralis, and
terminates distally at the level of the proximal ends of the femoral condyles.
Pars postica: This arises fleshily and tendinously (proximal end and deep
surface) from the posterolateral surface of approximately the distal half of the
femur, extends posteromedially to the posterolateral intermuscular line where
it contacts the origin of M. femorocruralis, and extends anteriorly to the level
of a line drawn diagonally across the femur from the proximal end of the
origin (at the posterolateral intermuscular line) to the proximal end of the
external condyle; the distal end is anterior (adjacent) to the attachment of the
proximal arm of the tendinous guide loop for M. extensor iliofibularis; the
origin is adjacent to, but distinct from, the origin of pars lateralis.
INSERTION.—Pars lateralis is fused indistinguishably with M. vastus medialis;
these two muscles form the main (middle) part of the patellar tendon, which
also receives contributions from pars postica and Mm. femoritibialis internus,
extensor iliotibialis lateralis, and extensor iliotibialis anticus; the patellar tendon
attaches to the entire rotular crest of the tibia; the patella is situated in the
proximal part of this tendon; some deep fleshy fibers of M. vastus lateralis pars
lateralis and M. vastus medialis attach to the proximal edge of the patella.
Pars postica forms a short narrow tendon that fuses to the lateral part of the
tendon of pars lateralis, forming the lateralmost part of the patellar tendon.
A broad flat vinculum extends from the lateral surface of the femorofibular
fascia (defined under M. flexor perforans et perforatus digiti Il) to the deep
surface of the lateral part of the patellar tendon; a similar vinculum extends
from the medial surface of the internal condyle to the deep surface of the
medial part of the patellar tendon.
INNERVATION.—Two or more branches of the middle division of the femoral
nerve penetrate the anterior surface of the fused Mm. vastus lateralis (pars
lateralis) and vastus medialis; short twigs emerge from the deep surface of
pars lateralis and penetrate the superficial surface of the anteroproximal part
of pars postica.
INDIVIDUAL VARIATION.—The proximal ends of M. vastus medialis and M.
vastus lateralis are usually separated by a deep notch. In some legs, a small
bundle of fibers forming the anteroproximal part of M. vastus lateralis attaches
to the lateral surface of M. vastus medialis anterior to this notch.
T. cupido
INDIVIDUAL VARIATION.—One leg shows the same variation found in T.
pallidicinctus (see above). In several legs, pars lateralis does not extend so
far proximally as usual, but begins at the level of insertion of M. piriformis
(does not contact the insertion of M. iliotrochantericus medius) and may not
overlap M. iliacus. In a few legs, no vincula are associated with the patellar
tendon.
P. p. jamesi
INDIvinUAL VARIATION.—Pars lateralis often begins proximally at the level
of the insertion of M. piriformis.
410 UNIVERSITY OF Kansas PuButs., Mus. Nat. Hist.
M. Vastus Medialis (Part of M. femoritibialis medius), Figs. 13, 14, 15
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Thick; on anteromedial surface
of femur medial to anterior part of M. vastus lateralis pars lateralis; bounded
medially by Mm. ambiens and extensor iliotibialis anticus (distally); bounded
posteromedially by M. femortibialis internus; proximal part medial to posterior
ends of Mm. iliacus, iliotrochantericus medius, and gluteus profundus; lateral
surface, except proximal part, fused with anterior part of M. vastus lateralis
pars lateralis; part of lateral surface of M. vastus medialis covered by sheet
of fascia attaching to anterior intermuscular line; M. vastus lateralis separable
from this fascia, but fascia absent anteriorly and distally and these two muscles
indistinguishably fused.
Oricin.—The proximal third is attached narrowly by its lateral edge; the
distal two thirds is attached broadly by its entire deep surface. The proximal
third arises tendinously from the trochanteric ridge and the proximal end of the
anterior intermuscular line and fleshily from a narrow area of the femur ad-
jacent (medial) to the latter; the distal part arises tendinously from the
anterior intermuscular line and fleshily from a broad adjacent area on the
anteromedial surface of the femur, terminating distally at the level of the
proximal end of the internal condyle; the posterior edge contacts the origin of
M. femoritibialis internus.
INSERTION.—Attachment is in common with M. vastus lateralis pars lateralis,
which see.
INNERVATION.—Two or more branches of the middle division of the femoral
nerve penetrate the anterior surface of the fused Mm. vastus medialis and
vastus lateralis pars lateralis; a variable number of branches of the same divi-
sion penetrate the medial surface of the proximal part of M. vastus medialis.
INprvipuAL VARIATION.—None of significance in any of the three species
studied.
M. Femoritibialis Internus, Figs. 13, 15
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Elongate; on posteromedial sur-
face of femur; bounded anteriorly by M. vastus medialis and posteriorly by M.
adductor profundus (overlapping anterior edge of latter); anteroproximal
part lateral to M. ambiens; anterodistal corer deep to distal end of M. ex-
tensor iliotibialis anticus; distal part of muscle split into superficial and deep
layers; superficial layer thin, narrow, and tendinous except for proximal end;
deep layer wider, much thicker, and fleshy except for distal end taking form
of flat tendon; anterior edge of latter somewhat fused to medial edge of tendon
of M. vastus medialis; deep layer widest near distal end of fleshy part; posterior
edge of superficial layer fused to underlying deep layer, and anterior edge
fused to (continuous with) posterior edge of tendon of M. extensor iliotibialis
anticus.
Oricin.—The origin is mostly fleshy from the posteromedial surface of the
femur between the origin of M. vastus medialis and the posterior intermuscular
line, terminating immediately proximal to the internal condyle.
MusciLes AND NERVES OF LEG OF GROUSE All
INSERTION.—The tendons of both superficial and deep layers attach to the
medial part of the rotular crest, forming the medialmost part of the patellar
tendon.
INNERVATION.—The posteriormost branch of the middle division of the
femoral nerve penetrates the medial surface of the muscle near the proximal
end.
INDIVIDUAL VARIATION.—None of significance in any of the three species
studied.
M. Extensor Iliofibularis (M. biceps femoris), Figs. 12, 14, 16, 17
The term extensor in the name of this muscle does not refer to its function.
Howell (1938) used the term extensor to indicate derivation of the muscle
from the primitive dorsal extensor muscle mass. (Likewise he used the term
flexor to indicate derivation from the primitive ventral flexor muscle mass.)
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Deep to M. extensor iliotibialis
lateralis and posterior to femur; broad proximally and narrow distally; posterior
to M. vastus lateralis and anterior to proximal part of M. flexor cruris lateralis
(superficial to distal part of latter); anteroproximal part aponeurotic, fused to
deep surface of aponeurosis of M. extensor iliotibialis lateralis; proximal part
of aponeurosis of M. extensor iliofibularis also fused to dorsal edges of under-
lying Mm. gluteus profundus and piriformis.
Ornicin.—The posterior part is fleshy from the ventromedial surface of
the entire lateral iliac process; the anterior part is aponeurotic from the pos-
terior part of the anterior iliac crest.
INSERTION.—The tendon forms along the posterodistal edge of the belly
and continues beyond the end of the belly as a cylindrical tendon that passes
through the tendinous guide loop (the belly terminates approximately at the
level of the guide loop), then extends anterodistally into the shank muscula-
ture; the tendon passes between the medial and lateral heads of M. flexor
perforatus digiti IV, between the medial and lateral heads of M. flexor
perforatus digiti II, lateral to the common tendon of the anterolateral heads
of Mm. flexor perforatus digiti IV, flexor perforatus digiti II, and flexor
perforatus digiti III, and between the posterior and lateral heads of M.
flexor digitorum longus, attaching to the fibular tubercle.
The tendinous guide loop has three arms—proximal femoral, distal femoral,
and fibular; the proximal and distal femoral arms unite posterior to the
tendon of M. extensor iliofibularis; the proximal arm is medial to, and the
distal arm is lateral to, the latter; the fibular arm joins the distal edge of the
distal arm lateral to the tendon of M. extensor iliofibularis. The proximal arm
extends anteroproximally lateral to the medial head of M. flexor perforatus
digiti IV and medial to M. vastus lateralis pars postica, attaching to a
narrow line on the anterolateral surface of the femur a short distance proximal
to the external condyle and adjacent (posterior) to the origin of M. vastus
lateralis pars postica. The distal arm extends anteriorly medial to the pos-
terior head of M. flexor perforans et perforatus digiti IT and medial to M.
vastus lateralis pars postica, attaching in common with the tendon of origin of
M. gastrocnemius pars externa to a small oval area on the posterolateral surface
4—5835
412 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
of the femur a short distance proximal to the fibular groove; the arm is also
fused to the underlying articular capsule. The fibular arm (broadest of the
three) passes deep to, and fused with, the common tendon of origin of the
lateral heads of Mm. flexor perforatus digiti IV and flexor perforatus digiti II,
superficial to the common tendon of origin of the anterolateral heads of Mm.
flexor perforatus digiti IV, flexor perforatus digiti II, and flexor perforatus
digiti III, and deep to the tendon of M. ambiens, attaching broadly to a
narrow line on the anterolateral surface of the proximal part of the fibula;
the arm is also fused to the underlying articular capsule.
INNERVATION.—A branch of the middle peroneal division of the sciatic
nerve sends twigs to the deep surface of the anteroproximal part; the dorsal
peroneal division of the sciatic nerve penetrates. the deep surface of the
proximal end.
INDIVIDUAL VARIATION.—In some instances a variable number of twigs
arises from the peroneal nerve near the middle of the thigh and enters the
deep surface of the muscle. They are difficult to expose without breaking
and may have been overlooked in some specimens.
T. cupido
INDIVIDUAL VARIATION.—The same variation is found as in T. pallidicinctus
(see above). In one leg, the tendon of insertion bifurcates into proximal and
distal arms before attaching.
P. p. jamesi
DIFFERENCES FROM TypicaL T. pallidicinctus.—It arises from the ventral
rather than the ventromedial surface of the lateral iliac process (there is no
ventromedial surface to this process ).
INnpIvipuAL VARIATION.—In nearly all of the legs, minute twigs to M. ex-
tensor iliofibularis come off the peroneal nerve near the middle of the thigh.
The insertional tendon tends toward doubleness in two legs.
M. Piriformis (M. gluteus medius et minimus), Figs. 16, 20H, I
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Small, thin, and triangular; lateral
to antitrochanter and posterior part of trochanter; deep to M. extensor ilio-
fibularis and posterior (adjacent) to M. gluteus profundus; distal half (or
more) tendinous.
Ornicin.—The muscle arises fleshily from the posterior end of the anterior
iliac crest (ventral to the origins of Mm. extensor iliotibialis lateralis and ex-
tensor iliofibularis) beginning adjacent to the posterior end of M. gluteus
profundus.
INSERTION.—The flat tendon narrows, overlaps the anteroproximal corner of
insertion of M. flexor ischiofemoralis, and attaches to the lateral surface of
the proximal part of the femur immediately anterior to the insertion of M.
flexor ischiofemoralis and posterior to the proximal end of M. vastus lateralis;
the attachment is posterodistal to the insertion of M. iliotrochantericus medius
and posteroproximal to the insertion of M. iliacus.
MusSCcLES AND NERVES OF LEG OF GROUSE 413
INNERVATION.—The small anterior peroneal division of the sciatic nerve
turns anteriorly immediately after emerging from the ilio-ischiatic fenestra
and passvs deep to M. piriformis, giving twigs to the deep surface.
InpIvipuAL VARIATION.—In both legs of one specimen, the insertion does
not overlap the insertion of M. flexor ischiofemoralis. The posteroproximal
corner of the muscle is tendinous in one leg.
T. cupido
INDIVIDUAL VARIATION.—The anterior border is somewhat fused with the
posterior edge of M. gluteus profundus in one leg, while in another there is a
slight gap between the origins of M. gluteus profundus and M. piriformis. In
one leg, the posterior edge of the origin is aponeurotic. On both sides of one
specimen, an accessory tendinous band arises several mm. posterior to the main
part of M. piriformis and joins the proximal part of the insertional tendon, thus
forming a Y-shaped unit (fig. 201); the accessory tendon arises from the
anterior end of the lateral iliac process (left side) or from the anterior part of
the lateral iliac fossa (right side). The insertion may be proximal (rather than
posterior) to the proximal end of M. vastus lateralis. In one leg, the in-
sertional tendon is partly fused to the insertional tendon of M. flexor ischio-
femoralis.
P. p. jamesi
INDIVIDUAL VARIATION.—There is often a gap between the origins of M.
gluteus profundus and M. piriformis. In one leg (fig. 20H), the postero-
proximal corner of the muscle is aponeurotic. The insertion is often proximal
(rather than posterior) to the proximal end of M. vastus lateralis. In one in-
stance, the insertion does not overlap the insertion of M. flexor ischiofemoralis.
M. Gluteus Profundus (M. iliotrochantericus posterior), Figs. 14, 16
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Large and thick; covering dorso-
lateral surface of entire preacetabular part of ilium; deep to Mm. extensor
iliotibialis lateralis and extensor iliotibialis anticus; bounded posteriorly by M.
piriformis and ventrally by M. iliacus; ventral edge fused with anterior part of
latter and with proximal accessory aponeurosis of M. extensor iliotibialis
lateralis; tough sheet of fascia strongly fused to anterior two thirds of lateral
surface; posterior to this, fascia overlying muscle but not attaching to it;
posterior half of fascia fused to overlying aponeurosis of M. extensor iliotibialis
lateralis; deep surface of muscle somewhat fused to proximal part of M.
iliotrochantericus medius.
Oricin.—The superficial surface is tendinous from the entire anterior iliac
crest except the posterior end and from the crest forming the anterior and
anterolateral edges of the ilium; the muscle arises fleshily from the entire
dorsolateral surface of the preacetabular ilium as far posteriorly as the level of
the pectineal process; the dorsal edge is adjacent (anterior) to the origin of
M. piriformis.
INSERTION.—The attachment is by a short, wide, thick tendon to a curved
line (convex anteriorly) on the lateral surface of the femoral trochanter.
INNERVATION.—The anterodorsal division of the femoral nerve turns dorsally
414 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist.
through the femoral notch of the ilium and penetrates the deep surface of the
ventral part of the muscle midway of its length; the anterior peroneal division
of the sciatic nerve passes deep to M. piriformis and terminates near the
posterodorsal edge of M. gluteus profundus.
INDIVIMUAL VARIATION.—On both sides of one specimen, the branch from
the femoral nerve passes lateral to the extreme anteroproximal corner of M.
iliotrochantericus medius instead of through the femoral notch.
T. cupido
INDIVIDUAL VARIATION.—In one leg, the insertional tendon is strongly fused
to the insertional tendon of M. iliotrochantericus medius.
P. p. jamesi
INDIVIDUAL VARIATION.—None of significance.
M. Lliacus (M. iliotrochantericus anterior), Figs. 13, 14, 15, 16, 17
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Adjacent ventrally to ventrolateral
edge of M. gluteus profundus; lateral edge much thicker than medial edge;
deep to M. extensor iliotibialis lateralis and anterolateral to M. iliotrochantericus
medius; distal (posterior) end passing between proximal ends of Mm. vastus
medialis and vastus lateralis pars lateralis; insertion overlapped by latter;
dorsal surface of anterior part fused with ventrolateral edge of M. gluteus
profundus and with ventral surface of proximal accessory aponeurosis of M.
extensor iliotibialis lateralis; ventral surface partly fused with distal accessory
aponeurosis of latter muscle.
Oricin.—The origin is fleshy and tendinous from the lateral edge of the
anterior part of the ilium.
INSERTION.—The attachment is by a short flat tendon to the lateral surface
of the femur distal to the trochanter and anterodistal to the insertion of M.
piriformis and deep to the proximal part of M. vastus lateralis pars lateralis.
INNERVATION.—The dorsal division of the femoral nerve penetrates the
ventral surface.
InpIvipuAL VARIATION.—The dorsal division of the femoral nerve may fuse
proximally with either the anterior or middle division. In one leg, there are
two separate branches to the muscle.
T. cupido
INDIVIDUAL VARIATION.—The insertion may not be overlapped by M. vastus
lateralis. The dorsal division of the femoral nerve is fused proximally with
the middle division in one leg.
P. p. jamesi
DIFFERENCES FROM TypicaL T. pallidicinctus——The fleshy origin is wider.
InptvipuaL VARIATION.—The dorsal division of the femoral nerve may fuse
proximally with either the anterior or middle division. In one leg, there are
two branches to M. iliacus, one fused with the anterior division and the other
with the middle division.
MuscLes AND NERVES oF LEG OF GROUSE 415
M. lliotrochantericus Medius, Figs. 17, 20A, B, C, D
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Small and triangular; ventral to
posterior half of M. gluteus profundus; all but posteroventral corner deep to
latter; posteromedial to M. iliacus, anterior to neck of femur, and dorsolateral
(adjacent proximally) to M. psoas; proximal end notched at level of femoral
notch for passage of anterodorsal division of femoral nerve; part anterior to
femoral notch mainly tendinous; dorsal surface of proximal part somewhat
fused to M. gluteus profundus, proximal accessory aponeurosis of M. extensor
iliotibialis lateralis split into two sheets enclosing and fusing with M. iliotro-
chantericus medius, ultimately attaching to lateral edge of ilium in common
with origin of latter muscle.
Oricin.—The muscle arises from the ventrolateral surface of the ilium
anterior to the acetabulum and posterior to the origin of M. iliacus; the
anterior part attaches to the ventrolateral edge of the ilium and the posterior
part attaches just above the ventral edge. The muscle is not attached to the
concavity of the femoral notch (the origin is notched here). The part at-
taching anterior to the femoral notch is narrow, tendinous, and continous
anteriorly with the accessory aponeurosis of M. extensor iliotibialis lateralis
(thus the anterior border of the muscle cannot be exactly delimited), The
part attaching posterior to the femoral notch is wider and fleshy (fig. 20A).
INSERTION.—The short fiat tendon attaches to the lateral surface of the
distal end of the trochanter slightly anterior and immediately distal to the
insertion of M. gluteus profundus; the attachment is proximal to the origin
of M. vastus lateralis, anteroproximal to the insertion of M. piriformis, and
several mm. proximal to the insertion of M. iliacus.
INNERVATION.—The small posterodorsal division of the femoral nerve
penetrates the ventral surface.
INDIviUAL VARIATION.—On both sides of one specimen, the femoral notch
is absent and the proximal end of the muscle is not notched; the proximal
part is entirely fleshy and the anterior border is well defined (fig. 20D).
T. cupido
INDIVIDUAL VARIATION.—The part attaching anterior to the femoral notch
has a fleshy origin in one leg (fig. 20B), but in another, no part attaches
anterior to the femoral notch (thus the muscle is not notched). In one leg,
the insertional tendon is strongly fused to, and continuous with, the ventral
edge of the insertional tendon of M. gluteus profundus,
P. p. jamesi
INDIvIUAL VARIATION.—The part attaching anterior to the femoral notch
may be mainly or entirely fleshy. In one leg, the part attaching anterior to
the femoral notch is entirely separate from, although overlapped by, the main
part of the muscle for the entire length of the fleshy belly (fig. 20C); both
parts have a common insertional tendon.
416 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
M. Psoas (M. iliacus), Figs. 138, 15, 18
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Small and slender; on medial
aspect of proximal end of thigh lateral to proximal end of M. ambiens; ventro-
medial to M. iliotrochantericus medius; proximal end visible from inside pelvis
(medial to inguinal ligament); passes dorsolateral to inguinal ligament.
Oricin.—The muscle arises fleshily from the ventrolateral edge of the ilium
posterior to the femoral notch and ventral (ona) to the origin of M.
iliotrochantericus medius,
INSERTION.—The attachment is tendinous to the medial surface of the femur
a short distance proximal to the origin of M. femoritibialis internus.
INNERVATION.—The posterior division of the femoral nerve, which spirals
completely around M. psoas, gives several twigs into the proximal part.
InprvmuAL VaARIATION.—None of significance.
T. cupido
INDIVIDUAL VARIATION.—In two legs the insertion is partly fleshy.
P. p. jamesi
InprvipuAL VarraTion.—In one leg the insertion is partly fleshy, The
posterior division of the femoral nerve perforates the muscle in one instance.
M. Flexor Cruris Lateralis (M. semitendinosus), Figs. 12, 13, 14, 15, 16, 17
This muscle represents only the main head of the muscle for which Fisher
and Goodman (1955) used the same name. Their accessory head of M.
flexor cruris lateralis is here termed M. femorocruralis.
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Large, thick, and strap-shaped;
on posterior surface of thigh; proximal part bounded anteriorly by Mm. ex-
tensor iliotibialis lateralis and extensor iliofibularis; anterodistal part deep to
latter; bounded medially by Mm. caudofemoralis (proximally) and flexor
cruris medialis (distally); proximal end much narrower than remainder and
posterior to ilium; fused to underlying tough membrane, which forms body
wall posterior to ilium; proximal half of narrow part aponeurotic; distal part
of muscle posterior to M. femorocruralis; separated from latter by common
raphe to which both attach; caudal muscle (M. transversoanalis) attached
aponeurotically to superficial surface of posteroproximal fleshy part of M.
flexor cruris lateralis.
Oricin.—The origin is tendinous (superficial surface) and fleshy from the
entire dorsolateral iliac ridge and fleshy from an area of the ilium below this
ridge, also tendinous from the posterior edge of the ilium medial to the
dorsolateral iliac ridge, and also tendinous from the transverse processes of
the first free caudal vertebra and the vertebra either anterior or posterior to
the latter.
INSERTION.—M. flexor cruris lateralis and M. femorocruralis insert broadly
on opposite sides of a long tendinous raphe that extends parallel to, but some
distance posterior to, the distal half of the femur; the distal end of this tendon
Muscies AND NERVES OF LEG OF GROUSE ANT.
broadens somewhat and fuses to the medial surface of M. gastrocnemius pars
media (continuous with the tendon of the latter); the superficial part of this
tendon continues toward the tibiotarsus, soon fusing to the deep surface of the
overlying tendon of M. flexor cruris medialis; thus the common tendon of M.
flexor cruris lateralis and M. femorocruralis insert in common with both M.
flexor cruris medialis and M. gastrocnemius pars media.
INNERVATION.—A branch of the middle tibial division of the sciatic nerve
enters the substance of M. caudofemoralis pars iliofemoralis, and emerges near
its ventral edge, then passes lateral to M. caudofemoralis pars caudifemoralis
and enters the anterior part of M. flexor cruris lateralis.
InprvinuaAL VartaTion.—In three legs, the nerve does not perforate M.
caudofemoralis pars iliofemoralis, but passes deep to it.
T. cupido
InprvipuaL VARIATION.—In one leg, a small accessory slip arises from the
ventrolateral surface of the caudal musculature and joins the posterior edge of
the main part of M. flexor cruris lateralis a short distance dorsal to the pubis.
In several legs, the nerve does not perforate M. caudofemoralis pars iliofemo-
ralis, but passes deep to it.
P. p. jamesi
DIFFERENCES FROM TypicaL T. pallidicinctus—The muscle is wider. The
extreme proximal end is fleshy up to its origin, which is fleshy and tendinous
from the vertebrae. The common insertional tendon of M. flexor cruris lateralis
and M. femorocruralis fuses with the distal end of the fleshy part (instead of
tendon) of M. flexor cruris medialis.
INDIVIDUAL VARIATION.—None of significance.
be Flexor Cruris Medialis (M. semimembranosus), Figs. 12, 13, 14, 15, 16,
17208
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Most posterior muscle on medial
surface of thigh; long and strap-shaped; bounded anteriorly by M. adductor
profundus; posteroproximal corner of latter medial to anteroproximal part of
M. flexor cruris medialis; bounded laterally by Mm. caudofemoralis (proxi-
mally) and flexor cruris lateralis (distally); anteroproximal corner adjacent to
posteroventral corner of M. flexor ischiofemoralis and lateral to extreme postero-
proximal corner of M. adductor superficialis; distal end tendinous, extending
into proximal part of shank; bounded medially by M. gastrocnemius pars
interna and laterally by Mm. gastrocnemius pars media and plantaris.
Oricin.—The muscle arises by a wide flat tendon from a narrow line on
the lateral surface of the ischium dorsal to the ventral ischiatic tubercle.
INsERTION.—The wide flat tendon attaches to a narrow line on the medial
surface of the proximal part of the tibiotarsus a short distance anterior to the
proximal part of M. plantaris and deep to M. gastrocnemius pars interna; the
proximal end attaches immediately anterior to the distal end of the medial col-
lateral ligament. Part of the common tendon of Mm. flexor cruris lateralis and
femorocruralis fuses with the lateral surface of the tendon of M. flexor cruris
medialis, inserting in common with it.
418 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
INNERVATION.—A branch of the middle tibial division of the sciatic nerve
passes deep to both heads of M. caudofemoralis and enters the anterior part
of M. flexor cruris medialis.
INDIVIDUAL _VARIATION.—In several legs, the anterior edge of the proximal
part fits into a deep longitudinal groove in the posterior edge of the proximal
part of M. adductor superficialis; the two muscles fuse slightly at this point.
T. cupido
INDIVIDUAL VARIATION.—In two legs, the extreme posterior end of the
origin is from the pubis. In two others, the proximal end is separated by a
slight gap from M. adductor superficialis. The nerve arises from the posterior
(rather than middle) tibial division in one leg.
P. p. jamesi
DIFFERENCES FROM TypicaL T. pallidicinctus——The origin is wider; the
posterior third to half of the origin is fleshy. The entire origin is from a
strongly curved line, the middle part of which attaches to the ventral edge of
the ischium posterior to the ventral ischiatic tubercle. The insertion is wider.
The insertional tendon attaches posterior (rather than anterior) to the distal
end of the medial collateral ligament; the proximal end of the insertion
attaches to the articular capsule (fig. 20E). The insertional tendon is shorter;
as a result, the common tendon of Mm. flexor cruris lateralis and femorocruralis
fuses with the distal end of the fleshy belly (instead of the tendon) of M.
flexor cruris medialis,
INDIvipUAL VARIATION.—In two thirds of the legs, the proximal part of the
insertion is fleshy rather than tendinous. In one leg, the middle part of the
insertional tendon spilts into two sheets, one attaching anterior to and one
attaching posterior to the distal end of the medial collateral ligament. The
nerve may arise from the posterior tibial division (two legs), from the middle
tibial division (one leg), or as an independent division of the tibial nerve
(three legs). In one leg, the nerve perforates the lateral part of M. flexor
ischiofemoralis.
M. Caudofemoralis (M. piriformis), Figs. 12, 13, 14, 15, 16, 20J
T. pallidicinctus
GENERAL DESCRIPTION AND ReELATIONS.—Posterior to proximal part of
shaft of femur and deep to M. extensor iliofibularis; posterior part deep to
M. flexor cruris lateralis; bounded medially by Mm. flexor ischiofemoralis
(dorsally), flexor cruris medialis (posteriorly), and adductor superficialis
(anteroventrally); anterior end distal to anterior end of M. flexor ischio-
femoralis; two distinct heads—pars iliofemoralis and pars caudifemoralis; pars
iliofemoralis dorsal to pars caudifemoralis; posteroventral corner of former
overlapped by latter; pars iliofemoralis wider and much shorter than pars
caudifemoralis; extreme posterior end of pars iliofemoralis fused to overlying
posteroproximal aponeurosis of M. extensor iliotibialis lateralis; small part of
ventral edge sometimes fused with underlying tendinous posteroproximal
corner of M. flexor cruris medialis; entirely fleshy except for smal] triangular
tendinous area along dorsal margin at point where branch of middle tibial
division of sciatic nerve passes deep to muscle; pars caudifemoralis long, thin,
MuscLes AND NERVES OF LEG OF GROUSE 419
narrow, and strap-shaped; overlapping posteroventral corner of ischium; pos-
terior end of fleshy belly narrowed and forming long slender tendon passing
into caudal musculature; anterior end forming short narrow tendon fused to
deep surface of ventral edge of pars iliofemoralis relatively near insertion;
tendon continuous to insertion; fleshy anterodorsal corner of pars caudifemoralis
slightly overlapped by ventral edge of pars iliofemoralis; some form of con-
nection usually present between anterior part of M. caudofemoralis pars caudi-
femoralis and dorsal end of raphe between Mm. flexor cruris lateralis and
femorocruralis, most often consisting of narrow weak tendon.
Oricin.—Pars iliofemoralis: This arises fleshily from the ventromedial sur-
face of the posterior part of the lateral iliac process, from the entire lateral
ischiatic ridge, and from the lateral surface of the ischium anterior to this
ridge nearly as far forward as the posterior edge of origin of M. flexor ischio-
femoralis; the posteroventral corner reaches the ventral edge of the ischium
and usually attaches to the ischiopubic membrane posterior to M. flexor
cruris medialis. Pars caudifemoralis: This arises by a narrow tendon from
the ventral surface of a broad, thick, tendinous sheet ventral to the pygostyle,
which, in tum, attaches to the ventral surface of the pygostyle.
INSERTION.—The common belly formed by the union of the two heads
narrows (width variable) and attaches to the posterolateral surface of the
femur distal to the level of insertion of M. iliacus and in contact with the
posterior edge of origin of M. vastus lateralis pars lateralis; the dorsal part
is fleshy and the ventral part is tendinous.
INNERVATION.—A branch of the middle tibial division of the sciatic nerve
gives several twigs to the deep surface of pars iliofemoralis; another twig
enters the substance of pars iliofemoralis and emerges from the ventral edge
of the latter, then enters the dorsal edge of pars caudifemoralis, The latter
twig was not found in all legs, but was probably destroyed during dissection.
INDIVIDUAL VARIATION.—The tendinous area in the dorsal margin of pars
iliofemoralis is lacking in one leg and extremely small in some others. In
both legs of one specimen, the connection between M. caudofemoralis pars
caudifemoralis and the raphe between Mm. flexor cruris lateralis and femoro-
cruralis consists of a small (11 x 2 mm.) but well developed and entirely
fleshy muscle slip (fig. 16). In one leg, the ventral third of this connection is
fleshy, the remainder tendinous; in another, this connection is completely
lacking.
T. cupido
INDIvIbUAL VARIATION.—The tendinous area in the dorsal margin of pars
iliofemoralis is lacking in one leg. The connection between pars caudifemoralis
and the raphe between Mm. flexor cruris lateralis and femorocruralis is lacking
in several legs. A conspicuous variation occurring in three legs is the presence
of a tendinous area in the belly of pars caudifemoralis, dividing the latter
into proximal and distal parts (fig. 20J). In one leg, the posteroventral commer
of pars iliofemoralis arises from the pubis. The origin of pars caudifemoralis
in three legs is directly from the anteroventral surface of the pygostyle. In
one instance, the insertional tendon of pars caudifemoralis is long and ex-
tremely slender and extends for some distance in a groove on the medial
surface of pars iliofemoralis before fusing with the latter.
420 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist.
P. p. jamesi
DIFFERENCES FROM TypicaL T. pallidicinctus—There is no connection
at all between pars caudifemoralis and the raphe between Mm. flexor cruris
lateralis and femorocruralis. The posteroventral corner of pars iliofemoralis
is some distance dorsal to the ventral edge of the ischium and, therefore, does
not attach to the ischiopubic membrane.
INDIVIDUAL VARIATION.—The insertion (narrow) is entirely tendinous in
one leg.
M. Flexor Ischiofemoralis (M. ischiofemoralis ), Figs. 16, 17
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Thick; on lateral surface of anterior
part of ischium; posterior end in lateral iliac fossa; deep to Mm. extensor
iliofibularis and caudofemoralis pars iliofemoralis; overlapping ventral extra-
pelvic part of M. obturator and anteroproximal part of M. adductor super-
ficialis (slightly fused to proximal edge of latter); posteroventral corner con-
tacting anteroproximal corner of M. flexor cruris medialis; extreme anterodorsal
corner usually overlapped by tendon of M. piriformis.
Oricin.—The muscle arises fleshily from a large area on the lateral surface
of the ischium extending ventrally to the origin of M. adductor superficialis,
anteriorly to the level of the posterior end of the obturator foramen, dorsally
to the ventral border of the ilio-ischiatic fenestra and to the depth of the
lateral iliac fossa, and posteriorly approximately to the level of the ventral
ischiatic tubercle.
INSERTION.—The short flat tendon attaches to the lateral surface of the
femur immediately posterior to the insertion of M. piriformis.
INNERVATION.—The posterior tibial division of the sciatic nerve penetrates
the dorsal surface.
INDIVIDUAL VARIATION.—The ventral part of the insertion may be fleshy.
T. cupido
INDIVIDUAL VARIATION.—None of significance.
P. p. jamesi
INDIVIDUAL VARIATION.—In all the legs except one, an additional twig arises
from the branch to M. flexor cruris medialis and penetrates the lateral surface
of M. flexor ischiofemoralis. The ventral part of the insertion is fleshy in one
leg.
M. Adductor Superficialis (M. adductor longus et brevis, pars externa), Figs.
1A 6 7,
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Posterior to femur, lateral to M.
adductor profundus, and medial to Mm. flexor ischiofemoralis, caudofemoralis,
and femorocruralis; proximal end (fleshy) fused to proximal tendinous end of
M. adductor profundus.
Oricin.—The origin is fleshy and tendinous from the proximal end of the
lateral surface of M. adductor profundus and from a narrow line on the ischium
MusSCLES AND NERVES OF LEG OF GROUSE 421
adjacent (dorsal) to the origin of the latter; the posterior part of the origin
sometimes extends farther dorsally on the lateral surface of the ischium; the
origin does not extend so far anteriorly nor so far posteriorly as the origin of
M. adductor profundus; the anterior edge is at the posterior border of the
obturator foramen.
INsERTION.—The attachment is fleshy and thick (distal end thin) to the
posterior surface of the middle part of the femur between the posterior and
posterolateral intermuscular lines; the attachment is adjacent (lateral) to the
insertion of M. adductor profundus and adjacent (medial) to the origins of
Mm. vastus lateralis (proximally) and femorocruralis (distally); the proximal
edge is approximately at the level of the distal edge of the insertion of M.
caudofemoralis.
INNERVATION.—A branch of the obturator nerve emerges from the obturator
foramen dorsal to the tendon of insertion of M. obturator pars postica, turns
ventrally (crossing latter), and passes deep to the anteroproximal corner of M.
adductor superficialis, extending posterodistally between the adductor muscles
and giving twigs to the medial surface of M. adductor superficialis and to the
lateral surface of M. adductor profundus.
INDIVIDUAL VARIATION.—The anterior edges of the two adductor muscles
are so firmly fused together in some cases that the boundaries cannot be
identified at this point. In several legs, there is a deep longitudinal groove in
the posterior edge of the proximal part of the muscle into which the anterior
edge of M. flexor cruris medialis fits.
T. cupido
INDIVIDUAL VARIATION.—In some cases, the anterior edges of the two ad-
ductor muscles are firmly fused together.
P. p. jamesi
DIFFERENCES FROM TypicaL T. pallidicinctus—The origin is narrower.
INDIvipuUAL VARIATION.—The anterior edges of the two adductor muscles
may be fused together. In one leg, the entire muscle is indistinguishably
fused with M. adductor profundus and they appear as a single muscle.
M. Adductor Profundus (M. adductor longus et brevis, pars interna), Figs.
SSS Leas
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Broad; on medial surface of thigh
immediately posterior to femur; bounded posteriorly by M. flexor cruris
medialis (medial to anteroproximal corner of latter), anteriorly by M. femori-
tibialis internus (anterior edge overlapped by latter), and laterally by Mm.
adductor superficialis and femorocruralis; proximal end tendinous (except an-
terior edge), fused to proximal fleshy end of M. adductor superficialis.
Oricin.—The muscle arises tendinously from the ventral edge of the
ischium extending from the posterior border of the obturator foramen to the
ventral ischiatic tubercle and (anterior edge) fleshily from the lateral surface
of the pubis ventral to the obturator foramen; the origin is adjacent (ventral)
to the origin of M. adductor superficialis.
4292, UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
INSERTION.—The attachment is fleshy and tendinous from the posterior
intermuscular line and (proximally and distally) from a narrow adjacent area.
Proximally there are often two approximately parallel lines a short distance
apart, representing points of attachment of the lateral and medial edges of the
muscle; if there is only one line proximally, it may represent the attachment
of either the lateral or medial edge of the muscle; distally there is usually
only one line, representing the lateral edge of the muscle. The distal end
extends onto the posterior surface of the proximal part of the internal condyle,
and is adjacent (lateral) to the origin of M. femoritibialis internus, adjacent
(medial) to Mm, adductor superficialis and femorocruralis, and adjacent
(proximal) to M. gastrocnemius pars media.
INNERVATION.—See M. adductor superficialis.
INDIVIDUAL VARIATION.—The anterior edges of the two adductor muscles
are strongly fused together in some cases.
T. cupido
INDIVIDUAL VARIATION.—The anterior edge may be fused with that of M.
adductor superficialis. The distal end is sometimes slightly fused with M.
gastrocnemius pars media. In one leg, the proximal two thirds of the insertion
is entirely tendinous, whereas in another the distal end of the insertion is
tendinous.
P. p. jamesi
INDIVIDUAL VARIATION.—The anterior edge (in one leg the entire muscle)
in some legs fuses with that of M. adductor superficialis.
M. Obturator (M. obturator externus + M. obturator internus), Figs. 16, 17,
18; 19C, D; 20K, LM
I am adopting the single name M. obturator for the complex that Fisher
(Fisher, 1946; Fisher and Goodman, 1955) subdivides into Mm, obturator
externus and obturator internus. The reasons for this change are given in
the section on terminology.
For ease of description, it is desirable to apply names to the subdivisions
of M. obturator. It has been customary to divide the obturator complex into
two parts—an obturator internus and an obturator externus; the latter has
often been further subdivided. The evidence given below demonstrates that
a primary division of the complex into only two parts is unsatisfactory.
I strongly suspect that comparable parts of the obturator complex have
been considered a part of the “intemus” in some birds and a part of the
“externus” in others. In their work on the Galliformes, Hudson, et al. (1959)
subdivide the obturator complex into only two divisions—obturator externus
and obturator internus. The extrapelvic part of this complex that arises from
the rim of the obturator foramen and inserts in common with the stout tendon
of the main intrapelvic part of the obturator internus is considered by them
to be a part of the obturator internus, Their obturator externus lies anterior
and deep to the extrapelvic part of the obturator internus and inserts separately
from the latter. (I also have found this same arrangement in Tympanuchus
and Pedioecetes. )
Berger (1952), in his description of the Black-billed Cuckoo (Coccyzus
erythrophthalmus), also divides the obturator complex into an obturator in-
Muscles AND NERVES OF LEG OF GROUSE 423
ternus and an obturator externus; the latter he subdivides into a dorsal and
a ventral part. He states (p. 530) that he did not find any measurable dif-
ferences in myology between C. erythrophthalmus and C. americanus. In
order better to compare this arrangement with that in Tympanuchus, I have
examined two specimens of C, americanus. My findings in the latter differ
from Berger’s description (p. 541) in one respect. Whereas Berger states
that the dorsal and ventral parts of M. obturator externus are distinct except
at their origin, I find them fused for their entire length; the muscle fibers that
connect these two parts lie deep to the tendon of M. obturator intemus. The
origin of all parts of the complex in Coccyzus is similar to that in Tym-
panuchus. The only notable difference in configuration is that the part in
Coccyzus that appears to correspond to the obturator externus of Hudson,
et al. (1959) is not separate from the remainder of the extrapelvic part of
the muscle. Berger (1952) considers all parts of the muscle having an extra-
pelvic origin to make up the obturator externus. It appears to me that the
dorsal part and a part of the ventral part of the obturator externus of Berger
correspond to the extrapelvic fleshy part of the obturator internus of Hudson,
et all,
From my limited study, it seems to me to be desirable to recognize four sub-
divisions of the obturator complex, for which I propose the terms pars antica,
pars dorsalis, pars ventralis, and pars postica. These parts exhibit various de-
grees of fusion in different groups of birds and some parts appear to be absent
in certain birds. A. study of a wide variety of birds will be required to de-
termine whether or not a subdivision into the four parts proposed here is
suitable for birds as a whole.
Applying these terms to Coccyzus, pars postica is equivalent to the entire
obturator internus of Berger (1952). Pars dorsalis is apparently equivalent to
the dorsal part of Berger’s obturator externus. The ventral part of the ob-
turator externus of Berger represents the fused pars antica and pars ventralis.
The main parts of the obturator muscle appear to be pars postica and pars
antica. Pars dorsalis and pars ventralis are more variable; in Coccyzus these
two parts are closely associated with pars antica whereas in Tympanuchus
they are most closely associated with pars postica. Apparently pars dorsalis
and pars ventralis may be absent in some birds.
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Deeply situated immediately
posterior to head of femur; part extending through obturator foramen and
lying inside pelvis; extrapelvic part deep to Mm. flexor ischiofemoralis and
piriformis; muscle partially divisible into four parts—pars antica, pars dorsalis,
pars ventralis, and pars postica (fig. 20K); pars postica: mostly inside pelvis;
much larger than other parts; broad (narrow anteriorly); on medial surface
of ischium; composed of several fascicles; anterior end forming narrow, heavy
tendon (with some fleshy fibers on posterior part of deep surface) passing
through obturator foramen; anteriormost fleshy fibers of ventralmost fascicle
fused with pars ventralis; pars ventralis: essentially extrapelvic (see origin);
mostly ventral to tendon of pars postica; superficial to pars antica; fused to
anterior fleshy part of pars postica; anterodorsal edge usually adjacent to, and
often slighily fused with, ventral edge of pars dorsalis (deep to tendon of pars
postica); pars dorsalis: entirely extrapelvic; mostly dorsal to tendon of pars
424 UNIVERSITY OF KANnsAS Pusts., Mus. Nar. Hist.
postica; superficial to dorsal part of pars antica; pars antica: extremely short
but relatively thick; entirely fleshy; entirely extrapelvic; between obturator
foramen and head of femur; anterior surface adjacent to articular capsule;
almost completely covered by other parts of muscle; proximal end of posterior
surface often slightly fused with adjacent parts of pars ventralis and pars
dorsalis.
Oricin.—Pars postica: This arises fleshily from the medial surface of the
entire ischium except the posterior end, from the dorsomedial and medial sur-
faces of the anterior half of the pubis as far forward as the obturator foramen,
from the internal ilio-ischiatic crest, from the medial surface of the ilium for
a short distance posterior to this crest, and from the iliac recess; the postero-
ventral corner usually arises from the medial surface of the ischiopubic mem-
brane. Pars ventralis: This arises fleshily from the dorsomedial edge of the
ventral border of the obturator foramen (fig. 20M) and (narrowly) from the
anterior border of the foramen; this part may or may not arise from the
lateral surface of the anteroventral border of the foramen and is usually
adjacent along the anterior border of the foramen to pars dorsalis; pars
ventralis is continuous along the ventral border of the foramen with the in-
trapelvic origin of pars postica. Pars dorsalis: This arises fleshily from the
lateral surface of the anterodorsal border of the foramen (fig. 20L) and may
extend posteriorly along the dorsal border of the foramen. Pars antica: This
arises fleshily from the depresssed area anterior to the obturator foramen (ad-
jacent to pars dorsalis and pars ventralis); the posteroventral corner may arise
from the lateral surface of the anteroventral border of the obturator foramen
(ventral to the anterior end of pars ventralis; fig. 20L).
INseRTION.—Pars postica: Several tendinous bands (intrapelvic) converge
and coalesce, forming a single strong tendon that passes through the obturator
foramen and attaches to the lateral surface of the femoral trochanter a short
distance posterior to the insertion of M. gluteus profundus and proximal to the
insertion of M. flexor ischiofemoralis. Pars ventralis: The attachment is fleshy
and tendinous to the ventral edge and the deep surface of the tendon of pars
postica. Pars dorsalis: The attachment is fieshy and tendinous to the dorsal
edge of the tendon of pars postica. Pars antica: The attachment is fleshy to
the posterior surface of the proximal end of the femur several mm. posterior to
the insertion of pars postica; the lateral edge attaches to the obturator ridge.
INNERVATION.—The muscle is supplied by the obturator nerve; several twigs,
which do not pass through the obturator foramen, penetrate the anterior
part of the medial surface of pars postica; several twigs pass through the
obturator foramen and supply pars dorsalis, pars ventralis, and pars antica.
INDiIvipuvAL VaRIATION.—In some cases the origin of pars postica does not
include the dorsal end of the internal ilio-ischiatic crest nor the ilium posterior
to it. Tiny but distinct accessory slips are sometimes present. In one leg
a tendinous slip of parts antica extends beyond the remainder of the muscle
and inserts independently on the trochanter close to the insertion of pars
postica. In another leg, a fleshy and tendinous slip of pars antica attaches
to the deep surface of the insertional tendon of pars postica. In still another
leg, a fleshy and tendinous slip of pars dorsalis inserts adjacent (anterior)
to the dorsal edge of the insertion of pars antica.
MuscLes AND NERVES OF LEG OF GROUSE 495
T. cupido
InpivipvaL VARIATION.—The variations are similar to those given above
for T. pallidicinctus except that there is no slip of pars antica attaching to
the tendon of pars postica.
P. p jamesi.
InpDIvipuAL VARIATION.—There are variations similar to those given above
for T. pallidicinctus except that there is no independent slip of pars antica
attaching on the trochanter close to the insertion of pars postica. Pars dorsalis
may be quite small. In several legs, pars dorsalis is more closely associated
with pars antica than with pars postica; in one of these, pars dorsalis is in-
distinguishably fused with pars antica (inserting with the latter) except for
a few fibers which insert with pars postica.
M. Femorocvuralis (M. accessorius semitendinosi), Figs. 14, 15, 16, 17
Fisher (Fisher, 1946; Fisher and Goodman, 1955) considers this muscle
as an accessory head of M. flexor cruris lateralis. The reasons for this change
in terminology are given in the section on terminology.
T. pallidicinctus
GENERAL DEscRIPTION AND RELATIONS.—Short and broad; posterior to
distal part of femur; deep to Mm. extensor iliofibularis and vastus lateralis
pars postica; bounded posteriorly by M. flexor cruris lateralis, medially by
Mm. adductor superficialis and adductor profundus, and distally by M. gastro-
cnemius pars media; fused to a variable degree with the latter (in some cases
these two muscles fused firmly together, appearing as single muscle); distal
and medial to proximal end of M. flexor perforatus digiti IV.
Oricin.—The muscle arises fleshily (thin proximally, thick distally) from
the posterior surface of approximately the distal half of the femur between
the posterior and posterolateral intermuscular lines. The ventral end is con-
tinuous with the origin of M. gastrocnemius pars media, adjacent (medial) to
the origin of M. vastus lateralis pars postica, and adjacent (lateral) to the
insertions of Mm. adductor superficialis and adductor profundus,
INSERTION.—The attachment is to the tendinous raphe in common with M.
flexor cruris lateralis (which see).
INNERVATION.—One or two tiny branches come off the tibial nerve near
the distal end of the main trunk of the sciatic nerve, pass anteriorly deep to
the peroneal nerve, and penetrate the lateral surface.
INDIVIDUAL VARIATION.—In two legs, the branch of the medial division of
the tibial nerve which supplies M. gastrocnemius pars media sends a twig to
the lateral surface of the distal end of M. femorocruralis (in addition to the
usual innervation ),
T. cupido
InprvipuaL _VARIATION.—None of significance.
426 UnIveRSITY OF Kansas Pusts., Mus. Nat. Hist.
P. p. jamesi
DIFFERENCES FROM TypicaL T. pallidicinctus——The muscle is much wider,
extending farther proximally on the femur.
INDIVIDUAL VARIATION.—None of significance.
M. Gastrocnemius, Figs. 12, 13, 15
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Divided into three distinct, widely
separated parts—pars externa, pars interna, and pars media; pars externa:
large; on posterolateral surface of shank; narrow proximally and distally;
bounded anterolaterally by M. flexor perforans et perforatus digiti II and
anteromedially by medial head of M. flexor perforatus digiti III; completely
separate from pars interna and media except for common tendon of insertion;
pars interna: large; on anteromedial surface of shank; narrow distally; bounded
anterolaterally by M. peroneus longus and posteromedially by pars media
(proximally) and medial head of M. flexor perforatus digiti III; broad sheet
of tough connective tissue extending between distal parts of pars externa and
pars interna; covering underlying M. flexor perforatus digiti III (medial head),
somewhat fused with anteroproximal edge of M. peroneus longus; pars media:
small and short; on medial surface of proximal part of shank; deep to tendon
of insertion of M. flexor cruris medialis; bounded anteromedially by pars
interna, posterolaterally by medial head of M. flexor perforatus digiti III, and
proximally by M. femorocruralis; fused to latter, and boundary between the
two difficult to locate.
Oricin.—Pars externa: The short cylindrical tendon fuses with the anterior
half of the distal arm of the tendinous guide loop for M. extensor iliofibularis
and attaches in common with the latter to the posterolateral surface of the
femur immediately proximal to the fibular condyle; the attachment is proximal
(adjacent) to the origin of M. flexor perforans et perforatus digiti II and distal
(adjacent) to the origin of M. flexor perforatus digiti IV and is fused to the
articular capsule.
Pars interna: The proximal end is partly separable into two layers—a super-
ficial longer one and a deep shorter one. The superficial layer attaches fleshily
to the ventral part of the anterior surface of the patella and to the medial half
of the superficial surface of the patellar tendon; this layer slightly overlaps the
distal fleshy end of M. extensor iliotibialis anticus. The deep layer (over-
lapped by the superficial layer) attaches to the medial surface of the inner
cnemial crest, to the rotular crest medial to the latter, to the medial surface of
the proximal part of the tibiotarsus, and (posteroproximal corner) to the
distomedial edge of the patellar tendon and to the articular capsule postero-
medial to the rotular crest; the entire ventral edge is tendinous, the remainder
fleshy.
Pars media: This arises fleshily from an oblique line beginning at the distal
end of the origin of M. femorocruralis (continuous with the latter) and ex-
tending distomedially across the proximal part of the popliteal area to the
proximal edge of the internal condyle, then attaching to the adjacent part of
the articular capsule; this part is adjacent (distal) to the insertion of M. ad-
ductor profundus and adjacent (proximomedial) to the medial head of M.
flexor perforatus digiti IV.
MusScLES AND NERVES OF LEG OF GROUSE 427
INSERTION.—Pars media narrows distally with a narrow tendon along the
posterior edge of the fleshy belly; approximately one third of the way down the
tibiotarsus the fleshy part terminates and the tendon joins the posterior edge
of pars interna, continuing distally in this position. The ossified tendon on
the superficial surface of the distal part of pars interna, continuous posteriorly
with the tendon of pars media, is joined approximately two thirds of the way
down the tibiotarsus by the tendon of pars externa; the fleshy belly of pars
interna ends just below the junction. The ossified tendon on the superficial
surface of the distal part of pars externa extends beyond the fleshy belly and be-
comes flexible before joining the tendon of pars interna and media. The com-
mon tendon (partly ossified) extends along the posterior surface of the
tibiotarsus and widens as it passes posterior to the tibial cartilage, bound to
the latter by a thin tough sheet of connective tissue which attaches to the
edges of the tibial cartilage, thus forming a sheath for the tendon; the tendon
attaches by its edges to the posterior edges of the calcaneal ridges of the
hypotarsus, then continues distally (much reduced in thickness) along the
posterior surface of the tarsometatarsus, enclosing the flexor tendons; the
lateral edge of the tendon attaches to the posterolateral edge of the tarsometa-
tarsus, terminating immediately above the level of the hallux; the medial edge
attaches to the edge of the posterior metatarsal crest; the tendon terminates
as a thin sheet that attaches to the fascia on the sole of the foot. (Hudson, et
al., 1959 consider the posterior metatarsal crest to be an ossified part of the
tendon of M. gastrocnemius. )
INNERVATION.—A branch of the lateral division of the tibial nerve penetrates
the proximal part of the medial surface of pars externa. One or two branches
of the medial division of the tibial nerve pass deep to M. plantaris and penetrate
the deep surface of the posterior part of pars interna. The most proximal
branch of the medial division of the tibial nerve penetrates the lateral surface
of pars media.
INDIVIDUAL VARIATION.—None of significance.
T. cupido
INDIVIDUAL VARIATION.—In one leg, the lateral edge of pars interna over-
laps the proximomedial edge of M. peroneus longus; some fibers attach to the
lateral surface of the inner cnemial crest.
P. p. jamesi
DIFFERENCES FROM TypicaL T. pallidicinctus—The proximal end of pars
interna does not reach the patella.
INprvipuAL VARIATION.—In one leg, an additional twig to pars media arises
from the distal branch to M. femorocruralis,
M. Flexor Perforans et Perforatus Digiti II, Figs. 12, 14
T. vallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Long, slender, and Y-shaped; on
lateral surface of shank; the two heads enclosing M. flexor perforans et per-
foratus digiti III; pasterior head bounded posteriorly by M. gastrocnemius pars
externa; extreme proximal end deep to M, vastus lateralis pars postica; anterior
surface fused to posterior surface of M. flexor perforans et perforatus digiti
5—5835
428 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
III; deep surface fused to tendinous part of lateral head of M. flexor perforatus
digiti IV; anterior head tendinous except for extreme distal end; covered by,
and fused to, posterior edge of M. peroneus longus; fused to anterior surface
of M. flexor perforans et perforatus digiti III; two heads join above middle of
shank; anteroproximal and posterodistal parts of common belly usually ten-
dinous.
Oricin.—Anterior head: This arises by a narrow tendon (partly ossified )
from the distal tip of the outer cnemial crest. The tendon is so intimately
fused with a connective tissue sheet fused to the deep and posterior surfaces
of M. peroneus longus and to the anterior surface of M. flexor perforans et
perforatus digiti III that M. flexor perforans et perforatus digiti II could be
considered to arise from these two muscles. Posterior head: This arises mostly
fleshily from the lateral surface of a compound sheet of tough connective tissue
formed by the fusion of the tendinous posteroproximal comer of M. flexor
perforans et perforatus digiti III, the proximal parts of the tendons of origin
of the lateral heads of Mm. flexor perforatus digiti IV and flexor perforatus
digiti II, the fibular and distal arms of the guide loop for M. extensor iliofib-
ularis, and the lateral part of the articular capsule; a part of the common
tendon of origin of the anterolateral heads of Mm. flexor perforatus digiti III,
flexor perforatus digiti IV, and flexor perforatus digiti II also contributes to
this sheet, which attaches to the lateral surface of the external condyle of the
femur and to the anterolateral surface of the head of the fibula; for conven-
ience in description, this complex connective tissue sheet will hereafter be
termed the femorofibular fascia. The anteroproximal corer of the posterior
head of M. flexor perforans et perforatus digiti II often attaches to the lateral
surface of the vinculum that passes from the femorofibular fascia to the deep
surface of the patellar tendon; the extreme proximal end usually attaches
fleshily to a small area on the femur immediately proximal to the fibular con-
dyle and adjacent (distal) to the attachment of the distal arm of the guide
loop for M. extensor iliofibularis,
INSERTION.—The common belly terminates approximately two thirds of the
way down the shank; the slender ossified tendon begins along the posteromedial
edge of the common belly, continues distally along the posterior surface of
the shank, and becomes flexible before passing through the canal in the tibial
cartilage that lies posteromedial to the canal for M. flexor digitorum longus.
The tendon passes with the tendon of M. flexor perforatus digiti II (medial
to the latter) through a canal in the hypotarsus (see M. flexor perforatus digiti
II); just below the hypotarsus, the tendon becomes superficial to the tendon
of M. flexor perforatus digiti II and farther distally becomes lateral and finally
deep to the latter; the tendon is ossified for most of the length of the tarsometa-
tarsus. At the distal end of this bone, the tendon expands before passing onto
the ventral surface of digit II between the tendons of Mm. flexor perforatus
digiti II and flexor digitorum longus; at the level of the first phalanx, the edges
of the tendon extend dorsally around the tendon of M. flexor digitorum longus
and fuse, forming a sheath around the latter; the latter emerges from the sheath
near the distal end of the first phalanx; the tendon attaches to the proximal
end of the subarticular cartilage ventral to the first interphalangeal joint (the
strongest attachment is on the medial side).
INNERVATION.—The lateral division of the tibial nerve sends twigs into the
posteromedial edge of the posterior head.
MuscLes AND NERVES OF LEG OF GROUSE 429
INDIVIDUAL VARIATION.—In one leg, the fleshy part of the anterior head is
unusually long. In another leg, the anterior head is entirely tendinous. In
one leg, a bundle of fibers of the posterior head attaches to the deep surface
of the distal part of the patellar tendon. In one leg, near the middle of the
tarsometatarsus a rather long and narrow but thick and strong vinculum arises
from the tendon of M. flexor perforatus digiti II and, farther distally, joins the
tendon of M. flexor perforans et perforatus digiti II.
T. cupido
INDIVIDUAL VARIATION.—In one leg, the posterior head arises in part from
the distolateral edge of the patellar tendon and in another, in part from the
superficial surface of the distolateral corner of the patellar tendon.
P. p. jamesi
InpivmuaAL VaRiIATION.—None of significance.
M. Flexor Perforans et Perforatus Digiti III, Figs. 12, 14
T. pallidicinctus
GENERAL DEscCRIPTION AND RELATIONS.—Thick, bipinnate; on lateral sur-
face of proximal part of shank between two heads of M. flexor perforans et
perforatus digiti II; bounded anteriorly by M. peroneus longus; anterior surface
fused with tendinous anterior head of M. flexor perforans et perforatus
digiti II; anterolateral edge somewhat fused to posterior edge of M. peroneus
longus superficial to latter tendon; posterior surface fused to posterior head of
M. flexor perforans et perforatus digiti II; distal part of belly covered by
common belly of latter muscle; posteromedial edge fused to underlying lateral
head of M. flexor perforatus digiti IV; anteromedial edge usually somewhat
fused to underlying M. flexor digitorum longus.
Oricin.—The origin is fleshy and tendinous from the edge of the outer
cnemial crest and fleshy from the superficial surface of the distolateral part
of the patellar tendon; the posteroproximal corner arises tendinously from the
femorofibular fascia.
INsERTION.—The belly narrows abruptly, terminating approximately at the
middle of the shank; the slender ossified tendon extends posterodistally along
the shank, becoming flexible before passing posterior to the tibial cartilage
deep to the tendon of M. gastrocnemius, medial to the tendon of M. flexor
perforatus digiti IV, and superficial to the medial half of the tendon of M.
flexor perforatus digiti III; a thin sheet of connective tissue covers the tendon
and attaches by its edges to the underlying tendon of M. flexor perforatus digiti
III (thus the latter tendon forms a sheath for the tendon of M. flexor perforans
et perforatus digiti II); the tendon is ossified for most of the length of the
tarsometatarsus; at midlength of the latter, the tendon lies between the
tendons of Mm. flexor perforatus digiti IV and flexor perforatus digiti III;
near the distal end of the tarsometatarsus, the tendon becomes lateral and then
deep to the tendon of M. flexor perforatus digiti III and is connected by
a vinculum to the latter (which see). The tendon enters the ventral surface
of digit III between the tendons of Mm. flexor perforatus digiti III and
flexor digitorum longus; after sending a dorsal slip (lateral to the tendon of
M. flexor digitorum longus) to the subarticular cartilage ventral to the first
430 UNIVERSITY OF KANsAs Pusts., Mus. Nat. Hist.
interphalangeal joint, the tendon divides into two branches, between which
emerges the tendon of M. flexor digitorum longus; the lateral branch attaches
to the subarticular cartilage of the second interphalangeal joint and to the
lateral surface of the distal end of the second phalanx; the medial branch
has similar attachments on the medial side of the digit,
INNERVATION.—A branch of the lateral division of the tibial nerve passes
deep to the posterior head of M. flexor perforans et perforatus digiti II and
enters the posteromedial edge of M. flexor perforans et perforatus digiti III.
INDIVIDUAL VARIATION.—In both legs of one specimen, the part arising
from the femorofibular fascia appears as a distinct but short accessory head.
There is no significant individual variation in T. cupido or P. p. jamesi.
M. Flexor Perforatus Digiti IV, Figs. 14, 16
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—On posterolateral aspect of shank
deep to M. gastrocnemius pars externa; bounded medially by medial head of
M. flexor perforatus digiti III, anterolaterally by posterior head of M. flexor
perforans et perforatus digiti II, and anteriorly by M. flexor digitorum longus;
divided into three heads—medial (largest), lateral, and anterolateral (small-
est); tendon of insertion of M. extensor iliofibularis passing between medial
and lateral heads; proximal and anteroproximal parts of lateral head an ex-
tremely thin, flat tendon; anterodistal part of tendon fused to lateral surface
of fleshy part of underlying lateral head of M. flexor perforatus digiti II;
proximal part of tendon fused indistinguishably to tendinous part of underlying
lateral head of M. flexor perforatus digiti II; fleshy part of anterolateral head
anterodistal to lateral head; proximal part of former a long slender tendon
anterior to lateral head; anterior surface of anterolateral head (both fleshy
and tendinous parts) fused to tendon of anterolateral head of M. flexor per-
foratus digiti III; deep surface fused to underlying anterolateral head
(fleshy) of M. flexor perforatus digiti II; common tendon of anterolateral
heads of M. flexor perforatus digiti IV and M. flexor perforatus digiti III
passing medial to tendon of insertion of M. extensor iliofibularis, to peroneal
nerve, and to fibular arm of guide loop for M. extensor iliofibularis; tendon
of M. ambiens inserting on anterolateral surface of this common tendon;
medial head entirely fleshy; medial surface fused to medial head of M. flexor
perforatus digiti III; deep surface fused to medial edge of underlying medial
head of M. flexor perforatus digiti II; medial and lateral heads joined, forming
bipinnate belly (pinnate structure most evident on deep surface); anterolateral
head joined to distolateral part of belly.
Oricin.—The medial head attaches fleshily to the proximal part of the
popliteal area proximal (adjacent) to the origin of M. flexor hallucis longus
and distolateral to the distal end of the origin of M. femorocruralis; the attach-
ment extends laterally onto the posterolateral surface of the femur proximal
(adjacent) to the common attachment of M. gastrocnemius pars externa and
the distal arm of the guide loop for M. extensor iliofibularis; the medial edge
of the origin is fused with part of the tendinous origin of the medial head of
M. flexor perforatus digiti II.
The broad flat common tendon of the lateral head and the lateral head of
MuscLes AND NERVES OF LEG OF GROUSE 431
M. flexor perforatus digiti II fuses to the superficial surface of the fibular arm
of the guide loop for M. extensor iliofibularis and contributes to the femoro-
fibular fascia; consequently the ultimate origin would be the external femoral
condyle and the head of the fibula.
The slender common tendon of the anterolateral head and the anterolateral
heads of Mm. flexor perforatus digiti II and flexor perforatus digiti III passes
deep to the insertional tendon of M. extensor iliofibularis and to the fibular arm
of the guide loop for the latter muscle (to which it partly fuses); the tendon
attaches to a narrow line on the head of the fibula adjacent to the attachment
of the fibular arm of the guide loop and to the deep part of the femorofibular
fascia.
InseRTION.—The slender ossified tendon becomes flexible before it passes
posterior to the tibial cartilage deep to the tendon of M. gastrocnemius, lateral
to the tendon of M. flexor perforans et perforatus digiti III, and superficial to
the lateral half of the tendon of M. flexor perforatus digiti III; a thin sheet of
connective tissue covers the tendon and attaches by its edges to the underlying
tendon of M. flexor perforatus digiti III (thus the latter tendon forms a sheath
for the tendon of M. flexor perforatus digiti IV; this sheath is separate from a
similar sheath surrounding the tendon of M. flexor perforans et perforatus digiti
III); the tendon is again ossified where it passes along the posterolateral sur-
face of the tarsometatarsus posterolateral to the tendon of M. flexor perforans
et perforatus digiti III; near the distal end of the tarsometatarsus the tendon
becomes flexible and expands greatly in width and thickness, and sends a
small slip dorsally, medial to the underlying tendons, that attaches to the sub-
articular cartilage ventral to the trochlea for digit IV; sometimes this slip is
continuous with the retinaculum ventral to the tendon at the level of the
proximal end of the digit. Several more or less distinct sheets of tough con-
nective tissue lie ventral to all of the flexor tendons at the level of the trochleae
and the proximal end of the digits, holding them in place. The tendon nar-
rows as it passes onto the ventral surface of digit IV and soon divides into
three branches; the tendon of M. flexor digitorum longus emerges between
the medial and middle branches. The lateral branch attaches to the sub-
articular cartilage ventral to the first interphalangeal joint and is also bound
by connective tissue to the ventrolateral surface of the first phalanx. A dorsal
slip arises at the point of divergence of the lateral and middle branches and
attaches to the subarticular cartilage of the first interphalangeal joint. The
middle branch attaches to the subarticular cartilage of the second joint. The
medial branch, after sending dorsal slips to each of the first two subarticular
cartilages, attaches to the subarticular cartilage of the third interphalangeal
joint.
INNERVATION.—The posterior division of the tibial nerve sends a branch
into the posterior edge of the medial head, then passes between the latter and
the medial head of M. flexor perforatus digiti III; as it extends distally it gives
off twigs to the medial surface of the medial head, to the deep surface of the
lateral head, and to the deep surface of the anterolateral head.
INDIVIDUAL VARIATION.—In one leg, an additional branch arises from the
tibial nerve at the level of origin of the posterior division and enters the
posterior surface of the medial head; a twig from this branch anastomoses
with the first twig of the posterior division to the same head; a branch of the
432 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
medial division joins the posterior division distal to the origin of the twigs to
the medial head but proximal to the origin of the twigs to the other heads.
T. cupido
InpivipuaL VARIATION.—None of significance.
P. p. jamesi
INDIvipUAL VARIATION.—In four legs, a tiny vinculum connects with the
tendon of M. flexor digitorum longus (which see).
M. Flexor Perforatus Digiti III, Figs. 13, 14, 15, 16, 17
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Divided into two widely separated
heads—medial and anterolateral—with completely separate bellies but with
common insertional tendon; small anterolateral head on lateral aspect of thigh
deep to M. flexor perforans et perforatus digiti II and posterior to M. flexor
digitorum longus; fleshy part of head distolateral to belly of M. flexor per-
foratus digiti IV; fleshy part fused to lateral edge of belly of M. flexor per-
foratus digiti II; proximal part of head a slender ossified tendon fused to
anterior edge of both fleshy and tendinous parts of anterolateral head of M.
flexor perforatus digiti IV and to lateral edge of anterolateral head of M.
flexor perforatus digiti II; this tendon passing deep to tendon of insertion of
M. extensor iliofibularis and to peroneal nerve; large medial head on postero-
medial surface of thigh anterior to medial edge of M. gastrocnemius pars ex-
terna, lateral to M. gastrocnemius pars media, and medial to M. flexor per-
foratus digit IV; fused to medial surface of medial head of latter and to
medial edges of Mm. flexor perforatus digiti II and flexor hallucis longus;
proximal end of head tendinous.
Oricin.—The medial head attaches tendinously to the medial part of the
popliteal area in common with the medial head of M. flexor perforatus digiti
II and with the medial edges of Mm. flexor perforatus digiti IV (medial head)
and flexor hallucis Jongus; and is also fused to the articular capsule. The
anterolateral head arises in common with the anterolateral heads of Mm.
flexor perforatus digiti II and flexor perforatus digiti IV (see account of
latter).
INSERTION.—The short unossified tendon of the anterolateral head and the
longer ossified tendon of the medial head join (after the latter becomes flexi-
ble) a short distance above the tibial cartilage, forming a broad flat common
tendon that passes posterior to the tibial cartilage (in a shallow groove of the
latter); the main part of the tendon is deep to the tendons of Mm. flexor per-
foratus digiti IV and flexor perforans et perforatus digiti III, but forms separate
thin sheaths around these two tendons at the level of the tibial cartilage. A
thin sheet of connective tissue covers these three tendons and attaches by its
edges to the tibial cartilage, forming a sheath for them. These three tendons
pass through the superficial groove in the hypotarsus deep to the tendon of M.
gastrocnemius; the tendon of M. flexor perforatus digiti III is ossified for most
of the length of the tarsometatarsus; a short distance below the hypotarsus,
the anterior branch of the tendon of M. peroneus longus attaches broadly to
the lateral edge of the tendon of M. flexor perforatus digiti III. In the proximal
MuscLes AND NERVES OF LEG OF GROUSE 433
part of the tarsometatarsus the tendon is deep to the tendon of M. flexor
perforans et perforatus digiti III, but farther distally becomes medial and then
superficial to the latter and lateral to the tendon of M. flexor perforans et
perforatus digiti II; near the distal end of the tarsometatarsus a narrow but
strong vinculum extends from the lateral edge of the tendon somewhat distally
to the lateral edge of the tendon of M. flexor perforans et perforatus digiti III.
At the distal end of the tarsometatarsus the tendon expands before entering
the ventral surface of digit III where it soon divides into two branches, be-
tween which emerge the tendons of Mm. flexor perforans et perforatus digiti
III and flexor digitorum longus; the lateral branch attaches to the subarticular
cartilage ventral to the first interphalangeal joint and to the lateral surface
of the distal end of the first phalanx; the medial branch has similar attach-
ments on the medial side of the digit.
INNERVATION.—The posterior division of the tibial nerve passes between
the medial heads of M. flexor perforatus digiti III and M. flexor perforatus
digiti IV and sends a twig to the lateral surface of the former, then passes
deep to the common belly of M. flexor perforatus digiti IV and sends a twig
to the posterior surface of the anterolateral head of M. flexor perforatus digiti
Ill.
INDIVIDUAL _VARIATION.—None of significance.
T. cupido
InprvmuAL VARIATION.—In one leg, an extra branch (immediately distal
to the branch to M. gastrocnemius pars media) of the medial division of the
tibial nerve penetrates the medial surface of the proximal end of the medial
head.
P. p. jamesi
InprvipuaAL VaRIATION.—None of significance.
M. Flexor Perforatus Digiti II, Figs. 15, 17
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Bipinnate; on posterior aspect
of shank deep to M. flexor perforatus digiti IV and between two heads of
M. flexor perforatus digiti III; bounded anteriorly by Mm. flexor digitorum
longus and flexor hallucis longus; proximal part divided into three small heads—
medial, lateral, and anterolateral; medial and proximal parts of medial head
tendinous and extremely thin except for ossified medial edge; proximal part of
lateral head tendinous and lateral to insertional tendon of M. extensor ilio-
fibularis; both tendinous and fleshy parts fused to overlying tendon of M. flexor
perforatus digiti IV; narrow anterolateral head fused to overlying anterolateral
head of latter muscle and (anterolateral edge) to ossified tendon of antero-
lateral head of M. flexor perforatus digiti III; lateral edge of common belly
fused to latter head; medial edge of muscle fused to medial heads of Mm.
flexor perforatus digiti IV and flexor perforatus digiti III and to M. flexor
hallucis longus.
Oricin.—The medial head attaches by a slender ossified tendon to the
medial part of the popliteal area in common with the medial head of M.
flexor perforatus digiti III and with the medial edges of Mm. flexor perforatus
434 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
digiti IV (medial head) and flexor hallucis longus; this head is also fused
to the articular capsule. The above-mentioned ossified part of the tendon is
situated at the junction of M. flexor perforatus digiti II and M. flexor perforatus
digiti III (medial head) and could be considered to be a part of the latter
rather than the former. The flat tendon of the lateral head arises in common
with the lateral head of M. flexor perforatus digiti IV (which see). The
anterolateral head arises in common with the anterolateral heads of Mm.
flexor perforatus digiti IV and flexor perforatus digiti III (see former).
INsERTION.—The short, slender, ossified tendon becomes flexible and passes
through the canal in the tibial cartilage that lies medial to the canal for M.
flexor hallucis longus and lateral to the canals for Mm. flexor digitorum longus
and flexor perforans et perforatus digiti II. The tendon passes with the tendon
of M. flexor perforans et perforatus digiti II (lateral to latter) through the canal
in the hypotarsus that is deep to the groove for M. flexor perforatus digiti III
and superficial to the canal for M. flexor digitorum longus; the former canal
has a bony floor and sides but a fibrous roof; a fibrous partition subdivides
the proximal half of this canal, forming a separate channel for each tendon.
The tendon is ossified for most of the length of the tarsometatarsus and is
situated lateral (adjacent) to the posterior metatarsal crest; immediately below
the hypotarsus, the tendon becomes situated deep to the tendon of M. flexor
perforans et perforatus digiti II and farther distally becomes situated medial
and finally superficial to the latter; at the distal end of the tarsometatarsus
the tendon expands greatly and its edges (thick) pass dorsally around the
underlying flexor tendons and become continuous with the subarticular cartilage
ventral to the trochlea for digit II. The tendon extends onto the ventral surface
of digit II and attaches by its edges to the ventromedial and ventrolateral sur-
faces of the proximal part of the first phalanx (the lateral edge extending
farthest distally); the tendons of Mm. flexor perforans et perforatus digiti II
and flexor digitorum longus emerge from the distal end of the tendon of M.
flexor perforatus digiti II.
INNERVATION.—The posterior division of the tibial nerve passes between
the medial heads of Mm. flexor perforatus digiti ITI and flexor perforatus digiti
IV and gives a twig to the superficial surface of each of the three heads of M.
flexor perforatus digiti If and sometimes gives another twig to the superficial
surface of the distal part of the common belly.
INpDIvipUAL VaARIATION.—In one leg, a vinculum connects the tendon with
that of M. flexor perforans et perforatus digiti II (which see).
T. cupido
InpivipvaAL VARIATION.—The canal in the hypotarsus through which the
tendon passes has a bony (instead of fibrous) roof in one leg.
P. p. jamesi
INDIvmUAL VARIATION.—The variation given above for T. cupido is found
in both legs of one specimen.
Muscies AND NERVES OF LEG OF GROUSE 435
M. Flexor Hallucis Longus, Figs. 15, 19A
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Elongate and tapering; on posterior
aspect of shank deep to M. flexor perforatus digiti II and to proximal end of
medial head of M. flexor perforatus digiti IV; bounded anterolaterally by M.
flexor digitorum longus and anteromedially by M. plantaris; tendinous antero-
medial surface of proximal end fused to common tendon of origin of medial
heads of Mm. flexor perforatus digiti III and flexor perforatus digiti II; belly
ending approximately halfway down shank.
Oricin.—The origin is fleshy and tendinous (anteromedial surface) from
the popliteal area immediately distal to the origin of the medial head of M.
flexor perforatus digiti IV, extending laterally to the area immediately proximal
to the external femoral condyle (medial to the origin of M. gastrocnemius
pars externa); the muscle also arises from the proximal end of the posterior
part of the articular capsule.
INsERTION.—The slender ossified tendon becomes flexible and passes through
the canal in the tibial cartilage that lies lateral to the canal for M. flexor
perforatus digiti II, then passes through a slight groove in the lateral surface
of the hypotarsus and becomes ossified again; midway of the tarsometatarsus,
the tendon becomes superficial to the tendon of M. flexor digitorum longus
and is connected with the latter by an extensive vinculum, which extends from
the deep surface and lateral edge of the tendon of M. flexor hallucis longus
distally to the superficial surface of the tendon of M. flexor digitorum longus;
the tendon continues, unossified and considerably reduced in size, distally
medial to the tendon of M. flexor digitorum longus, and passes through the
flexor groove of the first metatarsal anterolateral (adjacent) to the tendon of
M. flexor hallucis brevis, then passes deep to the terminal expansion of the
latter onto the ventral surface of the hallux; the tendon emerges from under
the end of the tendon of M. flexor hallucis brevis and attaches to the ventral
surface of the ungual phalanx; a weak dorsal slip attaching to the ventral sur-
face of the distal end of the first phalanx is usually present.
INNERVATION.—A branch of the medial division of the tibial nerve passes
along the medial edge of the muscle, giving several twigs into it.
InpivipvaL VaARIATION.—None of significance in any of the three species
studied.
M. Plantaris, Figs. 15, 19A
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Elongate and tapering; on pos-
teromedial surface of tibiotarsus; bounded medially by M. gastrocnemius pars
interna and tendon of M. flexor cruris medialis, posteriorly by M. gastrocnemius
pars media and medial head of M. flexor perforatus digiti III, posterolaterally
by M. flexor hallucis longus; medial to M. flexor digitorum longus; anterolateral
surface of proximal end often slightly overlapping and fused to posterior sur-
face of medial end of M. popliteus; belly terminating above middle of shank.
Oricin.—The origin is fleshy and tendinous (distal edge only) from an
elongate area on the posteromedial surface of the proximal end of the
tibiotarsus adjacent to the insertion of M. popliteus.
436 UnIversiTy OF Kansas Pusts., Mus. Nat. Hist.
INsERTION.—The long, slender, ossified tendon extends along the postero-
medial aspect of the tibiotarsus and becomes flexible just before attaching to
the proximomedial part of the tibial cartilage. The tibial cartilage is a large,
mostly cartilaginous pad fitting closely over the posterior surface of the intra-
tarsal joint; the distomedial corner is ossified. This cartilage is perforated by
the tendons of several flexor muscles; the distal end of the cartilage attaches
to the posteroproximal corner of the tarsometatarsus.
INNERVATION.—A branch of the medial division of the tibial nerve penetrates
the lateral surface.
INDIVIDUAL VARIATION.—In one leg, a small bundle of fibers separates from
the proximal end of the muscle, forming a short accessory head which at-
taches, separately from the remainder, to the articular capsule posteroproximal
to the main origin; a blood vessel passes between the main and accessory
heads.
T. cupido
INDIVIDUAL VARIATION.—In one leg, a small bundle of fibers arises from
the medial collateral ligament. In another leg, the nerve to M. gastrocnemius
pars interna passes through a gap in the origin of M. plantaris rather than
distal to its origin.
P. p. jamesi
INDIVIDUAL VARIATION.—The nerve branch supplying M. gastrocnemius
pars interna gives a minute twig to the deep surface of the free belly of M.
plantaris in one instance.
M. Flexor Digitorum Longus, Figs. 14, 16, 17, 19A
T. pallidicinctus
GENERAL DESCRIPTION AND ReELATIONS.—Relatively broad; bipinnate; on
posterolateral surface of tibiotarsus; bounded posteromedially by M. flexor
hallucis longus, posteriorly by M. flexor perforatus digiti II and anterolateral
head of M. flexor perforatus digiti III, laterally by Mm. flexor perforans et
perforatus digiti III and flexor perforans et perforatus digiti II, and antero-
laterally by Mm. peroneus brevis and tibialis anticus; anterior surface of lateral
part of distal half of common belly fused to M. peroneus brevis; divided into
three heads—posterior (largest), lateral, and medial; posterior head on pos-
terior surface of head of fibula; overlapping and fused to lateral end of M.
popliteus; proximomedial comer deep to latter; lateral head on lateral surface
of fibula; lateral and posterior heads separated by insertion of M. extensor
iliofibularis; these two heads joined immediately distal to insertion of latter;
medial head on posterior surface of tibiotarsus; group of blood vessels and
nerves passing between medial and posterior heads; these two heads joined
several mm. distal to junction of lateral and posterior heads; deep surface of
insertional tendon near distal end of tarsometatarsus serving as origin for M.
lumbricalis.
Oricin.—Posterior head: This arises fleshily from the posterior surface of
the fibula beginning almost at the proximal end and from the medial surface
of the fibula beginning deep to the distal part of M. popliteus. Lateral head:
This arises fleshily (sometimes partly tendinously) from the lateral surface of
the fibula proximal to the fibular tubercle. Some fibers arise from the distal
Muscies AND NERVES OF LEG OF GROUSE 437
edge of the tendon of insertion of M. extensor iliofibularis. Medial head: This
arises fleshily from the posterior surface of the tibiotarsus just medial to the
distal part of the posterior head, distal to M. popliteus, and either lateral or
distolateral to the origin of M. plantaris, Distal to the junction of the three
heads, the muscle arises fleshily from the posterior surface of the tibiotarsus
(except the distal part) and from the medial and posterior surfaces of the
fibula.
INsERTION.—The slender ossified tendon becomes flexible and _ passes
through the canal in the tibial cartilage that lies anterolateral to the canal
for M. flexor perforans et perforatus digiti II and anteromedial to the canal
for M. flexor perforatus digiti II, then passes through the bony canal of the
hypotarsus that is deep to all the other flexor tendons; the tendon ossifies
again and lies adjacent (lateral) to the posterior metatarsal crest; the vinculum
from the tendon of M. flexor hallucis longus fuses extensively to the superficial
surface of the present tendon a short distance below the midpoint of the
tarsometatarsus; the tendon is considerably broader below this point than
above it. At the level of the first metatarsal, the tendon divides into three
branches (unossified) that diverge, each passing through a groove on the
ventral surface of the subarticular cartilages ventral to the trochleae, then
pass onto the ventral surfaces of digits II, III, and IV. On digit IV the
tendon gives off two dorsal fibro-elastic slips before attaching to the ventral
surface of the ungual phalanx; one slip attaches to the subarticular cartilage
ventral to the third interphalangeal joint, the other to the subarticular cartilage
of the fourth joint and may also attach in part to the distal end of the fourth
phalanx. On digit III the tendon gives off two dorsal slips before attaching
to the ventral surface of the ungual phalanx; one slip attaches to the sub-
articular cartilage of the second interphalangeal joint, the other to the sub-
articular cartilage of the third joint and may also attach in part to the distal
end of the third phalanx. On digit II the tendon gives off one dorsal slip
before attaching to the ventral surface of the ungual phalanx; the slip attaches
to the subarticular cartilage of the second interphalangeal joint and may also
attach in part to the distal end of the second phalanx.
INNERVATION.—A branch of the medial division of the tibial nerve pene-
trates the medial surface of the posterior head.
InprvipuaL VariATION.—In half the legs, the proximal end of the lateral
head is notched for the passage of the peroneal nerve; the main part of the
head lies medial to this nerve; the short fleshy slip lateral to this nerve arises
by a long, slender, and extremely weak tendon from connective tissue sur-
rounding the femorotibiotarsal joint. In one leg, a bundle of fibers separates
from the lateral head and attaches to the terminal four mm. of the anterior
(proximal) edge of the tendon of M. extensor iliofibularis. Each of the
following variations occurs in several legs: a third dorsal slip on digit IV
attaches to the distal end of the fourth phalanx in some legs and to the sub-
articular cartilage of the fourth joint in other legs; a third dorsal slip on
digit III attaches to the distal end of the third phalanx in some legs and
to the subarticular cartilage of the third joint in other legs; a second dorsal
slip on digit II attaches to the distal end of the second phalanx in some legs
and to the subarticular cartilage of the second joint in other legs.
438 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
T. cupido
INDIVIDUAL VARIATION.—The dorsal slips of insertion show variations
similar to those noted above for T. pallidicinctus.
P. p. jamesi
INDIVIDUAL VARIATION.—In one leg, the proximal end of the lateral head
is notched for the passage of the peroneal nerve. The dorsal slips of insertion
show variations smiliar to those given above for T. pallidicinctus. In four legs,
a tiny vinculum extends from the lateral edge of the branch of the tendon on
digit IV to the lateral edge of the underlying medial branch of the tendon
of M. flexor perforatus digiti IV at the level of the second phalanx.
M. Popliteus, Fig. 19B
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Extremely short but relatively
broad and thick; on posterior surface of proximal end of tibiotarsus; extending
distomedially from proximal part of fibula; deep to M. flexor hallucis longus;
iateral end overlapped by, and fused to, posterior head of M. flexor digitorum
longus; medial end often slightly overlapped by, and fused to, M. plantaris;
medial end (insertion) much wider than lateral end (origin).
Oricin.—The origin is fleshy and tendinous (superficial surface) from the
medial surface of the fibula near the proximal end.
INsERTION.—The attachment is fleshy to the posterior surface of the
proximal end of the tibiotarsus adjacent (lateral) to the origin of M. plantaris.
INNERVATION.—A branch of the medial division of the tibial nerve penetrates
the posterior surface,
InpivipuaL VARIATION.—None of significance in any of the three species
studied.
M. Peroneus Longus, Figs. 12, 13
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Large; on anterolateral surface of
shank; bounded medially by M. gastrocnemius pars interna and posterolaterally
by Mm. flexor perforans et perforatus digiti III and flexor perforans et per-
foratus digiti II; proximal three fourths of posteromedial part (covered by M.
gastrocnemius pars interna) aponeurotic and tightly fused to medial surfaces
of underlying Mm. tibialis anticus and extensor digitorum longus; proximal
part of fleshy belly somewhat fused to anterior surface of underlying M.
tibialis anticus; posterolateral surface strongly fused to aponeurotic medial
head of M. flexor perforans et perforatus digiti II and slightly fused to antero-
lateral edge of M. flexor perforans et perforatus digiti III.
Oricin.—The muscle arises by fleshy and tendinous fibers from the edges
of the inner and outer cnemial crests; the extreme proximal end arises either
fleshily or aponeurotically from the rotular crest between the cnemial crests;
the posteromedial edge (aponeurotic except distal one fourth fleshy) arises
from the anteromedial intermuscular line.
INSERTION.—The narrow ossified tendon on the superficial surface of the
distal part of the fleshy belly extends several mm, beyond the belly where it
MuSCLES AND NERVES OF LEG OF GROUSE 439
becomes flexible and divides into two branches. The short, broad posterior
branch attaches broadly to the proximolateral corner of the tibial cartilage.
The narrow anterior branch passes along the lateral surface of the tibiotarsus,
through a strong retinaculum immediately proximal to the external condyle,
and crosses the lateral surface of the joint, where it is covered by connective
tissue nearly as tough as, and continuous with, the retinaculum; the tendon
attaches broadly to the lateral edge of the ossified tendon of M. flexor per-
foratus digiti III a short distance below the hypotarsus.
INNERVATION.—The peroneal nerve sends twigs to the deep surface.
InprivipvaL VARIATION.—In both legs of two specimens, the extreme
proximal end extends proximal to the rotular crest and attaches fleshily to
the superficial surface of the distal end of the patellar tendon.
T. cupido
INDIVIDUAL VARIATION.—None of significance.
P. p. jamesi
INDIVIDUAL VARIATION.—One leg shows the variation described above for
T. pallidicinctus.
M. Tibialis Anticus, Figs. 14, 15, 16, 19E, 20N
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Thick; on anterior aspect of
thigh deep to M. peroneus longus; bounded posteriorly by M. extensor digi-
torum longus and posterolaterally by Mm. flexor digitorum longus and peroneus
brevis; divided into two heads—tibial and femoral; small femoral head
adjacent to posterolateral surface of much larger tibial head; two heads
joined near midpoint of fleshy part of muscle, forming bipinnate belly (pinnate
structure most evident on deep surface); proximal part of femoral head
situated between outer cnemial crest and head of fibula; proximal part of
anterior surface of tibial head somewhat fused to overlying M. peroneus
longus; medial surface fused to aponeurosis of latter,
Oricin.—Tibial head: This arises by fleshy and tendinous fibers from the
edge of the inner cnemial crest, from the rotular crest between the inner and
outer cnemial crests, and from the anterior surface, distal edge, and posterior
surface of the outer cnemial crest; the attachment may or may not extend
onto the superficial surface of the distal part of the patellar tendon; the
attachment is adjacent to the origin of the underlying M. extensor digitorum
longus. Femoral head: This arises by a slender tendon from the notch in
the distal end of the external condyle of the femur,
INSERTION.—The slender ossified tendon extends along the anterior surface
of the distal end of the tibiotarsus and passes through a large, strong, ob-
lique retinaculum (superficial to the supratendinal bridge); the lateral end
of the retinaculum attaches to the lateral end of the supratendinal bridge;
the medial end attaches immediately proximal to the medial end of the
bridge. The tendon widens and becomes flexible as it passes across the an-
terior surface of the intratarsal joint, then narrows and attaches to the
tubercle on the anterior surface of the proximal part of the tarsometatarsus
between Mm. extensor hallucis longus and extensor brevis digiti IV. The
440 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
distalmost bundle of tendinous fibers does not attach to the tubercle, but
extends distally along the anterior surface of the tarsometatarsus and attaches
to the latter a few mm. distal to the tubercle, forming an accessory insertion.
A part of the peroneal nerve passes between the main and accessory inser-
tions,
INNERVATION.—A variable number of branches of the peroneal nerve pene-
trate the lateral surface of the femoral head; a variable number of branches of
the same division pass deep to the femoral head and enter the posterior edge
of the tibial head.
INDIVIDUAL VARIATION.—In one leg, the accessory insertion is absent.
T. cupido
INnDIvibUAL VARIATION.—None of significance.
P. p. jamesi
DIFFERENCES FROM Typicat T, pallidicinctus—The origin of the tibial head
does not extend onto the patellar tendon.
INDIVIDUAL _VARIATION.—The accessory insertion is absent in one leg.
M. Extensor Digitorum Longus, Figs. 15, 17
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Bipinnate; on anterior surface
of tibiotarsus deep to M. tibialis anticus; bounded laterally by M. peroneus
brevis; lateral edge usually slightly fused to proximal half of latter; medial
surface fused to aponeurosis of M. peroneus longus.
Oricin.—The muscle arises fleshily from the lateral surface of the inner
cnemial crest, from the rotular crest between the cnemial crests (deep to
the attachment of M. tibialis anticus), from the basal (medial) half of the
anterior surface of the outer cnemial crest, and from the anterior surface of
the tibiotarsus (except the distal part) between the anteromedial and an-
terolateral intermuscular lines; proximal to the anterolateral intermuscular
line, the origin usually extends almost to the lateral edge of the tibiotarsus.
INSERTION.—The ossified tendon extends along the mid-anterior surface
of the distal part of the tibiotarsus deep to the tendon of M. tibialis anticus
and passes under the supratendinal bridge, becoming flexible and widening
slightly as it crosses the anterior surface of the intratarsal joint; the tendon
narrows again and passes through a smal] but strong retinaculum on the
anterior surface (medial to midline) of the proximal part of the tarsometatar-
sus; the retinaculum is immediately proximal and medial to the insertion of
M. tibialis anticus. The tendon ossifies again as it passes down the anterior
surface of the tarsometatarsus and bifurcates near the midpoint of the latter;
the lateral branch soon bifurcates again; of these three branches, which are
ossified for some distance, the lateral one passes onto the dorsal surface
of digit IV, the middle one passes onto the dorsolateral surface of digit III,
and the medial one subdivides (at the level of the trochleae) into three
branches—one passing onto the dorsal surface of digit III and two passing
onto the dorsal surface of digit II. At the level of the metatarsophalangeal
joints, all of these tendons are interconnected by strong sheets of connective
tissue and it is often difficult exactly to delimit the tendons at this level, On
Muscles AND NERVES OF LEG OF GROUSE 44]
the digits, tough connective tissue binds the tendons to the phalanges; this
is most pronounced at the interphalangeal joints. The tendons are distinct
on the first phalanx of each digit, but are often poorly defined farther distally.
On digit IV the tendon subdivides into branches that attach to the proximal
ends of the ungual, fourth, third, and (usually) second phalanges. On
digit III the lateralmost tendon bifurcates, with one branch attaching to the
ungual phalanx and the other to the proximal end of the third phalanx; the
medial tendon attaches to the proximal end of the second phalanx, On
digit II the originally medial tendon passes underneath and then lateral to
the other tendon and attaches to the ungual phalanx; the other tendon at-
taches to the proximal end of the second phalanx.
INNERVATION.—One or more branches of the peroneal nerve enter the
lateral edge.
INpIvipvaAL VARIATION.—In four legs, the lateral branch of the trifurcated
tendon is not ossified at all.
T. cupido
INpIvmuAL VARIATION.—In a few cases, the muscle does not come in
contact with M. peroneus brevis,
P. p. jamesi
DIFFERENCES FROM TypicaL T. pallidicinctus——The belly is shorter. The
lateral branch of the tendon on the tarsometatarsus is not ossified (true also
of some legs of Tympanuchus).
INDIVIDUAL VARIATION.—In several legs, the muscle also arises from the
distal part of the posterior surface of the outer cnemial crest.
M. Peroneus Brevis, Figs. 14, 16, 17, 18, 19A
T. pallidicinctus
GENERAL DEsCRIPTION AND RELATIONS.—Small; on lateral surface of
distal part of tibiotarsus; mainly anterior to fibula; bounded posteriorly and
laterally by M. flexor digitorum longus (fused with latter), anteriorly by M.
tibialis anticus, and anteromedially by M. extensor digitorum longus (usually
slightly fused to latter).
Oricin.—The muscle arises by fleshy and tendinous fibers from the medial
and anterior surfaces of the fibula beginning a short distance below the
distal end of the fibular crest and from the anterolateral surface of the
tibiotarsus anterior to the fibula; the anteromedial edge attaches to the
anterolateral intermuscular line.
INsERTION.—The short, slender, ossified tendon passes along the antero-
lateral surface of the tibiotarsus and through a retinaculum immediately proxi-
mal and anteromedial to the retinaculum for the anterior branch of the
tendon of M. peroneus longus; the tendon becomes flexible and widens as it
passes across the lateral surface of the intratarsal joint deep to the tendon
of M. peroneus longus, turning posteriorly and attaching to the proximolateral
corer of the hypotarsus.
INNERVATION.—The superficial peroneal branch of the peroneal nerve gives
one or two twigs to the anterior surface of the proximal part.
INDIVIDUAL VARIATION.—None of significance.
442 UnrIverSITY OF Kansas Pusts., Mus. Nar. Hist.
T. cupido
InpIvipuAL VARIATION.—In a few legs, the muscle does not come in contact
with M. extensor digitorum longus.
P. p. james
INDIVIDUAL VARIATION.—None of significance.
o
M. Extensor Hallucis Longus, Figs. 19E, 20N
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Slender and elongate; proximal
part on anterior surface of tarsometatarsus medial to anterior metatarsal
groove; near midlength of tarsometatarsus, muscle twisted onto medial surface
of latter; divisible into two heads—proximal and distal; belly of proximal
head (largest) ending at level of twisting onto medial surface of bone; short
distal head beginning at this point deep to tendon of proximal head and
soon joining latter tendon.
Oricin.—Proximal head: This arises fleshily from the anterior surface of
approximately the proximal half of the tarsometatarsus medial to the anterior
metatarsal groove; the proximal end is partly medial to and partly deep to the
retinaculum for M. extensor digitorum longus; some fibers arise from the
extreme distal edge of the main insertion of M. tibialis anticus; the distal end
of the belly is unattached. Distal head: This arises fleshily from the medial
surface of the tarsometatarsus proximal to the first metatarsal and deep to the
tendon of the proximal head.
INSERTION.—The slender tendon of the proximal head, which begins along
the medial edge of the distal part of the belly, soon fuses with the superficial
surface of the distal head (ossified here); the common tendon (unossified)
passes onto the dorsal (proximal) surface of the first metatarsal, where it
passes through a retinaculum, then passes along the dorsal surface of the
hallux (bound by strong connective tissue to the metatarsophalangeal joint),
attaching to the dorsal surface of the ungual phalanx.
INNERVATION.—The branch of the deep peroneal nerve that passes medial
to the main insertion of M. tibialis anticus gives one or two twigs into the
proximal part of the proximal head. No supply to the distal head was found,
but see below.
InpivipvaL VaRIATION.—In one leg, the proximal end of the distal head
is fused to the distal end of the belly of the proximal head, whereas in three
legs, a distinct gap separates the fleshy parts of the two heads. The following
variations, each found in one leg, pertain to the relationship of the origin
of the proximal head to the retinaculum for M. extensor digitorum longus:
the origin does not extend proximally medial to the retinaculum; the origin
does not extend proximally deep to this retinaculum; a part of the proximal
end extends proximally lateral to this retinaculum (in this instance there is
an unusually wide gap between the retinaculum and the insertion of M.
tibialis anticus). In one leg, the distalmost fibers of the distal head do not
join the common tendon but insert independently on the articular capsule
of the metatarsophalangeal joint (deep to the common tendon).
MuscLes AND NERVES OF LEG OF GROUSE 443
T. cupido
InprvvAL VARIATION.—The relationship between the two heads varies
as follows: the proximal end of the distal head may be fused to the distal
end of the belly of the proximal head; the proximal end of the distal head
may begin anterior (adjacent) to the distal end of the belly of the proximal
head; there may be a distinct gap between the fleshy parts of the two heads.
In two legs, there is no origin from the insertion of M. tibialis anticus. In
one leg, a small accessory bundle of fleshy fibers arises from the proximal
end of the first metatarsal (widely separated from the origin of the distal
head), passes through the retinaculum deep to the common tendon and
attaches to the dorsal surface of the articular capsule of the metatarsopha-
langeal joint; thus this bundle is completely separate from the remainder
of the muscle. In two legs, the same nerve branch that gives twigs into
the proximal head also gives off (much farther distally) a twig that enters
the distal head.
P. p. jamesi
INDIVIDUAL VARIATION.—The proximal end of the distal head may begin
anterior (adjacent) to the distal end of the belly of the proximal head. In
four legs, the origin of the proximal head does not extend proximally medial
to the retinaculum for M. extensor digitorum longus; in one of these legs,
a part of the proximal end extends proximally lateral to this retinaculum.
The distalmost fibers of the distal head do not join the common tendon but
insert independently on the dorsal surface of the articular capsule of the
metatarsophalangeal joint in four legs; in another leg, the entire distal head
has the latter insertion (consequently the two heads are completely separate).
M. Abductor Digiti II, Figs. 19E, 20N
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Short; on medial surface of distal
part of tarsometatarsus; proximal end adjacent (anterior) to distal head of
M. extensor hallucis longus.
Onicin.—The origin is fleshy from the medial surface of the distal part
of the tarsometatarsus anterior (adjacent) to the first metatarsal and from
the anteromedial surface of the basal half of the first metatarsal.
INSERTION.—The flat tendon passes over the medial surface of the trochlea
for digit II and attaches to the medial surface of the proximal end of the first
phalanx of digit II; the tendon is fused with the articular capsule,
INNERVATION.—The compound nerve formed by the fusion of a branch
of the superficial peroneal nerve with a branch of the deep peroneal nerve
gives a twig to the anterolateral edge of the muscle.
InprvipvuaL VARIATION.—In some cases, the twig arises from the deep
peroneal branch alone (which is not joined by the superficial peroneal
nerve).
T. cupido
INpIviDUAL VARIATION.—In one leg, some of the fleshy fibers arising from
the first metatarsal insert independently on the medial surface of the trochlea
for digit II (deep to the main part of the muscle).
6—5835
444 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
P. p. jamesi
InpivipvAL VARIATION.—None of significance.
M. Extensor Brevis Digiti III (M. extensor proprius digiti III), Figs. 19E, 20N
T. pallidicinctus
GENERAL DESCRIPTION AND ReELATIONS.—Short and relatively broad (nar-
row proximally); on mid-anterior surface of distal part of tarsometatarsus;
tendon of insertion fused with articular capsule.
Oricin.—The origin is fleshy from the mid-anterior surface of the distal
part of the tarsometatarsus ending a short distance proximal to the trochlea
for digit III.
INSERTION.—The flat tendon passes over the trochlea for digit III and
attaches to the dorsal surface of the proximal end of the first phalanx of
digit III.
INNERVATION.—The compound nerve formed by the fusion of a branch
of the superficial peroneal nerve with a branch of the deep peroneal nerve
gives a twig to the proximal end of the muscle.
INDIVIDUAL VARIATION.—In some cases, the twig arises from the deep
peroneal branch alone (which is not joined by the superficial peroneal
nerve). The individual variation is insignificant in T. cupido and P. p. jamesi.
M. Extensor Proprius Digiti III (Not found by Hudson, et al.), Fig. 20N
T.. pallidicinctus and T. cupido
Absent in both species.
P. p. jamesi
This atypical muscle was found in only two legs (P.p. 1L and 4L). The
following description applies to P. p. 4L (Fig. 20N).
GENERAL DESCRIPTION AND RELATIONS.—Small but well developed; fleshy
part 14>< 13 mm.; proximal end narrower; on mid-anterior surface of tar-
sometatarsus between Mm. extensor brevis digiti IV and extensor hallucis longus
and mostly proximal to M. extensor brevis digiti III; tendinous distal part
superficial to latter; fleshy belly ending immediately distal to proximal end of
latter.
Oricin.—The origin is fleshy from a narrow elongate area on the mid-
anterior surface of the tarsometatarsus between Mm. extensor brevis digiti IV
and extensor hallucis longus, beginning at the distal end (bony) of the
elongate accessory insertion of M. tibialis anticus. The distal part of the
belly is free.
INSERTION.—The attachment is by a thin, wide (relative to belly) tendon
to the superficial surface of M. extensor brevis digiti III.
INNERVATION.—Not found.
INprIvipuaL VARIATION.—In P.p. 1L, the muscle is less well developed. The
fleshy belly is 1 X 5 mm. It arises from the lateral edge of M. extensor hallucis
longus. The extremely slender insertional tendon attaches as above.
Muscies AND NERVES OF LEG OF GROUSE 445
M. Extensor Brevis Digiti IV, Figs. 19E, 20N
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Slender and tapering; on lateral
part of anterior surface of tarsometatarsus; length of belly variable; middle
of medial edge in contact with M. extensor hallucis longus.
Oricin.—The origin is fleshy from the lateral part of the anterior surface
of the tarsometatarsus, including the anterior metatarsal groove.
INSERTION.—The long slender tendon enters the anterior aperture of the
distal foramen, passes through the intertrochlear canal, emerges from the
terminal foramen, and attaches to the medial surface of the proximal end of
the first phalanx of digit IV.
INNERVATION.—The superficial peroneal branch of the peroneal nerve sends
a twig into the proximal part of the muscle.
INDIVIDUAL VARIATION.—None of significance in any of the three species
studied.
M. Lumbricalis, Fig. 19F
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Small, thin, and _ strap-shaped;
on mid-posterior surface of distal end of tarsometatarsus deep to tendon of
M. flexor digitorum longus; belly partly fleshy and partly elastic connective
tissue,
Oricin.—The muscle arises from the deep (anterior) surface of the tendon
of M. flexor digitorum longus a short distance proximal to the trifurcation of
the latter.
INSERTION.—The muscle attaches to the proximal end of the subarticular
cartilage ventral to the trochlea for digit III.
INNERVATION.—A long but extremely small twig arises from the paraper-
oneal branch of the tibial nerve a short distance distal to the hypotarsus and
extends distally along the mid-posterior surface of the tarsometatarsus (parallel
to a Jarger nonmuscular branch) and enters the deep surface distal to the
middle. It was possible to follow this twig in only two legs; it was pre-
sumably destroyed in the course of dissection in the others,
INDIVIDUAL VARIATION.—In some cases, the “muscle” appears grossly to be
entirely connective tissue, although a distinct entity.
T. cupido
INDIVIDUAL VARIATION.—In some cases, the “muscle” appears grossly to be
entirely connective tissue. The innervation was found in only one leg, in
which the twig arises more distally than in T, pallidicinctus,
P. p. jamesi
The innervation was not found.
446 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
M. Abductor DigitiIV, Fig. 19F
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Slender and elongate; on posterior
surface of tarsometatarsus lateral to midline; in contact with M. flexor hallucis
brevis in midline.
Oricin.—The origin is fleshy from the posterior surface of the tarsometa-
tarsus lateral to the midline beginning near the proximal end (lateral to the
hypotarsus) and ending at the level of the first metatarsal.
INsERTION.—The slender tendon, which begins along the lateral edge of the
distal part of the belly, passes through a retinaculum on the posterolateral
surface of the tarsometatarsus immediately above the outer trochlea and
attaches to the lateral surface of the proximal end of the first phalanx of
digit IV.
INNERVATION.—The paraperoneal branch of the tibial nerve gives one or
two twigs to the proximal part of the muscle.
INDIVIDUAL VARIATION.—None of significance in any of the three species
studied.
M. Flexor Hallucis Brevis, Fig. 19F
T. pallidicinctus
GENERAL DESCRIPTION AND RELATIONS.—Slender and elongate; on posterior
surface of tarsometatarsus medial to midline; belly (except proximal end)
adjacent (lateral) to posterior metatarsal crest; proximal end passing under
latter (immediately distal to hypotarsus) and lying anteromedial to hypotarsus.
Oricin.—The origin is fleshy from the medial metatarsal depression and
from the posterior surface of the tarsometatarsus between the midline and
the posterior metatarsal crest beginning immediately below the hypotarsus
and ending a short distance above the first metatarsal (sometimes more
proximally ).
INsERTION.—The slender tendon, which begins along the medial edge of
the distal part of the belly, passes through the groove on the posterodistal
surface of the first metatarsal and onto the proximal end of the ventral surface
of the hallux; the tendon widens considerably and attaches by its edges
to the ventral surface of the proximal end of the first phalanx, forming a
short “tunnel” through which the tendon of M. flexor hallucis longus passes.
INNERVATION.—The paraperoneal branch of the tibial nerve sends one or
two twigs into the proximal part of the muscle (but distal to the hypotarsus).
INprIvipuAL _VARIATION.—In two legs, the muscle arises in part from the
distal end of the lateral calcaneal ridge. The individual variation is insignifi-
cant in T. cupido and P. p. jamesi.
DISCUSSION AND CONCLUSIONS
Analysis of Individual Variation
Considerable individual variation occurs in both the muscles
and the nerves of the leg of the three species studied. The amount
of variation reported by a worker depends in large part on the
degree of variation that he considers significant.
MuscLes AND NERVES OF LEG OF GROUSE 447
Individual variation in the muscles and in the nerves will be dis-
cussed separately; that of the muscles (excluding innervation)
will be considered first.
Muscles
Considering the number, rather than degree, of variations, the
most variable muscles are: Mm. flexor digitorum longus, obturator,
caudofemoralis, and extensor hallucis longus. The first-mentioned
muscle exhibits 14 different variations in the specimens studied.
Mm. vastus lateralis, flexor perforans et perforatus digiti II, and
piriformis also showed a considerable number of variations. The
following muscles did not exhibit any variations considered signifi-
cant in this study: Mm. vastus medialis, femoritibialis internus,
flexor perforatus digiti III, extensor brevis digiti II], and abductor
digiti IV.
Muscles showing a great degree of individual variation included
the following: M. extensor proprius digiti II] was present in two
legs of Pedioecetes but absent in the other legs studied. A fleshy
muscle slip connected M. caudofemoralis pars caudifemoralis with
the tendinous raphe between Mm. flexor cruris lateralis and femoro-
cruralis in two legs, whereas in others this connection was tendinous
or even absent altogether. M. caudofemoralis pars caudifemoralis
had a tendinous area within the belly in only three legs. A vin-
culum connected the insertional tendons of Mm. flexor perforans et
perforatus digiti II and flexor perforatus digiti II in only one leg.
The fleshy belly of M. iliotrochantericus medius was completely
split into two parts in one leg. M. flexor cruris lateralis had an
accessory slip arising from the caudal musculature in one leg.
Certain individual variations reported in the accounts of the
muscles formed a graduated series, as far as degree is concerned,
from the typical to the extreme condition. Therefore it was difficult
or impossible in some cases to state whether or not certain specimens
exhibited such a variation. Elimination of the doubtful instances of
variation leaves a total of 50 different variations (excluding varia-
tions between species ) which can be attributed to a definite number
of specimens. The remainder of the discussion of individual varia-
tion in the muscles concerns these 50 variations. See table 3.
The typical condition of any structure is considered to be the con-
dition of that structure in the majority of the legs studied. Some
conditions considered as typical in the present study might not be
so considered if a larger number of specimens had been studied. If
exactly half of the legs of one species shows a particular condition
448 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
of a structure, the condition typical for this species is considered
(for purposes of the following discussion) to be that found in the
majority of the legs of the other species.
In all instances except two (of 50) the typical condition of the
muscles in T. pallidicinctus was also the typical condition in T.
cupido. The majority of the legs in T. cupido had an additional
dorsal slip on the tendon of M. flexor digitorum longus in digits II
and III. In all instances except seven the typical condition in T.
pallidicinctus was also the typical condition in Pedioecetes. In
these seven instances a variation in the former was the typical con-
dition in the latter. These were: an additional dorsal slip on the
tendon of M. flexor digitorum longus in each of three digits, a
vinculum between the latter and M. flexor perforatus digiti IV, a
partly fleshy insertion of M. flexor cruris medialis, an unossified
lateral branch of the insertional tendon of M. extensor digitorum
longus, and an independent insertion of the distalmost fibers of the
distal head of M. extensor hallucis longus. For all characters except
the number of the dorsal slips on the tendon of M. flexor digitorum
longus in digits II and III, the typical condition in T. pallidicinctus
was also the typical condition for all species considered together.
To facilitate comparison, in the following discussion all of the above-
mentioned characters are considered in all species as variants from
the typical condition.
Certain legs showed a greater number of variations from the
typical condition than did others. The majority of legs showed
from four to seven variations in the muscles of the leg. The ex-
tremes were P.p. 1L, which showed 11, and T.c.p. 2L, which
exhibited only one variation.
Twenty-three of the 50 variations were found in only one leg
(out of 23). It would be expected that if additional specimens
were studied, more kinds of variations would be found. Nine varia-
tions were found in only two legs, five in three legs, five in four
legs, and four in five legs. One variation was found in nine legs,
one in ten legs, and two in 12 legs; the last four variations were in
the number of dorsal slips of the insertional tendon of M. flexor
digitorum longus in digits IJ, III, and IV and in the ossification of
the insertional tendon of M. extensor digitorum longus.
Five of the variations were found only in specimens in which
only one leg was dissected. Considering only those eight speci-
mens in which both legs were dissected, five of the 45 variations
were found in both legs of each specimen exhibiting the variation;
Muscies AND NERVES OF LEG OF GROUSE 449
28 variations were found in only one leg of each specimen exhibiting
the variation; 12 variations were found in both legs of some speci-
mens but in only one leg of other specimens. Of the six muscle
features showing the greatest degree of individual variation (de-
scribed previously), only two (both pertaining to M. caudofemo-
ralis) were found in both legs of the specimens exhibiting the varia-
tion.
For one leg (the one showing the most variations) of each speci-
men of which both legs were studied, the number of variations that
this leg had in common with every other leg (of all species) was
determined. Then the number of variations in common between
the two legs of one individual was compared with the number of
variations in common between one leg of this individual and each
leg of every other individual. See table 4. One leg of six of the
eight specimens showed at least as many variations in common with
a leg of another individual as with the other leg of the same in-
dividual. The two exceptions were T.p. 2R and T.c.a. 1R. Thus
for most specimens there was as much variation in the muscles be-
tween the right and left legs of one individual as there was between
individuals.
Of the 50 muscle variations seven were found only in T. palli-
dicinctus (eight legs), 16 were found only in T. cupido (nine legs),
and ten were found only in Pedioecetes (six legs). Two were
found in both species of Tympanuchus (but not in Pedioecetes).
Fifteen were found in both Tympanuchus and Pedioecetes; of these,
five were found in all three species studied, eight were shared by
T. pallidicinctus and Pedioecetes, and two occurred in T. cupido
and Pedioecetes.
Nerves
The lumbosacral plexus, femoral nerve, sciatic nerve, and tibial
nerve all showed numerous individual variations. The peroneal
nerve, however, was relatively constant. Variations in the obtura-
tor nerve were considered to be insignificant. See table 5.
In all instances except one (of 40) the typical condition in T.
pallidicinctus was also the typical condition in T, cupido. In most
of the legs of the latter the nerve to M. flexor cruris lateralis did
not perforate M. caudofemoralis. In all instances except four the
typical condition in T. pallidicinctus was also the typical condition
in Pedioecetes. These exceptions were: prefixation of the lum-
bosacral plexus, six roots of the sciatic nerve, femoral nerve formed
mainly from S2 to S4 and two twigs to M. flexor ischiofemoralis.
450 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
In all instances the typical condition in T. pallidicinctus was also
the typical condition for all species considered together.
Certain legs showed a greater number of variations from the
typical condition of the nerves than did others. The greatest num-
ber of variations was shown by P.p. 3L, which had 12. T.p. 1R
and T.c.p. 1L both showed only one.
All six variations in the lumbosacral plexus were found on both
sides of each specimen exhibiting the variation. In marked con-
trast to the other nerves, there was no significant variation in the
lumbosacral plexus between the right and left sides of one in-
dividual. (This might not always be true, however, if a larger
number of specimens were studied.) Of the variations in the
lumbosacral plexus, one was found in only one specimen (of 15),
one was found in three specimens, one in four specimens, two in
six specimens, and one in seven specimens. Of the 34 variations
found in the other nerves, 14 were found in only one leg (of 23),
six occurred in two legs, four in three legs, three in four legs, three
in five legs, two in six legs, one in seven legs, and one in nine legs.
Four of the variations were found only in specimens in which
only one leg was dissected. Considering only those eight specimens
in which both legs were dissected, and excluding the lumbosacral
plexus, ten of the 30 variations were found in both legs of each
specimen exhibiting the variation; 16 variations were found in only
one leg of each specimen exhibiting the variation; four variations
were found in both legs of some specimens but in only one leg of
other specimens.
The number of variations in common between the two legs of
one individual was compared with the number between individuals
in the same manner as for the muscles; the lumbosacral plexus
was excluded from consideration. See table 6. One leg of six
of the eight specimens showed at least as many variations in
common with a leg of another individual as with the other leg of
the same individual. The two exceptions were T.p. 2L and T.p.
8R. Thus for most specimens there was as much variation in the
nerves other than the lumbosacral plexus between the right and
left legs of one individual as there was between individuals.
Of the 40 nerve variations (including the lumbosacral plexus)
11 were found only in T. pallidicinctus, seven were found only
in T. cupido, and seven were found only in Pedioecetes. Four were
found in both species of Tympanuchus (but not in Pedioecetes).
Eleven were found in both Tympanuchus and Pedioecetes; of
MuscLes AND NERVES OF LEG OF GROUSE 451
these, four were found in all three species, three were shared by
T. pallidicinctus and Pedioecetes. and four occurred in T. cupido
and Pedioecetes.
The average number of variations per leg in both muscles and
nerves was 11 in T. pallidicinctus, nine in T. cupido, and 16 in
Pedioecetes. The high number in the last is in part the result of
these being variations from the typical condition of T. pallidicinctus
(rather than from Pedioecetes).
Analysis of Variation Between Species
No constant differences in the muscles or nerves was found be-
tween T. cupido pinnatus and T. cupido attwateri. Only one con-
stant difference was found between T. cupido and T. pallidicinctus:
a thicker fleshy origin of M. extensor iliotibialis lateralis in T. cupido
(associated with a thicker edge of the lateral iliac process).
Although no constant differences in the nerves were found be-
tween Pedioecetes and Tympanuchus (both species), 17 constant
differences in the muscles were found between these two genera.
Seven of these differences pertain to features of a single muscle—
M. flexor cruris medialis. Compared with the condition in Tym-
panuchus, M. flexor cruris medialis in Pedioecetes has a wider
origin, a partly fleshy (instead of entirely tendinous) origin, a more
pronounced curvature of the line of origin, a wider insertion, an
insertion posterior (rather than anterior) to the medial collateral
ligament, an insertion that attaches in part to the articular capsule,
and a shorter tendon of insertion (resulting in the fusion of the
common insertional tendon of Mm. flexor cruris lateralis and
femorocruralis with the fleshy belly rather than with the insertional
tendon). Other differences include the following. A more ex-
tensive posteroproximal aponeurosis of M. extensor iliotibialis
lateralis in Pedioecetes (resulting in a narrower fleshy origin); a
more nearly straight line of origin of this muscle (associated with
a less pronounced lateral iliac process); a thinner fleshy origin of
this muscle (associated with a thinner edge of the lateral iliac
process); a wider M. flexor cruris lateralis that is fleshy up to the
origin from the vertebrae; a wider fleshy origin of M. iliacus; the
origin of M. caudofemoralis pars iliofemoralis not reaching the
ventral edge of the ischium; a narrower origin of M. adductor super-
ficialis; a wider M. femorocruralis; and a shorter belly of M. ex-
tensor digitorum longus. Some additional differences between
these two genera, which are slight in degree, are given in the ac-
452 University oF Kansas Pusts., Mus. Nat. Hist.
counts of the muscles. If additional specimens were studied, some
of the differences listed above possibly would prove to be subject to
individual variation and so could not properly be listed as constant
differences between the two genera.
The picture of the differences between Tympanuchus and
Pedioecetes that the present study presents is radically different
from that presented by the study of Hudson, e¢ al. (1959). These
authors reported the following differences between these two
genera. (I am using my terminology.) The origin of M. piriformis
is narrower in Pedioecetes and is more posteriorly situated; the
belly of M. extensor iliotibialis anticus is broader in Pedioecetes;
the belly of M. tibialis anticus is longer; the belly of M. peroneus
brevis is shorter; the insertional tendon of the anterolateral head
of M. flexor perforatus digiti III is shorter; the belly of M. flexor
digitorum longus is shorter; only two (rather than three) of the
branches of M. extensor digitorum longus on the tarsometatarsus
are ossified; the posterior metatarsal crest is shorter; M. flexor per-
forans et perforatus digiti II has two heads in Pedioecetes but only
one in Tympanuchus; the roof over the hypotarsal canal enclosing
the tendon of M. flexor digitorum longus is bony in Pedioecetes but
fibrous in Tympanuchus; M. flexor cruris lateralis is wider in Pedio-
ecetes; and the origin of M. femorocruralis is wider. I paid par-
ticular attention in my study to these 13 features given by Hudson,
et al.; of these the only differences that I found to be constant were
the last two. The apparent reason for this great discrepancy is the
small number of legs of Tympanuchus studied by Hudson, et al.
They studied eight legs of Pedioecetes but only two legs of Tym-
panuchus. This emphasizes the danger of making comparisons
based on a very small number of specimens (a criticism which may
prove to apply to the present study as well). The reason why
Hudson, et al. did not report most of the differences found by me
is not so apparent. Either the specimens studied by the former
workers showed a greater variation in these characters than did
my specimens or else those workers overlooked the differences.
Probably both factors are involved. It remains to be determined
how many specimens need to be studied in order to obtain a fairly
accurate picture of variation.
Comparison with Other Studies of Innervation
I accept the following concept of muscle-nerve relationship. All
muscles of the pelvic limb of birds have developed phylogenetically
MuscLES AND NERVES OF LEG OF GROUSE 453
from either the dorsal extensor muscle mass or the ventral flexor
muscle mass. The former was (at least originally) supplied by
only the femoral and peroneal nerves (“dorsal” nerves), the latter
by only the obturator and tibial nerves (“ventral” nerves), The
best guide for determining which muscles are phylogenetically
dorsal and which are ventral seems to be their embryogeny (as
shown in the studies of Romer, 1927, and Wortham, 1948). In
the phylogenetic changes undergone by the muscles under con-
sideration, the innervation may have changed in some instances,
although this is less likely to occur than changes in the attachment
or function of the muscles, If a change in innervation has oc-
curred, it would be more likely to be a change from one dorsal
nerve to the other or from one ventral nerve to the other rather
than from a dorsal nerve to a ventral one or vice versa.
Thus, in my opinion, a report of a dorsal muscle supplied by a
ventral nerve, or vice versa, should be viewed with suspicion until
it is verified. I suspect that many previous workers have ignored
this concept of muscle-nerve relationship, or else do not accept
it, since they report, without comment, dorsal muscles (as deter-
mined embryologically ) innervated by ventral nerves, or vice versa.
Owing to the intimate association between the proximal parts of
the tibial and peroneal nerves, the true relationship may be difficult
to determine. I suspect that this relationship has been misinter-
preted by a number of workers. I found in Tympanuchus and
Pedioecetes a branch of the tibial nerve that is closely associated
with, and distributed with, the peroneal nerve and has been mis-
takenly considered a part of the peroneal nerve by some workers.
In the study here reported on, I have found no definite exceptions
to the expected innervation. The only possible exception is an
extra branch, which could not be traced to its origin, supplying
M. extensor iliofibularis in one leg. Thus my study of innervation
agrees with the embryological determination of the (phylogenetic)
dorsal and ventral muscles and lends strong support to the above-
stated concept of muscle-nerve relationship.
I have compared my findings on the nerves with those of other
workers, who have studied the nerves with a varying degree of
thoroughness, The important differences in innervation between
these studies and the present one are discussed below.
In neither of Gadow’s works did he distinguish tibial and peroneal
components in the thigh. In his later work (1891), covering a
wide variety of birds, he found that M. piriformis sometimes has
454 UNIVERSITY OF Kansas PusBts., Mus. Nar. Hist.
a femoral innervation in addition to the constant sciatic one and
that M. gluteus profundus may or may not have a sciatic supply in
addition to the femoral one. A comparison of Gadow’s terminology
of the sciatic nerve branches in the shank and foot (in both works)
with mine shows that his branch I represents my peroneal nerve
plus my paraperoneal branch of the tibial nerve (Ic); his branch II
represents my medial division of the tibial nerve; and his branch III
represents my posterior (IIIa) and lateral (IIJb) divisions of the
tibial nerve.
Gadow’s study (1880) on the ratites included Struthio, Rhea, and
Casuarius. Only in Casuarius did Gadow find a branch (Ile) of
the sciatic nerve supplying Mm. lumbricalis, adductor digiti II, and
abductor digiti II. The two former muscles are typically supplied
(as in Rhea) by the paraperoneal branch of the tibial nerve;
Gadow’s branch Ile presumably represents a segregated branch of
this nerve. More surprising is his finding that M. abductor digiti II
is innervated in Casuarius by both the deep peroneal nerve and
branch Ile and in Rhea by branch Ic (paraperoneal branch of tibial
nerve). The deep peroneal innervation is typical. Also unexpected
is his finding that the posterior division of the femoral nerve gives
minute twigs into M. gastrocnemius pars interna in Struthio and
Casuarius. Since the other terminal branches of this nerve in these
birds are nonmuscular, since this muscle is chiefly supplied by other
nerves, and since the innervation from the femoral nerve is ap-
parently atypical for most birds, the possibility should be considered
that the femoral twigs are sensory rather than motor.
Sudilovskaya (1931), studying Struthio, Rhea, and Dromaeus
(Dromiceius), used the same terminology as Gadow except that he
designates as branch III Gadow’s branch Ic. Sudilovskaya’s dis-
cussion of the main branches of the sciatic nerve is confusing. He
states that in Struthio, branches I, IJ, and III all pass through the
tendinous guide loop for M. extensor iliofibularis; this is hard to be-
lieve. As near as I can determine, he has mistakenly given the same
designation (branch III) to two separate branches (Gadow’s Ic
and III). There is no problem, however, in determining to which
of these two branches he is referring when he is describing the in-
nervation of a particular muscle, since one supplies only muscles
of the shank and the other only intrinsic foot muscles. Sudilovskaya
found M. abductor digiti II to be innervated by branch III (Ic of
Gadow); thus the innervation of this muscle corresponds to that
found in Rhea by Gadow. Although M. adductor digiti II had the
expected innervation from branch III (paraperoneal branch of
Muscles AND NERVES OF LEG OF GROUSE 455
tibial nerve) in Dromaeus, that muscle was found to be supplied by
branch II in Rhea. (Gadow, on the other hand, reports a typical
innervation for this muscle in Rhea.) Sudilovskaya found M.
peroneus brevis to be supplied by the deep peroneal branch (in
contrast to the superficial peroneal supply that I found in Tym-
panuchus and Pedioecetes). He found M. gastrocnemius pars in-
terna to be supplied in Struthio by twigs of the femoral nerve in
addition to its typical innervation from branch II of the sciatic
nerve; this agrees with Gadow’s findings in the same genus.
Sudilovskaya reports that M. gastrocnemius pars externa was in-
nervated by branches II and II in Struthio and Rhea and by
branches I and III in Dromaeus. (Gadow found only the typical
innervation—branch III.)
In the Whooping Crane, Fisher and Goodman (1955) found a
peroneal, rather than a femoral, nerve supply for pars postica of
M. vastus lateralis. They also report a peroneal nerve supply for
M. flexor ischiofemoralis (in contrast to the usual tibial nerve sup-
ply) and for M. adductor superficialis (in addition to the usual
supply from the obturator nerve). The innervation was not given
for the intrinsic foot musculature.
Fisher (1946), studying vultures, reports the following: tibial
branches, in addition to the main sciatic branch, supplying M. ex-
tensor iliofibularis (typically supplied by the peroneal nerve); an
obturator supply, in addition to the usual tibial supply, to M. flexor
cruris medialis; a tibial supply, in addition to the typical ob-
turator supply, to M. obturator pars postica; a possible obturator
supply, in addition to the typical femoral supply, to M. ambiens;
a possible peroneal supply, in addition to the typical tibial supply,
to M. flexor digitorum longus; and a peroneal supply to Mm. ab-
ductor digiti IV, flexor hallucis brevis, and adductor digiti II (which
are typically supplied by the paraperoneal branch of the tibial
nerve). Fisher’s postfibular branch of the peroneal nerve, which
supplies the latter three muscles, apparently represents the para-
peroneal branch of the tibial nerve.
Carlsson (1884) did not find a femoral nerve supply for M.
gluteus profundus. He found an obturator supply, in addition to
the usual sciatic supply, to M. flexor ischiofemoralis in Eudyptes
chrysolopha and Mergulus alle but not in the other two forms
studied. He reported a peroneal supply, rather than the expected
tibial (paraperoneal) supply, to Mm. abductor digiti IV and ad-
ductor digiti IV.
DeMan (1873) found a twig of the obturator nerve supplying
456 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
M. flexor ischiofemoralis, in addition to the typical innervation, in
Corvus monedula, but not in the few other forms studied. He
did not distinguish tibial and peroneal components in the thigh.
Wilcox (1948), studying a loon, did not find any peroneal supply
to M. extensor iliotibialis lateralis or to M. gluteus profundus. He
found a femoral, rather than a peroneal, supply to M. piriformis.
He found an obturator, instead of a tibial, supply to M. flexor ischio-
femoralis. (In some of my specimens I found a tiny blood vessel,
appearing much like a nerve, emerging from the obturator foramen
and entering M. flexor ischiofemoralis.) Wilcox reports an innerva-
tion of M. caudofemoralis pars caudifemoralis from the pudendal
plexus, in addition to the usual sciatic one. Wilcox did not dis-
tinguish tibial and peroneal components in the thigh. In the shank
and foot he misidentified the peroneal nerve as the tibial nerve
and therefore gave erroneous innervations for all the muscles sup-
plied by this nerve, except for M. adductor digiti IV, which actually
should be supplied by the tibial nerve.
Howell (1938) studied only the hip and thigh musculature of
the chicken. He overlooked the femoral nerve supply for M.
gluteus profundus.
Romer (1927) studied only the hip and thigh muscles of the
chick. He did not distinguish tibial and peroneal components in
the thigh. He did not mention any sciatic supply for M. gluteus
profundus,
Appleton (1928), studied (in various birds) only those muscles
of the hip and thigh that are innervated by the tibial and peroneal
nerves. He terms the former “ischiadicus ventralis” and the latter
“ischiadicus dorsalis.” His findings did not differ from mine.
Many differences in the innervation of specific muscles are re-
ported in the literature, even in the same species (by different
workers). Some of these differences may be real; others are prob-
ably misinterpretations. Consequently more work needs to be
done before a complete understanding can be obtained of the
innervation of the leg muscles of birds. Especially needed are
studies of the tibial-peroneal nerve relationship, perhaps approached
by a method other than gross dissection.
MuSCLES AND NERVES OF LEG OF GROUSE 457
SUMMARY
The muscles and nerves were dissected in eight legs of the Lesser
Prairie Chicken (Tympanuchus pallidicinctus), six legs of the
Greater Prairie Chicken (T. cupido pinnatus), three legs of Att-
water's Prairie Chicken (T. c. attwateri), and six legs of the Sharp-
tailed Grouse (Pedioecetes phasianellus jamesi) for the purpose of
obtaining information on individual variation as well as variation
between these closely related species. Relatively little information
is available regarding the nerves of the leg of birds and little is
known about individual variation and variation between closely re-
lated forms in the muscles of the leg of birds.
All osteological terms used in the present paper are defined and
those of the pelvis are illustrated. New terms were coined for some
structures for which no names could be found in the literature.
Terms were also coined for the major divisions of the femoral and
sciatic nerves. With three exceptions, my muscle terminology fol-
lows that of Fisher (1946) and Fisher and Goodman (1955). Their
term femoritibialis externus is not used here; the muscle so named
is considered to be a part of M. vastus lateralis. Fisher’s accessory
head of M. flexor cruris lateralis is considered to be a distinct muscle
—M. femorocruralis. Usage of the term obturator internus is
avoided because the muscle so named is considered not to be
homologous with the mammalian muscle of the same name; the
entire obturator complex is called M. obturator, and is subdivided
into four parts.
The typical (most common) condition of the nerves and muscles
in Tympanuchus pallidicinctus is described in detail. Variations
from this condition among the other birds studied are then de-
scribed. All muscles of one leg of T. pallidicinctus are illustrated.
Several variations in the muscles are also illustrated. The lum-
bosacral plexus and nerves of the leg in several specimens that
show variations are illustrated.
Considerable individual variation was found in both the muscles
and the nerves of the leg of the species studied. Certain muscles
were more variable than others. Mm. flexor digitorum longus,
obturator, caudofemoralis, and extensor hallucis longus showed the
greatest number of variations. Mm. vastus medialis, femoritibialis
internus, flexor perforatus digiti III, extensor brevis digiti III, and
abductor digiti IV did not exhibit any variations considered sig-
nificant. Certain legs showed a greater number of variations from
the typical condition than did others.
458 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
Although most of the variations were minor, some were major.
M. extensor proprius digiti III was present in two legs of Pedioecetes
but absent in the other legs studied. A fleshy muscle slip connected
M. caudofemoralis pars caudifemoralis with the tendinous raphe
between Mm. flexor cruris lateralis and femorocruralis in two legs,
whereas in others this connection was tendinous or even absent
altogether. M. flexor cruris lateralis had an accessory slip arising
from the caudal musculature in one leg. A vinculum connected the
insertional tendons of Mm. flexor perforans et perforatus digiti II
and flexor perforatus digiti II in one leg.
In most specimens there was as much variation between the
muscles of the right and left legs of one individual as there was be-
tween individuals. The same was true for the nerves, except for
the lumbosacral plexus, in which there was no significant variation
between the right and left sides of any individual. The peroneal
and obturator nerves varied less than the other nerves.
No constant differences in the muscles or nerves was found be-
tween T. cupido pinnatus and T. c. attwateri. One constant differ-
ence was found between T. cupido and T. pallidicinctus: the fleshy
origin of M. extensor iliotibialis lateralis in T. cupido was thicker
(associated with a thicker edge of the lateral iliac process ).
Although no constant differences in the nerves were found be-
tween Pedioecetes and Tympanuchus (both species), 17 constant
differences in the muscles were found between these two genera.
Study of additional specimens possibly would show enough indi-
vidual variation in some of these differences to reduce the number
of constant differences to fewer than 17. Seven of these differences
pertain to features of a single muscle—M. flexor cruris medialis.
Some of the other differences are associated with the thinner and
much less pronounced lateral iliac process in Pedioecetes. The
picture of the differences between Tympanuchus and Pedioecetes
that this study presents is radically different from that presented by
the study of Hudson, et al. (1959).
The important differences in innervation between previous
studies and the present one are discussed.
All of the muscles under consideration have been grouped as
either dorsal or ventral muscles, according to their embryonic
origin, as described by Romer (1927) and Wortham (1948). This
grouping probably represents accurately the phylogenetic origin
of these muscles. The dorsal muscles probably were originally sup-
plied by dorsal nerves—the femoral and peroneal—and the ventral
Musc.Les AND NERVES OF LEG OF GROUSE 459
muscles probably were originally supplied by ventral nerves—the
obturator and tibial. This primitive muscle-nerve relationship has
been relatively constant.
Several previous workers have reported some dorsal muscles sup-
plied by ventral nerves and vice versa. Those findings should be
viewed with suspicion until verified, because the proximal parts
of the tibial and peroneal nerves are intimately associated and
their relationship is easily misinterpreted. I found a branch of
the tibial nerve that is closely associated with, and distributed with,
the peroneal nerve. That branch of the tibial nerve has been mis-
takenly considered a part of the peroneal nerve by some workers.
My study revealed no definite exceptions to the expected innerva-
tion.
7—5835
UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
460
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MusSCLES AND NERVES OF LEG OF GROUSE
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UnIveRSITY OF Kansas Pusts., Mus. Nat. Hist.
462
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463
MuscLEs AND NERVES OF LEG OF GROUSE
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464 UNIversITy OF Kansas Pusts., Mus. Nat. Hist.
TABLE 2. RELATIVE Sizes (IN PERCENTAGES) OF SOME MUSCLES IN
TYMPANUCHUS AND PEDIOECETES
Muscle
Tliacus: width of fleshy origin
(divided by length of ilium).....
Flexor cruris lateralis: maximum
width of exposed part (divided
by length ofvilium)).....5.2:..5.-:%
Flexor cruris medialis: width of ori-
gin (divided by length of ilium). .
Flexor cruris medialis: width of in-
sertion (divided by length of
LIDIOCATSUS) hae stothe ee se ee
Adductor superficialis: width of ori-
gin (divided by length of ilium). .
Femorocruralis: distance of proxi-
mal end of origin from proximal
end of femur (divided by length
OifeMUM) has. ae Aas che tees
Extensor digitorum longus: length
of fleshy belly (divided by length
Ol tiblotarsus)mi acer dec ese oe. &
Tympanuchus
Ave.| Range | No.1} Ave.
10 | .08-.11 | 13 | .19
-22 } 19-27 | 13 1 .31
11 | .08-.16 | 18 |} .22
.09 | .08-.13 |} 13 | .17
20" 1F— 23) | ts) fs
.59 | .55-.63 | 13 | .40
.43 | 64-83 | 138 | .59
Pedioecetes
Range |No.!
17-.19 | 6
27-.36 | 6
19-.23 | 6
15-.17 | 4
10-.16 | 5
.38-.43 | 6
50-.62 | 4
1. No. = number of legs.
465
Muscles AND NERVES OF LEG OF GROUSE
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UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
466
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Muscles AND NERVES OF LEG OF GROUSE
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UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist.
468
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469
MuscLES AND NERVES OF LEG OF GROUSE
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UNIvERsITY OF Kansas Pusts., Mus. Nat. Hist.
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MuscLes AND NERVES OF LEG OF GROUSE 473
LITERATURE CITED
APPLETON, A. B.
1928. The muscles and nerves of the post-axial region of the tetrapod
thigh. PartsI and II. Jour. Anat., 62(3,4):364-438.
BERGER, A. J.
1952. The comparative functional morphology of the pelvic appendage
in three genera of Cuculidae. Amer. Midl. Nat., 47(3):513-605.
Bercer, A. J.
1956. The appendicular myology of the Sandhill Crane, with comparative
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IBOAS Jue, Ve
1933. Kreuzbein, Becken und Plexus Lumbosacralis der Végel. Det
Kongelige Danske Videnskabernes Selskabs Skrifter. Naturviden-
skabelig og Mathematisk Afdeling. 9 raekke, 5(1):1-74, 15 pls.
CaRLsson, A.
1884. Beitrige zur Kenntniss der Anatomie der Schwimmvégel. Bihang
till K. Svenska Vetenskapsakad. MHandlingar, 9(3):1-44, 5 pls.
Cuomiak, M.
1950. [Studies on the plexus lumbalis et sacralis in the domestic hen.]
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5(8):29-45,
FisHer, H. I.
1946. Adaptations and comparative anatomy of the locomotor apparatus
of new world vultures. Amer, Midl. Nat., 35(3):545-727, 18 pls.
Fisver, H. I., and GoopMan, D. C.
1955. The myology of the Whooping Crane, Grus americana. IIl. Biol.
Mono., 24(2):viii + 1-127.
Gapow, H.
1880. Zur vergleichenden Anatomie der Muskulatur des Beckens und der
hinteren Gliedmasse der Ratiten. Fischer, Jena, 56 pp., 5 pls.
Gapow, H. (with E. Selenka).
1891. Vo6gel. I. Anatomischer Theil. In Bronn’s Klassen und Ordnungen
des Their-Reichs, 6(4):1-1008. Winter, Leipzig.
Hotes, E. B.
1962. The terminology of the short extensor muscles of the third toe in
birds. Auk, 79(3):485-488.
Howarp, H.
1929. The avifauna of Emeryville shellmound. Univ. Calif. Publ. Zool.,
82(2):301-394, 4 pls.
HowEL.t, A. B.
1938. Muscles of the avian hip and thigh. Auk, 55(1):71-81.
Hupson, G. E.
1987. Studies on the muscles of the pelvic appendage in birds. Amer.
Midl. Nat., 18(1):1-108, incl. 26 pls.
Hupson, G. E., et al.
1959. Muscles of the pelvic limb in galliform birds. Amer. Midl. Nat.,
61(1):1-67.
JHERING (IHERING), H. V.
1878. Das peripherische Nervensystem der Wirbelthiere. Vogel, Leipzig,
xiv-238 pp., 5 pls.
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1873. Vergelijkende myologische en neurologische Studien over Amphib-
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RoMER, A. S.
1927. The development of the thigh musculature of the chick. Jour.
Morph., 43(2):347-385.
474 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
SupiLovskayA, A. M.
1931. [Study on the comparative anatomy of the musculature and in-
nervation of the pelvic region and the hind appendages of the
Ratitae (Struthio, Rhea, Dromaeus).] Acad. Sci. U.S.S.R., Len-
ingrad, 84 pp. (In Russian.)
Lom. EP; J; pu
1913. Untersuchungen iiber das Synsacrum und den Schwanz von Gallus
domesticus nebst Beobachtungen tuber Schwanzlosigkeit bei Kaul-
hithnern. Jenaische Zeitschr. Naturw., 49:149-312, 8 pls.
Wiicox, H. H., Jr.
1948. The pelvic musculature of the loon (Gavia immer). Univ. Micro-
films, Ann Arbor, 95 pp., 26 pls.
WorrTHAM, R. A.
1948. The development of the muscles and tendons in the lower leg and
foot of chick embryos. Jour. Morph., 83(1):105-148.
YasupA, M., et al.
1959. [Comparative and topographical anatomy of the fowl. XI. On the
nervous supply of the hind limb.] In Proc. of 47th Meeting of Jap.
Soc. 7 Vet. Sci. Jap. Jour. Vet. Sci., 21(6):386. (Japanese ab-
stract.
Transmitted October 30, 1962.
XO
29-5835
Vol. 10.
Vol. 12. 1.
Vol. 18.
18.
19.
20.
21,
22.
23.
(Continued from inside of front cover)
Conspecificity of two pocket mice, Perognathus goldmani and P. artus. By
. Raymond Hall and Marilyn Bailey Ogilvie. Pp. 513-518, 1 map. Janu-
ary 14, 1960.
Records of harvest mice, Reithrodontomys, from Central America, with de-
scription of a new subspecies from Nicaragua. By Sydney Anderson and
J. Knox Jones, Jr. Pp..519-529. January 14, 1960.
Small carnivores from San Josecito Cave (Pleistocene), Nuevo Leén, México,
By E. Raymond Hall. Pp, 531-538, 1 figure in text. January 14, 1960.
Pleistocene pocket gophers from San Josecito Cave, Nuevo Leén, México.
By Robert J. Russell. Pp. 589-548, 1 figure in text. January 14, 1960.
Review ofthe insectivores of Korea. By J. Knox Jones, Jr., and David H.
Johnson. Pp. 549-578. February 23, 1960.
Speciation and evolution of the pygmy mice, genus Baimoys. By Robert L.
Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960.
Index. Pp. 671-690
Ms
2.
3.
4,
5.
9.
10.
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.
Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.
The forest habitat of the University of Kansas Natural History Reservation.
By Henry §. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures
in text, 4 tables. December $31, 1956.
Aspects of reproduction and development in the prdirie vole (Microtus ochro-
gaster). By Henry S. Fitch. Pp. 129-161, 8 figures in text, 4 tables. Decem-
ber 19, 1957.
Birds found on the Arctic slope of northern Alaska. By James W. Bee.
Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958.
The wood rats of Colorado: distribution and ecology. By Robert B. Finley,
Jr. Pp. 213-552, 34 plates, 8 figures in text, 85 tables. November 7, 1958.
Home ranges and movements of the eastern cottontail in Kansas. By Donald
W.. Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, 1959.
Natural history of the salamander, Aneides hardyi. By Richard F. Johnston
and Gerhard A. Schad. Pp. 573-585. October 8, 1959.
A new subspecies of lizard, Cnemidophorus sacki,-from Michoacan, México.
By William E. Duellman. Pp. 587-598, 2 figures in text. May 2, 1960.
A taxonomic study of the middle American snake, Pituophis deppei. By
William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May-2, 1960.
Index. Pp. 611-626.
Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960.
*2.
Functional morphology of three bats: Sumops, Myotis, Macrotus. By Terry
A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. July 8, 1959. )
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.
The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49
figures in text. February 19, 1960.
A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By
Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in
text. May 2, 1960.
Natural history of the bell vireo. By Jon C, Barlow. Pp. 241-296, 6 figures
in text. March 7, 1962.
Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C.
Fox. Pp. 297-307, 6 figures in text. May 21, 1962.
Vertebrates from the barrier island of Tamaulipas, México. By Robert K.
Selander, Richard F. Johnston, B, J. Wilks, and Gerald G. Raun. Pp. 309-
345, pls. 5-8. June 18, 1962.
Teeth of Edestid sharks. By Theodore H. Eaton, Jr. Pp. 847-362, 10 fig-
ures in text. October 1, 1962.
Variation in the muscles and nerves of the leg in two genera of grouse
(Tympanuchus: and Pedioecetes). By E. Bruce Holmes. Pp. 363-474, 20
figs. October 25, 1963.
More numbers will appear in volume 12.
Five natural hybrid combinations in minnows (Cyprinidae). By Frank B.
Cross and W.\L. Minckley. Pp. 1-18: June 1, 1960.
A distributional study of the amphibians of the Isthmus~ of Tehuantepec,
México. By William E. Duellman. Pp. 19-72, pls. 1-8, 8 figures in text.
August 16, 1960.
A new subspecies of the slider turtle .(Pseudemys scripta) from Coahulia,
Sate By John M. Legler. Pp. 73-84, pls. 9-12, 3 figures in text. August
Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, pls. 13-20,
26 figures in text. November 30, 1960.
Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and
Rocky Mountains. By Henry S.’Fitch and T, Paul Maslin. Pp. 289-308,
4 figures in text. February 10, 1961.
Fishes of the Wakarusa river in Kansas. By James E. Deacon and Artie L.
Metcalf. Pp. 809-322, 1 figure in text. February 10, 1961.
Geographic variation in the North American cyprinid fish, Hybopsis gracilis.
By Leonard J. Olund and Frank B. Cross. Pp, 3238-348, pls. 21-24, 2 figures
in text. February 10, 1961.
(Continued on outside of back cover)
8.
9.
10.
(Continued from inside of back cover)
Decriptions of two species of frogs, genus Ptychohyla; studies of Ameri-
can hylid frogs, V.. By William E. Duellman. Pp. 349-357, pl. 25, 2 figures
in text. April 27, 1961.
Fish populations, following a drought, in the Neosho and Marais des Cygnes
rivers of Kansas. By James Everett Deacon. Pp. 359-427, pls. 26-30, 8 figs.
August 11, 1961. ;
Recent soft-shelled turtles of North America (family Trionychidae). By
a ee Webb. Pp. 429-611, pls. 31-54, 24 figures in text. February
Index. Pp. 613-624.
Vol. 14. 1.
2.
8.
Neotropical bats from. western México; By Sydney Anderson. Pp. 1-8.
October 24, 1960.
Geographic variation in the harvest mouse. Reithrodontomys megalotis, on
the central Great Plains and in adjacent regions. By J. Knox Jones, Jr.,
and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961.
Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson.
Pp. 29-67, pls. 1 and 2, 3 figures in text. July 24, 1961,
A new subspecies of the black myotis (bat) from, eastern Mexico. By E.
Faynond Hall anad Ticul Alvarez. Pp. 69-72, 1 figure in text. December
North American yellow bats, “‘Dasypterus,’” and a list of the named kinds
of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr.
Pp. 73-98, 4 figures in text. - December 29, 1961.
Natural history of the’ brush mouse (Peromyscus boylii) in Kansas with
description of a new subspecies. By Charles A. Long. Pp. 99-111, 1 figure
in text. December 29, 1961.
Taxonomic status of some mice of the Peromyscus: boylii group in eastern
Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 118-
120, 1 figure in text. December 29, 1961.
A new subspecies of ground squirrel (Spermophilus spilosoma) from Ta-
maulipas, Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962.
Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J.
” Knox Jones, Jr., and Ticul Alvarez. / Pp. 125-133, 1 figure in text. March 7,
1962.
A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene,
of Texas. By E. Raymond Hall and Walter W. Dalquest. Pp. 135-138,
2 figures in text. April 30, 1962.
A new subspecies of wood rat (Neotoma) from northeastern Mexico. By
Ticul Alvarez. Pp. 139-143. April 30, 1962.
Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul
aes and M. Raymond Lee. ~Pp. 145-159, 1 figure in text. May 18,
A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164,
1 figure in text. May 21, 1962.
The mammals of Veracruz. By E. Raymond Hall anad Walter W. Dalquest.
Pp. 165-862, 2 figures. May 20, 1963. S :
The recent mammals of Tamaulipas, México. By Ticul Alvarez. Pp. 868-
473, 5 figures in text.’ May 20, 1963.
More numbers will appear in volume 14.
The amphibians and reptiles of Michoacan, México. By William E. Duell-
man. Pp. 1-148, pls. 1-6, 11 figures in text. December 20, 1961.
Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox
Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962.
A new species of frog (Genus Tomodactylus) from western México. By
Robert G. Webb. Pp.-175-181, 1 figure in text. March 7, 1962.
Type specimens of amphibians and reptiles in the Museum of Natural His-
tory, the University of Kansas. By William E. Duellman and Barbara Berg.
Pp. 183-204. October 26, 1962.
Amphibians and Reptiles of the Rainforests of Southern El Petén, Guatemala.
By William E. Duellman. - Pp. 205-249, pls. 7-10, 6 figures in text. October
4, 1963.
A revision of snakes of the genus Conophis (Family Colubridae, from Middle
scicey By John Wellman. Pp. 251-295;.9 figures in text. October 4,
638.
A review of the Middle Américan tree frogs of the genus Ptychohyla. By
vate Duellman. Pp. 297-349, pls. 11-18, 7 figures in text. October
18, 1963.
More numbers will appear in volume 15.
UNIVERSITY OF KANSAS PUBLICATIONS mt Addai
i i ¥cROLET
MvuSsEUM OF NATURAL HISTORY ev nencrornints Soe
toe
Volume 12, No. 10, pp. 475-501, 7 figs.
October 25, 1963
A New Genus of Pennsylvanian Fish
(Crossopterygii, Coelacanthiformes)
from Kansas
BY
JOAN ECHOLS
UNIVERSITY OF KANSAS
LAWRENCE
1963
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HisToRY
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 12, No. 10, pp. 475-501, 7 figs.
Published October 25, 1963
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY
JEAN M. NEIBARGER, STATE PRINTER
TOPEKA, KANSAS
A New Genus of Pennsylvanian Fish | bal shies
(Crossopterygii, Coelacanthiformes )
from Kansas
BY
JOAN ECHOLS
INTRODUCTION
In 1931 and 1932, H. H. Lane, C. W. Hibbard and W. K. Mc-
Nown collected the specimens that Hibbard (1933) described and
made the basis of two new species. These were from the Rock
Lake shale member of the Stanton formation, six miles northwest
of Garnett, Anderson County, Kansas. In 1954, from a locality
(KAn-1/D, see page 480) approximately one fourth mile southwest
of the first locality, specimens were quarried by F. E. Peabody,
R. W. Wilson and R. Weeks. In 1955 R. R. Camp collected addi-
tional blocks of Rock Lake shale from this second locality. Study
of all of the materials from the above mentioned localities reveals
the existence of an hitherto unrecognized genus of coelacanth. It
is named and described below.
I wish to thank Prof. Theodore H. Eaton, Jr., for suggesting the
project and for much helpful advice. I am indebted to Dr. E. I.
White of the British Museum (Natural History) for furnishing a
cast of the endocranium of Rhabdoderma elegans (Newberry) for
comparison, and to Drs. Donald Baird (Princeton University ), Bobb
Schaeffer (American Museum of Natural History) and R. H. Deni-
son (Chicago Natural History Museum) for loans and exchanges
of specimens for comparison. I am grateful to Dr. Bobb Schaeffer
for advice on the manuscript. Mr. Merton C. Bowman assisted with
the illustrations. The study here reported on was made while I
was a Research Assistant supported by National Science Founda-
tion Grant G-14013.
(477)
478 UNIVERSITY OF KANsAs PuBLs., Mus. Nar. Hist.
SYSTEMATIC DESCRIPTIONS
Subclass CROSSOPTERYGII
Superorder CoELACANTHI
Order Coelacanthiformes
Suborder DIPLOCERCIDOIDEI
Family DrpLocerRcIDAE
Subfamily Rhabdodermatinae, new subfamily
Type genus.—Rhabdoderma Reis, 1888, Paleontographica, vol. 35, p. 71.
Referred genus.—Synaptotylus new, described below.
Horizon.—Carboniferous.
Diagnosis——Sphenethmoid region partly ossified, and consisting of basi-
sphenoid, parasphenoid, and ethmoid ossifications; paired basipterygoid process
and paired antotic process on basisphenoid; parasphenoid of normal size, and
closely associated with, or fused to, basisphenoid; ethmoids paired in Rhab-
doderma (unknown in Synaptotylus).
Discussion—Because of the great differences in endocranial
structure between the Devonian and Pennsylvanian coelacanths,
they are here placed in new subfamilies. The two proposed sub-
families of the family Diplocercidae are the Diplocercinae and the
Rhabdodermatinae. The Diplocercinae include those coelacanths
having two large unpaired bones in the endocranium (at present
this includes Diplocercides Stensié, Nesides Stensi6 and Euporos-
teus Jaekel). The subfamily Rhabdodermatinae is composed of
coelacanths having reduced endocranial ossification, as described in
detail above, and now including Rhabdoderma Reis and Synapto-
tylus n. g.
Members of this subfamily differ from those of the subfamily
Diplocercinae in having several paired and unpaired elements in
the sphenethmoid region of the endocranium, instead of only one
larger ossification. They differ from those of the suborder Coela-
canthoidei in the retention of basipterygoid processes.
Synaptotylus is more closely related to Rhabdoderma than to the
Diplocercines because the anterior portion of the endocranium
contains only a basisphenoid, parasphenoid, and probably ethmoids.
The sphenethmoid region was certainly not a large, unpaired unit
as in the Diplocercines. Probably the posterior part, the otico-
occipital region (not known in Synaptotylus), was much more
nearly like that of Rhabdoderma, which consisted of unpaired basi-
occipital and supraoccipital, and paired prootics, exoccipitals, and
anterior and posterior occipital ossifications (Moy-Thomas, 1937:
A New GENUS OF PENNSYLVANIAN FIsH 479
figs. 3, 4). Moy-Thomas (1937:389) points out that in Rhabdo-
derma the occipital region is “considerably more ossified” than in
any coelacanths other than the Devonian forms. Berg (1940:390)
thought that the Carboniferous coelacanths should be placed in a
separate family because they did not have two large, unpaired
bones in the endocranium. Rhabdoderma and Synaptotylus repre-
sent another stage in evolution of the endocranium in coelacanths,
and, if classification is to be based on endocranial structure, then
this stage (represented by the two genera) may later be given
family rank as Berg suggested. Because Rhabdoderma and Synap-
totylus have only part of the sphenethmoid region ossified and be-
cause they retain basipterygoid processes, they are considered to be
related and are included in the subfamily Rhabdodermatinae.
Synaptotylus, new genus
Type species.—Synaptotylus newelli (Hibbard).
Horizon.—Rock Lake shale member, Stanton formation, Lansing group,
Missouri series, Upper Pennsylvanian.
Diagnosis.—Late Pennsylvanian fishes of small size, having the
following combination of characters: on basisphenoid, knoblike
antotic processes connected by a low ridge to basipterygoid proc-
esses; entire ventral surface of parasphenoid toothed; anterior mar-
gin of parasphenoid notched and no evidence of hypophyseal
opening. Dermal bones of skull smooth or with low, rounded
tubercles and striae; fronto-ethmoid shield incompletely known
but having one pair of large rectangular frontals with postero-
laterally slanting anterior margins; intertemporals large, the lateral
margins curving laterally; postorbital triangular, apex downward;
subopercular somewhat triangular; squamosal carrying sensory
canal that curves down posteriorly and extends onto a ventral pro-
jection; opercular generally triangular; supratemporals elongate,
curving to fit lateral margin of intertemporals; circumorbital plates
lightly ossified. Palatoquadrate complex consisting of endoptery-
goid and ectopterygoid (both toothed on medial surface), quadrate,
and metapterygoid, the latter smooth and having widened border
for articulation on anterodorsal margin. Pectoral girdle consisting
of cleithrum and clavicle (supracleithrum not seen); small projec-
tion on medial surface of posterior portion of cleithrum; horizontal
medial process on clavicle. Pelvic plate bearing three anteriorly
diverging apophyses, and one denticulate ventromedian process
for articulation to opposite plate. Lepidotrichia jointed distally,
480 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
but not tuberculated. Scales oval, having posteriorly converging
ridges on posterior exposed parts.
The name refers to the most distinctive character of the genus,
the connected antotic and basipterygoid processes on the basi-
sphenoid, and is derived from Greek, synaptos—joined, tylos
(masc. )—knob, projection.
Synaptotylus is excluded from the advanced suborder Coela-
canthoidei by the retention of basipterygoid processes on the basi-
sphenoid. Synaptotylus differs from Rhabdoderma in several
characters of the basisphenoid, the most important being: knoblike
antotic processes (those of Rhabdoderma are wider, more flattened
and more dorsal in position); small, lateral basipterygoid processes
(in Rhabdoderma these are larger and farther ventral in position).
Synaptotylus newelli (Hibbard )
Gece as newelli Hibbard, 1933, Univ. Kansas Sci. Bull., 21:280, pl. 27,
eS PAO
Coelacanthus arcuatus Hibbard, 1933, Univ. Kansas Sci. Bull., 21:282, pl. 26,
fig. 8; pl. 27, fig. 1.
Rhabdoderma elegans Moy-Thomas, 1937 (in part), Proc. Zool. Soc. London,
107 (ser. B, pt. 3) :399.
Type.—k. U. no. 786F.
Diagnosis.—Same as for the genus.
Horizon.—Rock Lake shale member, Stanton formation, Lansing group,
Missouri series, Upper Pennsylvanian.
Localities—The specimens studied by Hibbard (K.U. nos. 786F, 787F,
788) and no. 11457 were taken from the Bradford Chandler farm, from the
original quarry in SW, SEX, sec. 82, T.19S, R.19E. The remainder were
collected from University of Kansas Museum of Natural History locality
KAn-1/D, a quarry in sec. 5, T.19S, R.19E. Both of these are approximately
six miles northwest of Garnett, Anderson County, Kansas.
Referred specimens.—K. U. nos. 786F, 787F, 788, 9939, 11424, 11425,
11426, 11427, 11428, 11429, 11480, 114381, 114382, 1143838, 11484, 11449,
11450, 11451, 11452, 11458, 11454, 11455, 11457.
Preservation.—Preservation of many of the specimens is good, few are
weathered, but most of the remains are fragmentary and dissociated. One
specimen (the type, no. 786F) and half of another were nearly complete.
Specimens are found scattered throughout the Rock Lake shale (see p. 498).
Morphology—Terminology used for bones of the skull is that of Moy-
Thomas (1937) and Schaeffer (1952).
Endocranium and parasphenoid
The basisphenoid (see fig. 1) has been observed in only one
specimen (K.U. no. 9939) in posterodorsal and ventral views.
The basisphenoid, although somewhat crushed, appears to be fused
to the parasphenoid. Both antotic and basipterygoid processes are
A New GENwus OF PENNSYLVANIAN FIsH 481
frontal frontal
alisphenoid alisphenoid
sphenoid basipterygoid sphenoid
condyle process condyle
antotic : antotic
process process
basisphenoid parasphenoid notochordal socket
A B
basisphenoid
anfotic
process
basipterygoid
process
parasphenoid
Fic. 1. Synaptotylus newelli (Hibbard). Restoration of the basisphenoid,
based on K. U. no. 9939, x 5. A, lateral view, B, posterior view, C, ventral
view.
present, and are connected by a low, rounded ridge. The antotic
processes are large, bulbar projections. These processes in Rhab-
doderma are wider and more flattened (Moy-Thomas, 1937:figs.
3, 4). The antotic processes are at mid-point on the lateral surface,
not dorsal as in Rhabdoderma, and both the processes and the
ridge are directed anteroventrally. The basipterygoid processes
are smaller, somewhat vertically elongated projections, situated at
the end of the low connecting ridge extending anteroventrally from
the antotic processes, and are not basal as are those of Rhabdo-
derma. The sphenoid condyles, seen in posterior view, issue from
the dorsal margin of the notochordal socket. The margins of the
socket are rounded, and slope down evenly to the center. A slight
depression situated between and dorsal to the sphenoid condyles
is supposedly for the attachment of the intercranial ligament
(Schaeffer and Gregory, 1961:fig. 1). The alisphenoids extend
482, UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
upward, anterodorsally from the region above the sphenoid con-
dyles, and may connect to ridges on the ventral surface of the
frontals. The lateral laminae are not preserved, and their extent
is unknown.
In viewing the changes in the endocranium of Carboniferous
and Permian coelacanths, it would be well to consider the me-
chanical relationship of the loss of the basipterygoid processes to
the effect on swallowing prey. Evidently many of the coela-
canths, Latimeria for example, are predators (Smith, 1939:104); to
such fishes a more efficient catching and swallowing mechanism
would be an adaptive improvement. Stensid (1932:fig. 14) pre-
sents a cross section of the ethmosphenoid moiety of the endo-
cranium of Diplocercides kayseri (von Koenen) showing the meta-
pterygoid of the palatoquadrate loosely articulated to both the
antotic and basipterygoid processes. According to Tchernavin
(1948:1387) and Schaeffer and Rosen (1961:190) the swallowing
of large prey depends on the ability of the fish to expand its oral
cavity by allowing the posteroventral portion of the palatoquadrate
and the posterior end of the mandible to swing outward. Where
the palatoquadrate articulates with the basisphenoid at the antotic
and basipterygoid processes, as in the Devonian coelacanths, it
can not swing so far laterally as where it articulates with only the
dorsal, antotic process. Perhaps the loss of the basipterygoid articu-
lation reflects the development of a more efficient mechanism for
swallowing prey in these fishes. Schaeffer and Rosen (1961:191,
193) show that in the evolution of the actinopterygians several
changes improved the feeding mechanism: some of these changes
are: (1) freeing of the maxilla from the cheek, giving a larger
chamber for the action of the adductor mandibulae; (2) develop-
ment of a coronoid process on the mandible; and (3) increase in
torque around the jaw articulation. In coelacanths, at least some
comparable changes occurred, such as: (1) loss of the maxillary,
thus increasing the size of the adductor chamber; (2) develop-
ment of the coronoid bone, affording a greater area for muscle
attachment; (3) development of an arched dorsal margin on the
angular; (4) modification of the palatoquadrate complex, with
resultant loss of the basipterygoid processes. In Synaptotylus the
basipterygoid processes are small, not basally located, and perhaps
not functional. A more efficient feeding mechanism developed
rapidly during the Carboniferous and has remained almost un-
altered.
A New GENUs OF PENNSYLVANIAN FIsH 483
Fic. 2. Synaptotylus newelli (Hibbard). Restoration
of the parasphenoid, based on K. U. nos. 9939, 11451,
x5. A, ventral view, B, dorsal view and cross
sections.
The parasphenoid (see fig. 2) is a shovel-shaped bone having a
wide anterior portion and a narrower posterior portion of nearly
uniform width. Most of the ventral surface is covered with minute
granular teeth. The anterior margin is flared and curved postero-
medially from the lateral margin to a median triangular projection.
The lateral margins curve smoothly from the greatest anterior width
to the narrow central portion, where the margins become somewhat
thickened and turned dorsally. Posterior to this the lateral margins
are probably nearly straight. The external surface of the anterior
section is nearly flat and has a central depressed area the sides of
which slope evenly to the center. The internal surface is smooth
and centrally convex. Because of the fragmentary nature of all
four observed specimens, total length was not measured but is
estimated to be 15 to 20 mm. The opening of the hypophyseal
canal was not present, possibly because of crushing. Ethmoidal
ossifications were not preserved in any of the specimens studied.
The parasphenoid differs from that of Rhabdoderma elegans (New-
berry) in being more flared and widened anteriorly and more con-
cave centrally.
484 UNIVERSITY OF KANnsAs PuBsLs., Mus. Nar. Hist.
Dermal bones of the skull
Various portions of the cranial roof are preserved in several
specimens (see fig. 3). For comparisons with Rhabdoderma ele-
gans, see Moy-Thomas (1937:fig. 1).
The premaxillaries and rostral elements are not preserved in any
of the specimens. Only one pair of relatively large frontals have
been observed; they are 5.5 to 9.0 mm. long and 2.0 to 3.5 mm.
wide. These are nearly flat bones, with the greatest width pos-
teriorly 0.1 to 1.0 mm. wider than the anterior portion. The mid-
line suture is straight, the lateral margins are nearly straight, the
anterior margin slopes evenly posterolaterally, and the posterior
margin is slightly convex to straight. The anterior margin in R.
elegans is essentially straight. Ornamentation consists of sparse,
unevenly spaced, coarse tubercles or short striae. In one specimen
both bones have small clusters of tubercles near the lateral margins
and about 2.0 mm. from the posterior margin. None of these
bones has alisphenoids or ridges on the ventral surface, as Stensi6
(1921:65, 97) described for Wimania and Axelia.
Only six supraorbitals have been preserved (see fig. 3). These
are nearly square, flat, thin bones lying nearly in place adjacent to
intertemporal
frontal
lacrimo-jugal
dentary coronoid subopercular
splenial angular
Fic. 3. Synaptotylus newelli (Hibbard). Diagram of the dermal bones
of the skull, in lateral view, based on K. U. nos. 788 and 11432. x 2%
approximately.
A New GENUus oF PENNSYLVANIAN F'IsH 485
a frontal on K. U. no. 788. The smallest is anterior; the margins
of all are nearly straight. The bones are unornamented. Each
bears a pore of the supraorbital line just below the midline. The
supraorbitals of R. elegans have a triangular outline and do not
bear pores.
Intertemporals (fig. 3) on several specimens vary from approxi-
mately 9.0 to 15.0 mm. in length, 2.0 to 2.7 mm. in anterior width,
and increase to 4.5 to 8.0 mm. in maximum posterior width. The
midline suture is straight, the anterior margin is concave and the
lateral margin proceeds laterally in a concave curve to the widest
portion. In R. elegans only the anterior half of the corresponding
margin is concave. The posterior margin is slightly rounded and
slopes anteriorly toward the lateral margin. Ornamentation is
usually of randomly oriented tubercles and striae, although striae
are more common in the posterior third and may be longitudinal,
whereas tubercles occur mainly on the anterior section. No evi-
dence of sensory pores, as seen on the intertemporal of R. elegans,
has been found.
The supratemporals were observed on only one specimen (K. U.
no. 788), (fig. 3). Sutures were difficult to distinguish but the
medial margin is presumed to curve to fit and to articulate with the
lateral margins of the intertemporals. Lateral margins are smoothly
curved but the anterior and posterior margins were broken off.
There appears to be no ornamentation on this bone. The supra-
temporals are much more elongated and curving than those in R.
elegans.
The cheek region is nearly complete in one specimen (K. U. no.
788), and scattered parts occur in a few others (see fig. 3). The
lacrimojugal of no. 788 is elongate, with both ends curving dorsally.
It differs from the lacrimojugal in R. elegans, in which the anterior
end extends anteriorly and is not curved dorsally. The posterior
and anterior margins are not preserved; the greatest height ap-
pears to be posterior. Pores of the suborbital portion of the infra-
orbital sensory canal are seen on the dorsal surface of the bone.
In R. elegans the pores are on the lateral surface. A section of
the lacrimojugal on specimen no. 11425, broken at both ends, shows
a thin layer of bone perforated by the pores and covering a groove
for the canal within the dorsal margin of the bone. Both speci-
mens are unornamented.
A nearly complete postorbital (fig. 3) on specimen no. 788 is
nearly triangular, with the apex ventral. The concave anterior
486 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
margin bears pores of the postorbital part of the infraorbital line.
Ornamentation consists of widely spaced, coarse tubercles.
Part of one squamosal is preserved. It is somewhat triangular
and its apex is ventral. This bone is associated with the postorbital,
subopercular and lacrimojugal on no. 788. The preopercular sen-
sory line passes down the curving ventral margin of this bone, and
extends ventrally onto a narrow projection. A low ridge, nearly
vertical, passes dorsally from about midpoint of the canal to the
dorsal portion. The anterior margin is nearly straight, the ventral
margin is concave, and the dorsal margin is convex dorsally but
may be incomplete. Perhaps the squamosal and preopercular are
fused. The surface appears smooth; the view may be of the
medial side. The squamosal of R. elegans is nearly triangular and
notably different from that of Synaptotylus newelli.
The subopercular (fig. 3) shows closely spaced tubercles on the
lateral surface. The bone is an elongated, irregular triangle with
the apex pointing anterodorsally. The margins are incomplete,
except for the concave, curving anterior margin.
Numerous operculars (fig. 3) occur in the suite of specimens,
both isolated and nearly in place. Each is subtriangular; the apex
of the triangle is ventral. A slight convexity projects from the
anterodorsal border. The posterior margin is broadly but shallowly
indented. Otherwise the margins are smooth. Maximum height
ranges from 8.0 to 11.0 mm., and maximum width from 8.0 to 18.0
mm. Ornamentation varies from a few widely spaced, randomly
oriented tubercles to closely spaced tubercles merging posteriorly
into striae. On some specimens these are parallel to the dorsal
border, and oblique in the central portion. On the posterior
margins of several operculars the striae break up into tubercles.
A few operculars have closely spaced tubercles over much of the
surface. The internal surface is smooth.
Visceral skeleton
The palatoquadrate complex, best seen on K. U. no. 9939 (fig. 4),
consists of endopterygoid, ectopterygoid, metapterygoid and quad-
rate. No trace of epipterygoids, dermopalatines or autopalatines,
such as Moy-Thomas (1937:392, fig. 5) described for Rhabdoderma,
has been observed.
The endopterygoid has a long, ventral, anteriorly-directed process,
and an anterodorsal process that meets the metapterygoid in form-
ing the processus ascendens. The suture between the endoptery-
A New GENUS OF PENNSYLVANIAN FIsH 487
processus
ascendens
metapterygoid
endopterygoid
quadrate
processus
ascendens
metapterygoid
quadrate
endopterygoid
Fic. 4. Synaptotylus newelli (Hibbard). Restoration of the palatoquadrate
complex, based on K. U. no. 9939, x 5. A, medial view, B, lateral view.
goid and metapterygoid, seen in lateral view, is distinct in some
specimens and has an associated ridge; these bones appear to be
fused in others, without regard to size. This suture curves dorsally
from a point anterior to the quadrate and passes anterodorsally to
the extremity of the processus ascendens. The suture is visible
on the medial side only near the processus ascendens, for it is
covered by a dorsal, toothed extension of the endopterygoid. The
endopterygoid has a smooth lateral surface; the medial surface is
covered with tiny granular teeth, in characteristic “line and dot”
arrangement. The teeth extend onto the ventral surface of the
ventral process.
Two long, narrow, splintlike bones covered on one surface with
granular teeth are interpreted as ectopterygoids. These are 13.0
and 16.0 mm. long and each is 1.5 mm. wide. Orientation of these
is unknown, but they probably fitted against the ventral surface
488 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Fic. 5. Synaptotylus newelli (Hibbard). A, cerato-
hyal, lateral (?) view, based on K. U. nos. 11429 and
11457, x 5. B, urohyal, based on K.U. no, 11457,
xo.
of the ventral process of the endopterygoid (Moy-Thomas, 1937:
fig. 5).
The metapterygoid has a smooth surface in both views. The
dorsal edge has a thickened, flared margin that presumably articu-
lated with the antotic process of the basisphenoid. No articular
surface for the basipterygoid process has been observed.
The quadrate is distinct and closely applied to the posteroventral
margin of the complex. In medial view the margin is nearly straight
and continues to the ventral edge. The ventral surface is thick-
ened and forms a rounded, knoblike articular surface. In lateral
view the surface is smooth; the anterior margin is irregular (or
perhaps broken on all specimens), and proceeds in an irregular
convex curve from the posterior to the ventral margin.
The general shape of the palatoquadrate complex is most nearly
like that of Rhabdoderma elegans (Moy-Thomas, 1937:fig. 5).
The orientation of the complex in the living fish was probably
oblique, with the processus ascendens nearly vertical, the quadrate
oblique, and the ventral process of the endopterygoid extending
dorsoanteriorly and articulating with the parasphenoid.
Of the hyoid arch only the ceratohyals (see fig. 5A) are preserved
in several specimens. These are long, curved bones with a postero-
ventral process and widened, flaring posterior margin. The medial
(?) surface is concave in one specimen. The lateral (?) surface
displays a distinct ridge on several specimens, arising on the dorsal
A New GENUS OF PENNSYLVANIAN FISH 489
surface opposite the posteroventral process and extending diagon-
ally to the anteroventral end of the anterior limb. The impression
of one other specimen appears to have a central ridge because of
greater dorsal thickness and narrowness. Both surfaces are un-
ornamented.
The urohyal (see fig. 5B) is an unornamented, Y-shaped bone,
with the stem of the Y pointing anteriorly. Orientation with re-
spect to dorsal and ventral surfaces is uncertain. In one view a
faint ridge, also Y-shaped, occurs on the expanded posterior por-
tion, and the surface is convex. The anterior process has a convex
surface, sloping evenly off to the lateral margin; the opposite side
of the process has a concave surface. The posterior portion has a
slightly depressed area (see fig. 5B) at the junction of the “arms”
of the Y.
The five branchial arches are represented by the ceratobranchials,
several of which are preserved on K. U. no. 11431. These are long
bones with anteriorly curving ventral ends. The medial surfaces
are partly covered with minute granular teeth; only the dorsal
part is without teeth. The dorsal articular surface is convex dor-
sally and rounded.
The mandible (fig. 3), the best specimens of which are K. U.
nos. 788 and 11425, is seen only in lateral and ventral views, with
only angular, splenial and dentary visible.
The angular forms the main body of the mandible, and is similar
to that of Spermatodus. The dorsal margin of the angular is ex-
panded in the central region, with some variation. One specimen
has an expanded portion slightly anterior to that of the opposite
angular. The articular surface near the posterior end has not been
observed; the posterior end of the angular slopes off abruptly. The
anterior sutures are seen in only two specimens, K. U. nos. 788,
11425. The dentary meets the angular in a long oblique suture;
the dentary gradually tapers posterodorsally and ends on the
dorsal surface of the angular. The splenial fits into a posteriorly
directed, deep V-shaped notch on the ventral surface. The latero-
ventral surface of the angular contains sensory pores of the man-
dibular line. The ventral surface extends medially into a narrow
shelf, approximately 1.0 mm. wide, which extends the full length
of the bone; the external surface of this shelf is smooth and slightly
concave dorsally. Ornamentation of the angular consists of tu-
bercles and longitudinal or oblique striae, occurring mostly on the
expanded portion. The medial surface is not seen. Several broken
490 UNIVERSITY OF KAnsAS PuBts., Mus. Nat. Hist.
specimens show a central canal filled with a rod of calcite; in one
of these the sensory pores are also calcite-filled and appear to be
connected to the rod. Thus the pores originally opened into a
central canal.
The dentary is an unornamented bone with the anterior half
curving medially; the greatest height is anterior. This bone in
specimen K. U. no. 11425 bears irregularly spaced, simple, recurved,
conical teeth; nine were counted, but there is space for many
others. One other specimen, no. 11429, seems to have tiny tubercles
on the surface. The dentary meets the splenial dorsally in a
straight suture.
The splenial also curves medially, and as stated, meets the dentary
in a straight suture. Ornamentation on this bone was not observed.
The posterior margin is V-shaped and fits the notch in the angular.
The ventral surface bears three or more sensory pores of the man-
dibular line.
The gular plates are oval. The medial margin is straight to
slightly curved, the lateral margin curved crescentically, the pos-
terior end is blunt, and the anterior end somewhat rounded. Orna-
mentation varies greatly; some bones show only a few tubercles,
whereas others exhibit an almost concentric pattern of closely
spaced striae. Typically there are some tubercles in the anterior
quarter or third of the total length; these pass into longitudinally
oriented striae in the posterior section. A few have only randomly
oriented, widely-spaced striae. The internal surface is smooth.
The coronoid (K. U. no. 11428) is a triangular bone, with the
apex pointing dorsally. The lateral surface is smooth; no teeth
were observed. Moy-Thomas (1937:292, 293) mentions several
tooth-bearing coronoids in Rhabdoderma, but as yet these have
not been seen in Synaptotylus.
Axial skeleton
Only three specimens (K. U. nos. 786F, 787F, 11450) show parts
of the vertebral column, but isolated neural and haemal arches are
numerous. All are of the coelacanth type, having Y-shaped neural
and haemal arches, without centra. A total count of 88 was ob-
tained, but this was incomplete; the actual number was probably
near 50, Counts of 10 and 16 haemal arches were obtained in two
of the specimens. Total height of neural arches ranges from 7.5
to 12.0 mm., and of haemal arches, from 9.0 to 12.0 mm. The
shorter arches are anterior and the height increases gradually to a
maximum in the caudal region. Height of the spines varies from
A New GENUS OF PENNSYLVANIAN FIsH 49]
4.0 to 9.0 mm., or from twice the height of the arch in the anterior
to three times the height in the caudal region. Total width of the
base, measured in isolated specimens because lateral views in other
specimens prevented measuring width, ranges from 0.7 to 4.2 mm.
The short, broad arches having short spines occur at the anterior
end of the spinal column; the narrower arches having tall spines
occur toward the caudal end. Broken neural and haemal arches
show a thin covering of bone with a central, calcite-filled cavity,
which in life may have been filled with cartilage (Stensid, 1932:58,
fig. 20).
No ossified ribs have been observed, either isolated or in place.
For further description of the axial skeleton, see Hibbard (1933).
Girdles and paired fins
A nearly complete pectoral girdle on specimen K. U. no. 11483
(see fig. 6A) has only a cleithrum and clavicle. No evidence of an
extracleithrum or supracleithrum has been observed, but the extra-
cleithrum may be fused to the cleithrum. The two bones form a
boot-shaped unit, with the anteroventral part turned medially to
form a horizontal process which meets the opposite half of the
girdle. In lateral view the surface is unornamented, and convex
in the ventral half. The suture between the cleithrum and clavicle
begins on the expanded posterior portion, the “boot-heel,” at a point
immediately below the greatest width on the posterior margin,
cleithrum
clavicle
Fic. 6. Synaptotylus newelli (Hibbard). Paired fin girdles. A, pectoral
girdle, lateral view, based on K. U. no. 11433, x 3.5. B, pelvic girdle basal
plate, medial (?) view, based on K. ae no. 788, < 8. Anterior is toward the
eft.
492 UNIVERSITY OF Kansas PuB3s., Mus. Nat. Hist.
passes anteriorly, then turns sharply and parallels the anterior
margin. The shape of the cleithrum resembles that in Rhabdo-
derma and the internal surface is not ridged (see Moy-Thomas,
1937:fig. 9). The exact orientation in the fish is uncertain, but if
the median extension is really horizontal, then the posterior ex-
pansion is directed caudally. The medial surface is concave, steep-
est near the anterior margin, and then slopes outward evenly. In
medial view one specimen (K. U. no. 11426) shows a small, cau-
dally directed projection of bone, evidently for articulation of the
fin-skeleton, at the widest portion of the cleithrum. Sutures on
several specimens were indistinct. Broken specimens show sutural
faces, but many nearly complete specimens show little or no indica-
tion of sutures, without regard to size of the girdles. The internal
structure of the fin was not observed.
Numerous isolated basal plates of the pelvic girdle have revealed
details of structure but no information on the orientation. Pre-
sumably the basal plates of Synaptotylus had essentially the same
orientation as those of other coelacanths (Moy-Thomas, 1937:395).
The most complete basal plate is K. U. no. 788 (see fig. 6B). The
three apophyses diverge anteriorly; the horizontal one is best de-
veloped and the dorsal one is least well developed. A median
process (Schaeffer, 1952:49), denticulate on several specimens,
articulates with the corresponding process of the opposite plate.
The expanded part that articulates with the skeleton of the fin
extends caudally. The posterior expanded part is nearly square in
outline, resembling the dorsal, rectangular projection. One side
bears ridges leading to the extremities of the apophyses, and faint
crenulations on the median process. This may be the medial view.
The other view displays a smooth surface, usually without indica-
tion of the ridges seen in the reverse view. These specimens differ
somewhat from the basal plates of Rhabdoderma and appear to be
intermediate between Rhabdoderma and Coelacanthus (Moy-
Thomas, 1937:fig. 10A, B). The apophyses are not free as in
Rhabdoderma but webbed with bone almost to their extremities,
as in Coelacanthus.
The pelvic fin is seen in only two specimens (K. U. nos. 786F,
788). That on no. 788 is lobate and has 25 lepidotrichia, jointed
for approximately the distal half, and 2.5 to 13.0 mm. in length.
Total length of the fin is 25.0 mm. There is no trace of the internal
skeletal structure or of the articulation to the basal plate in either
specimen. For a description of the fin on no. 786F, see Hibbard
(1988:281).
A New GENUws OF PENNSYLVANIAN FIsH 493
Unpaired fins
A few isolated bones on K. U. no. 788 (fig. 7) are interpreted as
basal plates of the unpaired fins. For additional description of the
unpaired fins on the type, K. U. no. 786F, see Hibbard (1933).
Two of these bones are flat, smooth and oblong, bearing a diag-
onal ridge that extends in the form of a projection. Orientation is
completely unknown. These may be basal plates of the anterior
dorsal fin. The fin on no. 786F that Hibbard (1933:281) interpreted
as the posterior dorsal fin is now thought to be the anterior dorsal
fin.
Fic. 7. Synaptotylus newelli (Hibbard). Basal
plates of unpaired fins. A, anterior dorsal fin,
based on K.U. no. 788, <x 10. B, posterior
dorsal fin, based on K. U. no. 788, x 12. C,
anal fin, based on K. U. no. 11450, x 5. An-
terior is toward the left.
494 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
One distinctive bone may represent the basal plate of the pos-
terior dorsal fin. This incomplete specimen shows two projecting
curved processes, bearing low but distinct ridges, which diverge,
probably anteriorly. The central portion is narrow. The two
ridges continue onto the posterior portion. This has been broken
off, but shows that the ridges diverge again. The surface is smooth,
except for the ridges. As before, orientation is uncertain. On no.
786F this fin was interpreted by Hibbard (1933:281) as the anal
fin.
Only part of one basal plate of the anal fin was preserved
on K.U. no. 11450. That plate is oblong and has an expanded
anterior end. The narrow, constricted part bears two oblique
ridges and a few tubercles. The posterior part has nearly straight
margins (represented by impressions) and the posterior margin
is oblique, sloping anteroventrally. The flared anterior part has
a smooth surface. This basal plate is more nearly like those of
Coelacanthus, according to the descriptions given by Moy-Thomas
(1937:399). The basal plate is associated with seven apparently
unjointed, incomplete lepidotrichia. The anal fin on no. 786F is
interpreted as the anterior dorsal fin (Hibbard, 1933:281).
The caudal fins are preserved on K. U. nos. 786F, 787F, and have
a total of 24 lepidotrichia, 12 above and 12 below. These are
jointed for the distal half or two-thirds, and are up to 16.0 mm.
in length. In specimen no. 787F the supplementary caudal fin
has at least seven lepidotrichia, the longest of which is 11.0 mm.
but incomplete. Anterior lepidotrichia appear unjointed but the
posterior ones are jointed for the distal two-thirds (?) (these are
broken off). The supplementary caudal fin is approximately 1.5
mm. long and 8.0 mm. or more wide. The supplementary caudal
fin on K.U. no. 786F described by Hibbard (1933:281) could
not be observed; this part of the caudal fin is missing.
Squamation
In the suite of specimens isolated scales are numerous, but patches
of scales are rare. Only two specimens (K. U. nos. 786F, 787F ) are
complete enough for scale counts, but preservation permits only
partial counts. In general the scales resemble those of Rhabdo-
derma elegans (Newberry ).
The scales are oval. The exposed posterior part of each bears
posteriorly converging ridges; the anterior part is widest and shows
a fine fibrillar structure. There are at least six scale-rows on either
_A New GENws OF PENNSYLVANIAN FIsH 495
side of the lateral line. Lateral line scales show no pores, and
except for slight irregularities in the orientation and length of the
posterior ridges, closely resemble the others. Central ridges on
the lateral line scales are shorter and tend to diverge from the
center of the impression of the canal. The lateral line canal shows
only as the impression of a continuous canal 0.7 mm. in diameter.
Preservation is poorest in scales along the line of the neural and
haemal arches; therefore lateral line scales are rarely preserved.
Isolated scales are of two types: those on which the posterior
ridges converge sharply and form the gothic arch configuration
mentioned by Hibbard (1933:282), and those which do not. Both
types of scales can be present on one fish, as shown by specimen
no. 788. This is not apparent on nos. 786F and 787F; all of the
scales on these specimens appear to be much alike. Both Moy-
Thomas (1937:385) and Schaeffer (1952:51, 52) have remarked on
the variation of the scales on different parts of the same fish. Be-
cause the number of ridges and amount of convergence of the
ridges is not related to size of the scale, it is concluded that these
characters are not of taxonomic significance.
The strong resemblance of the scales of the Garnett specimens
to those of Rhabdoderma elegans (Newberry) caused Moy-Thomas
(1937:399) to add Hibbard’s two species to the synonymy of R.
elegans. But at that time only the scales could be adequately
described. If the shape of the scale and the number and pattern
of ridges can vary with age, size and shape of the scale, it follows
that assignment of isolated scales to a species should not be at-
tempted. Assignment to genus should be made only with caution.
Discussion.—The relationship of Synaptotylus to other coela-
canths is obscure at present. The knoblike antotic processes on the
basisphenoid are unlike those of any other known coelacanth. The
palatoquadrate complex is shaped like that of Rhabdoderma elegans
but consists of fewer bones, probably because of fusion. The scales
resemble those of Rhabdoderma. With regard to general shape
of fin girdles, the pectoral girdle resembles that of Eusthenopteron
more than that of Rhabdoderma, but the cleithrum is more nearly
like the cleithrum of Rhabdoderma. The pelvic girdle appears to
be midway between those of Rhabdoderma and Coelacanthus in
general appearance. Regarding the basal plates of the remaining
fins, those of Synaptotylus appear to resemble basal plates of both
Rhabdoderma and Coelacanthus. Considering the structure of the
sphenethmoid region of the braincase, Synaptotylus is probably
496 UNIVERSITY OF KANSAs Pusts., Mus. Nat. Hist.
more closely related to Rhabdoderma than to other known coela-
canth genera.
COMMENTS ON CLASSIFICATIONS
Classification of Carboniferous coelacanths has been difficult,
partly because the remains are commonly fragmentary, and sig-
nificant changes in anatomy did not become apparent in early
studies. In general, coelacanths have been remarkably stable in
most characters, and it has been difficult to divide the group into
families. As Schaeffer (1952:56) pointed out, definition of coela-
canth genera and species has previously been made on non-
meristic characters, and the range of variation within a species has
received little attention. For example, Reis (1888:71) established
the genus Rhabdoderma, using the strong striation of the scales,
gular plates and posterior mandible as the main characters of this
Carboniferous genus. Moy-Thomas (1937:399-411) referred all
Carboniferous species to Rhabdoderma, redescribed the genus and
compared it to Coelacanthus, the Permian genus. He cited as
specific characters the ornamentation of the angulars, operculars
and gular plates (Moy-Thomas, 1935:39; 1937:385). Individual
variation in some species has rendered ornamentation a poor cri-
terion. This variation is apparent in Synaptotylus newelli (Hib-
bard), some specimens having little or no ornamentation; others
having much more. The number of ridges and pattern of ridges
on the scales also varies. Schaeffer (1952:56) has found this to
be true of Diplurus also. Moy-Thomas (1935:40; 1937:385) real-
ized that the type of scale is a poor criterion for specific differentia-
tion. In the search for features useful in distinguishing genera of
coelacanths, Schaeffer and Gregory (1961:3, 7) found the structure
of the basisphenoid to be distinctive in known genera, and thought
it had taxonomic significance at this level. Higher categories
should have as their basis characters that display evolutionary
sequences. A recent classification (Berg, 1940), followed in this
paper, reflects two evolutionary trends in endocranial structure of
coelacanths: reduction of endocranial ossification and loss of the
basipterygoid processes. Because there has been little change in
other structures in coelacanths, Berg’s classification is the most
useful. Berg (1940:390) includes Rhabdoderma in the suborder
Diplocercidoidei because of the presence of the basipterygoid proc-
esses, and in the single family, Diplocercidae, but remarks that
because of the reduced amount of endocranial ossification the Car-
A New GENUs OF PENNSYLVANIAN Fis 497
boniferous Diplocercidae “probably constitute a distinct family.”
In considering this concept of classification, the subfamilies Diplo-
cercinae and Rhabdodermatinae of the family Diplocercidae are
proposed above. The subfamily Rhabdodermatinae includes at
present Rhabdoderma and Synaptotylus. The principal characters
of the subfamily Rhabdodermatinae, named for the first known
genus, are the retention of the basipterygoid processes and the
reduction of endocranial ossification. Application of this classifica-
tion based upon endocranial structure would probably change
existing groupings of species of Carboniferous coelacanths; the
entire complex of Carboniferous genera should be redescribed and
redefined. It will be necessary to consider endocranial structure
in any future classification.
The greater part of the evolution previously mentioned appears
to have been accomplished during the Carboniferous; thereafter
coelacanth structure became stabilized. The trend progressed from
Devonian coelacanths which had two large unpaired bones in the
endocranium, and both antotic and basipterygoid processes on the
basisphenoid, to Carboniferous fishes in which ossification was
reduced to a number of paired and unpaired bones embedded in
cartilage, and retaining both processes, and then post-Carboniferous
kinds with reduced ossification and no basipterygoid processes.
The Pennsylvanian was evidently the time of greatest change for
the coelacanths, and they have not changed significantly since, in
spite of the fact that since the Jurassic they have shifted their en-
vironment from shallow, fresh water to moderate depth in the sea
(Schaeffer, 1953:fig. 1). The changes in endocranial structure
appear to be significant, and are perhaps related to higher efficiency
of the mouth parts in catching and swallowing prey (see p. 482).
ENVIRONMENT
The coelacanth fishes from the Rock Lake shale are part of the
varied fauna collected from Garnett. Peabody (1952:38) listed
many elements of the fauna and flora, and concluded that the de-
posits are of lagoonal origin. In addition to numerous inverte-
brates (including microfossils) and arthropods, a number of verte-
brates other than coelacanths have been found. These include at
least one kind of shark, Hesperoherpeton garnettense Peabody, one
or more kinds of undescribed labyrinthodonts and the reptiles
Petrolacosaurus kansensis Lane, Edaphosaurus ecordi Peabody, and
Clepsydrops (undescribed species ). This is indeed a rich vertebrate
498 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
fauna, and the earliest known reptilian fauna. Much of the rock
contains plant remains. The flora that has been identified is adapted
to growing in a well-drained soil; although it contains some elements
considered characteristic of the Permian, it is of Pennsylvanian age
(Moore ef al., 1936). Peabody (1952:38-39) discusses the features
of these lagoonal sediments. Much of the fauna and flora suggests
continental origin, but the many marine invertebrates at some
horizons indicate that at least some of the sediments were of
marine origin.
Little can be said about the actual environment of the living
fishes of the genus Synaptotylus. Remains of these fishes occur
in layers containing marine invertebrates, as well as in those con-
taining plant remains and vertebrate skeletal parts, and in those
nearly completely composed of dark carbonaceous material. Most
of the remains are fragmentary and consist of isolated bones, iso-
lated scales, and dissociated skulls; only one specimen and half of
another are nearly complete. Many published statements on Rhab-
doderma, a related genus, indicate both marine and fresh-water
environments. Wehrli (1931:115) regarded Rhabdoderma elegans
(Newberry) as a euryhaline species, and cited its occurrence with
both marine and fresh-water fossils. Aldinger (1931:199) also
found this to be the case with other species, and Fiege (1951:17)
quotes others as giving the same information. Keller (1934:913)
thought that few Carboniferous fishes were exclusively marine, and
stated that the majority of them became adapted to fresh water
during the late Carboniferous. Later, Schaeffer (1953:175) stated
that all Carboniferous and Permian coelacanths were fresh-water
fishes, and that many were from swamp deposits. If Keller is
correct, then members of the genus Synaptotylus may have in-
habited the lagoon, the adjacent sea, or the streams draining into
the lagoon. Perhaps these fishes swam upstream, as modern salmon
and tarpon do, although there is no direct evidence for this. Possi-
bly they lived in the lagoon at times of scant rainfall and little
runoff, when the salinity of lagoon water approached normal marine
values or the fishes may have lived in the streams, and after death
were washed into the lagoon. As numerous remains of land plants
and animals were washed in, perhaps this best accounts for the
presence of the fish in nearly all layers of the deposits, not only
the marine strata.
A New GENUs OF PENNSYLVANIAN FIsH 499
SUMMARY
A new genus of Pennsylvanian coelacanths, Synaptotylus, is de-
scribed and a previously named species, Coelacanthus newelli
Hibbard, 1933 (C. arcuatus Hibbard, 1933, is a junior synonym),
is referred to this genus. All specimens of Synaptotylus newelli
(Hibbard) were collected from the Rock Lake shale member of
the Stanton formation, Lansing group, Missouri series, six miles
northwest of Garnett, Anderson County, Kansas. Synaptotylus is
distinguished from all other coelacanths by a basisphenoid having
large, knoblike antotic processes each connected by a low ridge
to a small basipterygoid process. Synaptotylus is most closely
related to Rhabdoderma, but is intermediate between Rhabdoderma
and Coelacanthus in shape of the fin girdles and basal plates. Two
new subfamilies, Diplocercinae and Rhabdodermatinae, of the
family Diplocercidae, are proposed. Synaptotylus and Rhabdo-
derma are included in the subfamily Rhabdodermatinae, because
both exhibit reduced ossification in the endocranium and retain
basipterygoid processes.
Loss of the basipterygoid processes in post-Carboniferous coela-
canths may reflect the development of a more efficient feeding
mechanism, by allowing the palatoquadrate complex and mandible
to swing farther laterally and expand the oral cavity.
Synaptotylus newelli (Hibbard) may have occupied either the
sea or fresh water; these fishes occur in lagoonal deposits with rep-
tiles and amphibians, arthropods, marine invertebrates and remains
of land plants.
Because scale patterns on Synaptotylus and Rhabdoderma are so
nearly similar and vary with size of the scale and its location on the
fish, it is recommended that isolated scales not be assigned to a
species, and to a genus only with great caution.
500 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hisr.
LITERATURE CITED
ALDINGER, H.
1931. pees karbonische Fische aus Westfalen. Paleont. Zeit., 13:186-
Bere, L. S.
1940. Classification of fishes, both Recent and fossil. Moscow and
Leningrad, 1940 (J. W. Edwards, Ann Arbor, Michigan, 1947,
offset reproduction, pp. 1-345, 197 figs., plus English translation
of text, pp. 346-517, 1947.
Firce, K.
1951. Eine Fisch-Schwimmspur aus dem Culm bei Waldeck. Neues
Jahrb. Geol. and Paliont. Jahrgang 1951:9-81.
HIBBArD, C. W.
1933. Two new species of Coelacanthus from the middle Pennsylvanian
of Anderson County, Kansas. Kansas Univ. Sci. Bull., 21:279-287.
KELLER, G.
1934. eet aus dem oberkarbon des Ruhrgebiets. Gluckauf, 70:
13-917.
Moore, R. C., Extas, M. K., and NEwE tL, N. D.
1936. A “Permian” flora from the Pennsylvanian rocks of Kansas. Jour.
Geol., 44:1-31.
Moy-THomas, J. A.
1935. A synopsis of the coelacanth fishes of the Yorkshire Coal Measures.
Ann. Mag. Nat. Hist., 15 (ser. 10): 87-46.
1937. The Carboniferous coelacanth fishes of Great Britain and Ireland.
Proc. Zool. Soc. London, 107 (B): 383-415.
Prasopy, F. E.
1952. Petrolacosaurus kansensis Lane, a Pennsylvanian reptile from Kan-
sas. Kansas Univ. Paleont. Contrib., 1:1-41.
Reis, O. M.
1888. Die Coelacanthinen mit besonderen Beriicksichtigung der im Weis-
sen Jura Bayerns verkommenden Arten. Palaeontographica, 85:
1-96.
SCHAEFFER, B.
1952. The Triassic coelacanth fish Diplurus, with observations on the
evolution of the Coelacanthini. Bull. Amer. Mus. Nat. Hist., 99:
art. 2, 29-78.
1953. Latimeria and the history of the coelacanth fishes. New York
Acad. Sci. Trans., (2) 15:170-178.
SCHAEFFER, B., and Grecory, J. T.
1961. Coelacanth fishes from the continental Triassic of the western
United States. Amer. Mus. Novitates, 20386:1-18.
ScHAEFFER, B., and Rosen, D. E.
1961. Major adaptive levels in the evolution of the actinopterygian feed-
ing mechanism. Am. Zool., 1:187-204.
Smiru, J. L. B.
1939. A living coelacanthid fish from South Africa. Trans. Roy. Soc.
South Africa, 28:1-106.
STENSIO, E. A.
1921. Triassic fishes from Spitzbergen. Part I. Vienna, Adolf Holz-
hausen: 1-307.
1932. Hea fishes from East Greenland. Meddel. om Grgnland, 38:
1-305.
A New GENws OF PENNSYLVANIAN FIsH 501
TCHERNAVIN, V. V.
1948. On the mechanical working of the head of bony fishes. Proc. Zool.
Soc. London, 118:129-143.
WEHRLI, H.
1931. Die Fauna der Westfilischen Stufen A und B der Bochumer Mulde
zwischen Dortmund und Kamen (Westfalen). Palaeontographica,
74:93-184,
Transmitted March 29, 1962.
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UNIVERSITY OF KANSAS PUBLICATIONS ** |
MvuSEUM OF NATURAL HISTORY
Volume 12, No. 11, pp. 503-519
October 25, 1963
Observations on the Mississippi Kite
in Southwestern Kansas
BY
HENRY S. FITCH
UNIVERSITY OF KANSAS
LAWRENCE
1963
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HisToRY
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 12, No. 11, pp. 503-519
Published October 25, 1963
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY
JEAN M. NEIBARGER, STATE PRINTER
TOPEKA, KANSAS
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Observations on the Mississippi Kite
Thy ry
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in Southwestern Kansas Seecetenate |
BY
HENRY S. FITCH
The Mississippi kite (Ictinia misisinpiensis) is one of the common
raptors of Kansas, occurring regularly and abundantly in summer
in that part of the state south of the Arkansas River. In 1961, in an
attempt to find out more about the ecology of the species in Kansas,
I made several trips to parts of the state where kites could be found
in numbers, notably to Meade County State Park in the south-
western part of the state, 7% miles south and five miles west of
Meade. Little has been written regarding the species in this ex-
treme northwestern part of its breeding range, where it thrives
under ecological conditions much different from those that prevail
elsewhere in its range. Also, the social behavior and food habits
have been given relatively little attention.
In my field study I was helped by my son, John H. Fitch, who
climbed to many kite nests and spent many hours observing in the
field. My daughter, Alice V. Fitch, likewise aided me by keeping
nests under surveillance. Dr. Claude W. Hibbard of the University
of Michigan and Mr. Harry Smith, superintendent of Meade State
Park, also kindly provided much useful information concerning
the history of the colony of Mississippi kites at the Park. Mr.
William N. Berg analyzed pellets, and Dr. George W. Byers kindly
checked many of the identifications, and provided generic and
specific determinations for some of the insects.
In general, the range, habits and ecology of the Mississippi kite
are already well known through the publications of Audubon
(1840), Chapman (1891), Bendire (1892), Ganier (1902), Wayne
(1910), Nice (1931), Bent (1936), Sutton (1939) and Eisenmann
(1963). The breeding range is the southeastern United States,
chiefly within the Austroriparian Life-zone, but extending north-
west through much of Oklahoma and into southern Kansas. The
species is highly migratory. Wintering Mississippi kites are known
from Argentina and Paraguay (Eisenmann, op cit.:74), and most of
the population probably winters in southern South America, but
records outside the breeding range are few.
The Mississippi kite is perhaps one of the most social raptors. It
is highly gregarious, not only in its migrations but in breeding
(505)
506 UNIVERSITY OF KANSAS PuB3s., Mus. Nat. Hist.
colonies. All breeding pairs seen were closely associated with other
individuals, with no territorial hostility; signs of intraspecific intoler-
ance are rare, even where the kites are abundant. In the nesting
season many of both sexes perch together in the same tree, and
groups tend to keep together as they forage.
Secondary sexual differences are slight. Seven males in the Uni-
versity of Kansas Museum of Natural History collection average
351 (342 to 360) millimeters in length, and six females average 361
(348 to 370) millimeters. Sutton (op. cit.:44) collected 16 breeding
kites near Arnett, Oklahoma in 1986 and 1987 and recorded that
eleven males averaged 245 (216 to 269) grams and five females
averaged 311 (278 to 339) grams. As indicated by Sutton, the
head is paler in the adult male than in the female, and at close range
this difference will serve for identification of the sexes. The differ-
ence in size is scarcely noticeable in the field.
Habitat
In Kansas this kite seems to prefer open and even barren terrain,
in contrast with its habitat in forests of the southeastern states.
Typical habitat of Kansas is that of the High Plains, dominated by
a short-grass climax of blue grama (Bouteloua gracilis) and buffalo
grass (Buchloé dactyloides), with sagebrush (Artemisia sp.),
prickly pear (Opuntia sp.) and other somewhat xerophytic vege-
tation. In the Gypsum Hills of south-central Kansas near the
Oklahoma border, the Mississippi kite finds habitat conditions ex-
ceptionally favorable. This is an area of broken topography, dis-
sected by small steep-sided ravines, often with brush and scrubby
trees on the slopes.
At Meade County State Park groves of cottonwoods (Populus del-
toides) provided abundant places for perching and nesting. At
this locality an artesian well provided an abundant year round
water supply, which was impounded into an artificial lake half a
mile long and a little less than a quarter mile wide. Water was
also impounded in a series of small ponds maintained for the benefit
of fish and waterfowl. Along with other improvements extensive
plantings of cottonwoods and other trees were made with relief
labor in the nineteen thirties. Trees were scarce on the area orig-
inally, but by 1961 there were almost continuous groves in an area
nearly two miles long and three quarters of a mile wide encom-
passing the lake and ponds and adjacent areas. In conversation
at the Park in August 1961, Dr. C. W. Hibbard told me of his
observations on the colony of kites since 1936 when his paleonto-
MISsSISSIPPI KITE 507
logical field work in that area was begun. He indicated an area of
less than two acres west of the artesian well to which the colony
had been limited in its nesting in 1936, because at that time few
trees were available as nest sites. In subsequent years, as the trees
in the artificially established groves increased in size and height, and
other trees became established naturally where the impoundments
had created favorably moist conditions, the nesting colony ex-
panded in all directions, and the number of kites increased tre-
mendously. When my observations were made in 1961, the nesting
area was co-extensive with the cottonwood groves, and there were
literally thousands of trees within the area that provided adequate
sites for nests.
Numbers
The maximum number of kites seen flying at one time at the
Park was 44, on August 22, 1961. Probably almost all there were
adults, because fledglings, even though able to fly strongly by this
date, were still spending most of their time perched. The colony
of kites was usually scattered over at least two square miles, and
at most times some were perched, others were flying low and
solitarily, hence it is improbable that the total population or a high
percentage of it could be seen together at any one time or place.
More than 40 nests were located in 1961, and probably at least as
many more were overlooked. There must have been a breeding
population of at least 100 kites, and probably as many as 150 in the
Park in 1961. H. B. Tordoff recorded on the label of K. U. Mus.
Nat. Hist. no. 30514, taken on September 1, 1951, in Barber County,
Kansas, that it was one of at least 200 at a communal roost.
Feeding
The Park and its vicinity stood out as a veritable oasis in an
almost treeless region of open rolling topography, with a short-grass
type of vegetation dominating. The kites displayed versatility in
their choice of places to forage. Often they soared over the cotton-
wood groves, the lake, or the ponds, but at other times they flew
far out over the plains, and seemed to prefer such open situations.
A small herd of buffalo was maintained at the Park, and their closely
grazed pastures of several hundred acres were favorite foraging
grounds for the kites. Often the kites and buffalo were seen in
close association, and at times the kites must have benefited from
the movements of the buffalo, serving to flush certain insects such
as grasshoppers. The latter were probably the chief food source
of the kites in the heavily grazed pastures. Bent (1936:67) stated:
508 UNIVERSITY OF KANSAS PuBis., Mus. Nat. Hist.
“A flock of from 3 to 20 will sail about a person, a horseman or a
team, traveling through grassy flats or bushy places, and seize the
cicadas as they are scared up.” Dr. Hibbard told me that on one
occasion when he had caught a number of cicadas, he fed them to
a pair of kites by tossing them into the air one by one, and each
was seized by a kite which was flying nearby waiting expectantly.
Mississippi kites are noted for their buoyant and seemingly al-
most effortless flight, and their prey is caught while they are on
the wing. In extended flights the kites soar, drift and circle with
frequent easy flapping, at variable heights. Sometimes they are
several hundred feet above the ground. Doubtless the height is
influenced by the types of insects that are flying, and where they
can be found most readily. Even at close range the catching of
prey by a kite is likely to be overlooked by an observer. After
being snatched from the air, the prey is usually eaten while the kite
is still in flight, and the movements of the head in pecking at the
objects held in the talons are much more noticeable than the slight
veering from the course of flight that signals the actual capture.
Kites were often watched while they were hunting in the open
areas around the Park. On June 1, 1961, my son and I observed
16 perched together in a small tree. From time to time each kite
would leave the tree in a short flight low over the surface of a
nearby pool, where it would snatch up prey, probably a dragonfly
in many instances, and would return to a perch to feed. Most of
the time one or several kites were in flight while the majority were
perched. Similar observations were made on smaller groups
perched on fence posts along the edges of large pastures. Gre-
garious tendencies were evident from the fact that two or more of
the kites perched fairly near together on separate but sometimes
adjacent fence posts. Each kite in turn would glide from its post,
skim low over the ground surface for a few seconds, seize its prey
with a sudden slight swerving, and return to the fence (usually to
a different post from the one it had left) to feed upon the insect
captured. Grasshoppers of many species were abundant in the
area. It seemed that grasshoppers were flushed from the ground
by the bird flying near them and were picked off before they were
well underway. In any case the prey was taken from the air rather
than from the ground in all observed instances. Ganier (1902:86)
mentioned seeing one of these kites alight on the ground in a
cotton field, where it stayed for more than a minute, but perching
on the ground is unusual.
Most often kites that were catching their prey by skimming close
MuississrePi KiTE 509
to the ground did not return to a perch but ate while they were
flying. Associations of groups on posts at edges of fields, in trees
or in flight were ephemeral as each bird seemed driven by a restless
urge to be in motion. The kites generally gave the impression of
catching their prey effortlessly and casually in the course of their
flights. However, on July 20, 1961, one flying over a pond was
seen to swoop three times in rapid succession at a dragonfly without
catching it. The kite then flew higher, circled, and swooped three
times more at the dragonfly, catching it on the last attempt. Most
of the insects preyed upon are slower and less elusive than dragon-
flies, which are largely immune to the attacks of flying predators
because of their great prowess in flight.
Only on rare occasions could the kind of prey captured be ob-
served in the field. Food habits were studied by collecting pellets
of the kites at the Park, and analyzing them. The pellets were
usually disgorged early in the morning while the kites were still
on their night roosts in large cottonwoods. Often several kites
roosted in the same tree. The pellets were of characteristic appear-
ance, elliptical, approximately 15 millimeters in diameter, 30 milli-
meters long, pinkish or purplish, composed of insects’ exoskeletons
compacted, and comminuted to about the consistency they would
have after passing through a meat grinder.
A total of 205 pellets was collected—37 on August 20, 1960;
56 on July 18, 1961; 60 on August 4 and 5, 1961, and 52 on August
21 to 23, 1961. A total of 453 separate items was tentatively identi-
fied. Obviously the material was far from ideal for the identifica-
tion of prey, which had to be reconstructed from minute fragments.
The kites are dainty feeders and discard the larger and less digesti-
ble parts such as wings, legs, and heads. Often it was uncertain
how many individuals or how many kinds of insects were repre-
sented in a pellet. Probably most pellets contained many indi-
viduals of the same species, but these were not separable. Hence,
only 2.2 items per pellet were found, whereas Sutton found an
average of 22.2 items in each of the 16 stomachs that he examined.
Best information concerning kinds of prey utilized was obtained
soon after the fledglings had left the nest; on various occasions
these still clumsy young dropped nearly intact insects that were
delivered to them by the adults. These insects, recovered from
beneath the perches, were the basis for all specific and generic
determinations; other material was determinable only to order or
to family.
One of the most significant outcomes of the examination of pellets
510 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
was the finding that vertebrates were scarcely, if at all, represented
in the food. Three pellets contained shreds that seemed to be
mammal hairs, but in the absence of other remains, the diagnosis
is somewhat doubtful. Many species of small mammals, birds,
reptiles and amphibians were common in the Park or its vicinity,
but insects made up nearly all the recorded prey. Audubon (1840:
73) mentioned lizards and small snakes in the food and gave a
dramatic but perhaps imaginative account of a kite swooping and
snatching a lizard (anole) from the topmost branch of a tree. Goss
(1891:251) stated: “I have seen them swoop down, and, with their
claws, snatch lizards from the ground, rocks and old logs, sometimes
stopping to eat them, but, as a rule, feeding on the wing.” Bendire
(1892:179) stated that the food was mostly insects “probably
varied with a diet of small rodents, lizards and snakes.” Wayne
(1910:71) stated that the food consisted almost entirely of insects
and lizards. Bent (1936:67-68), after stating that small snakes,
lizards and frogs were sometimes taken, cited a statement in the
notes of G. W. Stevens that the latter had found the remains of
toads, mice and young rabbits in nests with young. However,
Sutton (op cit.:51) in a detailed analysis of the stomach contents
of 16 kites in Oklahoma, found only insects and remains of one
small fish among a total of 358 prey items. Predation on vertebrates
must be rare, and perhaps requires further verification in view of
the rather vague character of the records so far published.
The following list includes both the prey found beneath perches
of fledglings and that identified from pellets, the latter mostly from
adult kites.
coleopteran orthopteran
unspecified ..............-. 187 unspecitied: )s). a4a48scieeneeee 120
Carabige 6 vA, ss Set 89 locustid }
ea WUNSPECIEd 4 uk oe ciene oe. 84
aa rary Arphia Crass@ 2.3. 6.0 1
unspecified .... ......... 18 Melanoplus cf, differentialis, 2
Cicindela sp. : 2 Schistocerca cf. lineata .... 1
hydrophilid Xanthippus corallipes ..... 2
unspecified” BA. Usa. aise 18 oe A 3
WUNSPECiMEd: Hess. 05 Lous ah
Hyanous iiscakl hia ei ; Dathinig isp. oo. ciaostya aot 1
scarabaei d homopteran
MHSPECHIE: W3..5 ., Aum ebede aus 1 eed
Canthonitspsdss, soe tant 8 unspecified .............. 15
silphid Tibicen cf. pruinosa ...... 1
Necrophorus sp. .......... 1
lepidopteran (unspecified moth), 3
Mississippi KITE 511
At Meade State Park I gained the impression that much of the
foraging is carried on near the nest. The short time lapse between
successive feedings was one indication, and from time to time while
keeping nests under observation, I saw kites that were individually
recognizable as the owners coursing back and forth in the vicinity.
However, only a few individuals were recognizable. For several
minutes before and after delivering food, such an adult was often
seen soaring within 200 to 300 yards of the nest, or sometimes
much closer. A somewhat different impression was received on
August 23, 1961, at Natural Bridge, south of Sun City, Barber
County, Kansas, where I observed two pairs of kites feeding fledg-
lings. One fledgling was seen to be fed ten times in a 1% hour
period. The transfer of food from the adult usually required less
than a minute. Then the adult would leave the tree, in a ravine,
and drift away. Circling and soaring, it seemed to be wandering
aimlessly, but within two or three minutes it was usually out of
sight over the horizon. In what appeared to be slow, lazy, flight
it usually drifted off to the west, to more upland areas of short grass
and sage brush. Once, watching from a high knoll I succeeded in
keeping it in view for almost five minutes, and during most of this
time it appeared to be between one and two miles away, but it
finally moved off even farther. Dr. Hibbard mentioned seeing
kites in the vicinity of the Jinglebob Ranch eight to ten miles from
the Park, and he believed that these individuals had come from
the Park since there was no suitable habitat in the intervening
areas. Actually, the distance could have been covered in a few
minutes’ flying time, but it is unlikely that these individuals were
feeding young at the Park, else they would not have wandered so
far. On several occasions groups of from three to 20 individuals
were seen in open terrain as much as four or five miles from the
Park.
Breeding Cycle
Probably kites arriving from their northward migration are al-
ready paired. In those observed at the Park in the first week of
June, there was no indication of courtship, or of sexual rivalry. On
June 1, 1961, incubation had begun. The birds had arrived some
three weeks earlier, according to Smith. Although arriving from
the south long after most raptors have begun their nesting, the
kites are not further delayed by establishment of territories and
choosing of mates, and nesting is underway soon after their arrival.
512 UNIVERSITY OF KANSAS Pusts., Mus. Nar. Hist.
According to Sutton (1939:45) the nest-building is an exceedingly
leisurely process. In the first two weeks after their arrival he ob-
served that the kites only occasionally bring a twig to the nest,
usually repairing last year’s structure rather than starting a new one.
Sutton recorded egg-laying on May 17 and 18 and hatching on
June 18 in northwestern Oklahoma, and the timing of these events
must be similar in Meade County, Kansas.
Shortly before sunset on June 1 a pair was observed at close
range from a parked automobile as the kites perched on roadside
fence posts about 50 feet apart at the Park boundary. At this
time the birds lacked their usual restlessness and were perching
quietly, neither preening nor attempting to find prey. With no
preliminaries the male flew to the female and lit on her back to
copulate. The female was receptive but did not crouch in a hori-
zontal position. The mounting lasted for approximately a minute.
During the first 30 seconds the male was fully occupied with bal-
ancing and positioning himself, and copulation occurred only during
the latter half of the mounting. During this interval cloacal contact
was effected three times, but was only momentary each time. The
birds were silent. After the male left, the female continued to
perch until flushed by my movements.
Judging from the nests that were examined, the kites of the
Meade Park area are well synchronized in their nesting, as all
arrive at approximately the same time. Bent (1936:66) stated that
if a kite’s nest is robbed, the birds will lay a second set, either in
the old nest or a new one, about two weeks later. All young seen
at Meade State Park seemed to represent an age range of consider-
ably less than two weeks, and, presumably, no renestings were
involved.
Nests were variable in size. Some were remarkably small in
relation to size of the kites, and would scarcely have been credited
to this species, had not the kites been seen sitting on them. Nests
were from 10 to 18 (average 14) inches long and from 10 to 14
(average 11.7) inches wide, in forks or crotches of branches. The
branches supporting the nests were from 1% to 10 inches in diameter.
The nests were constructed of twigs of approximately pencil size.
Of 87 nests at the Park, 29 were in cottonwoods, six were in willows,
and two were in elms. The figures probably reflect the relative
numbers of each of these species of tree rather than any clear-cut
preference of the kites. By the time nesting has begun the trees
have leafed out, and the nests are well concealed.
Mississippi Kite 513
At the time of my visit to the Park, July 18 to 22, nestlings were
well grown, and were beginning to feather out. On August 4 and
5 the young were well feathered, but flight feathers were not fully
grown and the young remained in the nest or perched on nearby
branches. On August 21 to 24 the young were fully fledged, and
were able to fly strongly but they still spent most of their time
perching and those of a brood tended to stay near together, usually
in the nest tree.
In a total of 26% hours of observation, 148 feedings were observed
—on the average one per 10.7 minutes. The interval changed from
an average of 12.8 minutes for 62 feedings on July 19 to 21, to 8.5
minutes for 59 feedings on August 4, and to 10.8 minutes for 27
feedings on August 21. The longer interval on July 19 to 21 may
have resulted from the greater furtiveness of the adult kites at this
stage in their nesting cycle. Nests usually were watched through
field glasses at distances of 50 to 100 feet. Ordinarily kites are not
disturbed by the presence of a person at these distances, but when
delivering food to the nest they seemed somewhat distracted and
sometimes stopped only momentarily then left, still carrying the
food. Usually they swooped at the observer when leaving; rarely
they swooped at him as they approached the nest. All observations
were between 10 a. m. and 5 p. m., and there was no obvious trend
according to time. Earlier and later in the day the rate of delivery
is probably less. The kites are notably late risers, and their activity
increases gradually after sunrise; in late afternoon activity tapers
off again. In 89 feedings, the average visit to the nest lasted 51
seconds but this average included a few relatively long stops, up to
four minutes in length, and 60 per cent of the visits were for inter-
vals of 30 seconds or less.
Insects often protruded from the bills of the adult kites delivering
food, but most of the food was carried in the throat. Sometimes
the gorge was much distended, although nothing protruded from
the mouth. The adult upon alighting sometimes would pass food
to the nestling, and sometimes would disgorge a mass of food in
the nest in front of the nestling. When the young were small, the
adult after having disgorged a food mass, remained to pick up the
food, bit by bit, and place it in the mouth of the nestling. However,
after the young were partly feathered out the adult merely left the
food for them. The nestling sometimes would peck at the dis-
gorged material for several minutes ~ “ter the adult left before all
of the food was eaten.
514 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
The small nestlings are generally silent, but when handled or
otherwise disturbed, they give soft lisping peeps. By early August,
when the young have ventured from the nest bowl to nearby
branches, they become vocal and their calls can be heard more
often than those of the adults. The call of the adult has been well
rendered by Sutton (1939:43) with the syllables “phee phew’—a
whistle in which the first syllable is short (lasting only about one-
fourth of a second) with a rising inflection, clipped off short, while
the second syllable has a downward inflection, and is drawn out to
two or three times the length of the first syllable. The call of the
fledgling is soft, with a lisping quality; that of the adult is much
like it but is sharper and more piercing. Fledglings call frequently
while waiting to be fed, but as an adult approaches with food, the
calls are given in rapid succession and slurred to a high thin
squablike sqeaking or squealing.
When fledglings are able to fly and have left the nest, the adults
generally pass food to them directly, rather than dropping the
regurgitated mass, which might fall to the ground and be lost. On
August 22 a fledgling was seen following an adult in flight, and was
also seen to eat while it was flying. At this stage, when an adult
fed one young of a brood, the other would sometimes fly to the
spot in an attempt to share the meal. However, the transfer of
food was usually rapid and the adult would leave within a few
seconds. Young often were seen to fly out from the nest tree and
maneuver in the vicinity, flying in a roughly circular course perhaps
100 feet in diameter and then returning to the nest tree, thereby
familiarizing themselves with their surroundings.
According to the consensus of published accounts, there are
usually two eggs per clutch, occasionally one or three. However,
Ganier (1902:89), who studied the species in Mississippi, wrote:
“Of all the nests I have examined [number unspecified] only one
was found to contain more than a single egg.” Nice (1931:69)
recorded 19 sets of two each and seven of one each in Oklahoma.
In the course of my observations, 12 clutches of two were recorded.
A group of four fledglings were observed concentrating their activi-
ties at a nest more than 200 feet from any other known nests;
possibly all belonged to the same brood, but this was not definitely
determined.
Many of the nests that were in use in 1961 appeared to be relics
from earlier years, as the material was darkened and disintegrating,
but probably a new layer of sticks had been added on the top.
MIssIssrpPI KITE 515
Bent (op. cit.:65) mentioned this kite’s habit of frequently using
the same nest in successive years. On one occasion as I drove over
a little-used road in the Park and passed a cottonwood grove where
kites were nesting, one of the birds swooped down and struck the
top of the automobile. In a subsequent conversation, Harry Smith
asked me if this had happened, and said that this particular kite
had struck his truck frequently when he drove past its nest. This
had occurred at the same place in three successive years, and Smith
was convinced that the same kite had used the nest each year,
although the bird was not recognizable except by its unusually
aggressive behavior. On dozens of occasions in the course of my
observations kites swooped at me when I was near their nests,
but, except for this one individual, they always veered away at a
distance of several feet or several yards.
At the time of my visit to the Park in early June, kites were
relatively silent and secretive in their behavior. Approximately half
of those that were incubating flushed when a person walked near
the tree, but others continued to sit on their eggs until a person
had climbed to within a few feet of the nest. Upon being flushed,
such a kite, in 50 per cent of observed instances, swooped at least
once at the intruder, but some of the kites would soar overhead,
watching without making any active defense. At the time of my
next visit, July 18 to 21, when the kites were feeding well grown
nestlings, behavior at the nest was much different. As soon as a
nest was located the parents began scolding and swooping. At
the first nest observed, a group of eight kites had congregated
within two minutes to scold and harass the intruders. Even kites ~
whose nests were kept under observation frequently, never became
fully reconciled to the intrusion but there was much difference
between individuals in this respect. Some were reluctant to deliver
food and, having secured prey, would fly about in the vicinity
without coming to the nest.
Mortality Factors and Defense
Joint defense against a common enemy was noted on July 21,
1961, when 21 kites were seen swooping at a Swainson’s hawk
perched near the top of a large cottonwood, where it was partly
protected by foliage and branches. When I flushed the hawk, it
was pursued and harrassed by the kites, some of which followed
it for nearly a quarter mile although there were no nests of the
kites nearby. On August 4 a group of six kites was seen heckling
516 UNIVERSITY OF KANSAS PuUBLS., Mus. Nat. Hist.
a fledgling Swainson’s hawk, which crouched among thick foliage
in the top of a tall cottonwood, as the kites swooped at it, sometimes
brushing it with their wings when they swept past. Dr. Hibbard
mentioned an instance in which a horned owl was flushed, and was
chased and heckled by a red-tailed hawk and by a group of kites.
The latter seemed to regard the owl as the greater enemy, but
ordinarily any large raptor arouses their hostility.
Because of their exceptionally swift and skillful flight, the adult
kites have few natural enemies, but the eggs or nestlings are vul-
nerable to such enemies as crows, jays, the larger hawks and owls,
and to certain mammalian predators, notably raccoons. Also, many
nests probably are destroyed by the sudden and violent summer
storms that are characteristic of the High Plains. Bendire (1892:
178) cited observations by Goss that in a hailstorm in Barber
County, Kansas, eggs were destroyed in many kites’ nests and some
of the nests were almost completely demolished. Several nests
found by me to have incubating eggs in the first week of June were
abandoned or had disappeared completely by July 18, but the
cause was not evident. One nest that was under observation on
July 22 had nestlings approximately two-thirds grown on that date,
but on August 4 only a few sticks remained, and the carcass of a
fledgling dangled from a limb ten feet below the nest. Even at
the Park where firearms are prohibited, kites are sometimes shot
by ignorant or malicious persons. In general, Kansas ranchers
recognize the harmless and beneficial habits of kites, appreciate
their esthetic appeal and protect them, but many persons use them
as convenient targets, with utter disregard for the Federal laws
protecting them. Because of the strong popular prejudice against
raptorial birds in general, laws protecting them are usually not
enforced. Law enforcement officers do not take action even when
clear-cut violations come to their attention. Arrest and prosecution
for the killing of any kind of raptor is almost out of the question
in Kansas.
Ratio of Immatures to Adults
In the juvenal plumage flight feathers of the kites are brown,
barred with white, much different in appearance from the dark,
slaty plumage of adults. Bent (op. cit.:67) stated that these barred
flight feathers are retained through the second summer, and he
quoted Mr. G. W. Stevens as having found kites breeding in this
immature plumage. On June 2, 1961, I attempted to determine
the ratio of these yearling kites to others in the population at the
MISSISSIPPI KITE tay h7/
Park. Most of the kites seen were in flight too far away to discern
definitely whether or not they were juveniles, and records were
limited to those seen at relatively close range. In a total of 108
records only 11 pertained to these yearlings and the remaining 97
were identified as of adults. Beyond doubt in the course of my
counts some individuals were recorded repeatedly, therefore the
counts are not entirely acceptable. However, on each occasion that
kites were seen in numbers in early summer, the adults greatly out-
numbered the juveniles. The approximate nine to one ratio of
adults to yearlings seems much too high. Even if the difference
is much less than indicated, the high ratio of adults to yearlings
would seem to imply that the adults have a long life expectancy.
A rather improbable alternative is that some of the yearlings remain
in winter quarters or wander elsewhere rather than accompanying
the adults on the return migration to their breeding grounds. Still
another alternative is that the breeding season of 1960 was relatively
unsuccessful, but this idea is negated by my own observations at
the Park in late 1960, as recently fledged young were numerous then.
At the time of my visit to the Park August 21 to 24, 1961, all young
had recently left the nests and were able to fly. However, their
behavior was so much different from that of the adults that a
reliable ratio could not be obtained. The fledglings tended to re-
main in the nest tree, or to make relatively short flights near it,
while the adults occupied with catching of prey for themselves and
their young, spent much of their time aloft. The adults were hence
far more conspicuous than the fledglings. However, it is my im-
pression that the fledglings were from one-third to one-fourth as
numerous as the adults. If this ratio is correct, and if all adults had
bred, from two-thirds to three-fourths of the eggs and/or nestlings
must have been destroyed. This rate of loss seems reasonable in
view of the known histories of nests observed in June and again
in July, and of the fates of birds’ nests in general.
Summary
Mississippi kites were studied in southwestern Kansas in the
summer of 1961, at various localities, especially at Meade State
Park. At this locality, near the northwestern limit of the breeding
range, the kite thrives in typical High Plains habitat dominated by
a short-grass type of vegetation, but availability of trees suitable
for nests is a limiting factor. Since maturing of extensive groves
of cottonwoods and other trees planted at Meade State Park, the
518 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
colony of kites has increased tremendously and the breeding popu-
lation probably exceeded 100 in 1961.
The kites are social in all their activities and do not maintain
territories. The sexes differ little in appearance, but males are
slightly smaller than females and have paler heads. Food consists
almost entirely of flying insects, and these are usually eaten while
the kite is in flight. Kites that are feeding nestlings may travel up
to two miles from the nest or perhaps considerably farther in the
course of their foraging. For 148 feedings of nestlings the observed
intervals averaged 10.7 minutes. Most published references to the
food habits mention predation on small vertebrates, especially
lizards, but including also snakes, toads, rodents, and even rabbits.
In my study a total of 205 pellets were collected and 453 insects
were tentatively identified but the total number of insects in the
pellets was much larger. No vertebrates were identified from this
sample and among 358 prey items identified from kite stomachs
collected in Oklahoma, by Sutton, vertebrae of a small fish were
the only vertebrate remains. Further verification of predation on
mammals, reptiles and amphibians by this species is needed. Of
the insects distinguished in pellets, beetles including carabids, ci-
cindelids, hydrophilids, scarabaeids, and silphids were most numer-
ous (270) and grasshoppers (164) were second; also there were
16 cicadas and three moths.
Kites arrive in Kansas about the second week in May. Often
old nests are repaired and used over again. Hatching is about
mid-June. Normally there are two eggs per clutch. By mid-August
the fledglings are learning to fly. By the latter part of August they
are learning to capture their insect prey, and in early September
southward movement of the entire population begins.
Eggs and/or young in many nests are destroyed by hail or high
wind in the sudden violent storms that are characteristic of the
High Plains. Mississippi kites are often shot by misguided persons,
and benefit little from the protection supposedly provided by Fed-
eral law. However, the adults probably have few natural enemies.
The high ratio of older adults to yearlings indicates that the life
expectancy is long. Through their second summer the kites retain
their barred immature plumage, and can be readily distinguished
from adults. Only ten per cent of the kites recorded in 108 June
sight records at the Park were in juvenile plumage.
MIssIssIPpP!I KITE 519
Literature Cited
AUDUBON, J. J.
1840. The birds of America. Philadelphia, pp. xv + 246.
BENDIRE, C. E.
1892. Life histories of North American birds. U.S. National Mus. Spec.
Bull. 1, viii + 446 pp.
BEnT, A. C.
1937. Life histories of North American birds of prey. Bull. U. S. Nat.
Mus., 167, x + 409 pp. 102 pls.
CHAPMAN, F. M.
1891. On the birds observed near Corpus Christi, Texas, during parts of
March and April, 1891. Bull. Amer. Mus. Nat. Hist., 3:315-328.
EISENMANN, E.
1963. Mississippi kite in Argentina, with comments on migration and
plumage in the genus Ictinia, Auk, 80:74-77.
Ganiep, A. F.
1902. The Mississippi kite (Ictinia mississippiensis). The Osprey, vol. 1
(new series ), No. 6:85-90.
Goss, N. S.
1891. History of the birds of Kansas. Geo. W. Crane and Co., Topeka,
692 pp.
Nice, M. M.
1931. The birds of Oklahoma (rev.). Publ. Univ. Oklahoma, vol. 3,
Biol. Surv. No. 1, 261 pp.
Sutton, G. M.
1939. The Mississippi kite in spring. Condor, 41(2):41-52.
Wayne, A. T.
1910. Birds of South Carolina. Contr. Charleston Mus., No. 1, viii +
254 pp. The Daggett Printing Co., Charleston, S. C.
Transmitted June 8, 1963.
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UNIVERSITY OF KANSAS PUBLICATIONS
MusEuM OF NaTURAL HISTORY
Volume 12, No. 12, pp. 521-551, 22 figs.
October 25, 1963
Jaw Musculature
Of the Mourning and White-winged Doves
15356
ROBERT L. MERZ
UNIVERSITY OF KANSAS
LAWRENCE
1963
UNIVERSITY OF KANSAS PUBLICATIONS, MuSEUM OF NATURAL HisToRY
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 12, No. 12, pp. 521-551, 22 figs.
Published October 25, 1963
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY
JEAN M. NEIBARGER, STATE PRINTER
TOPEKA, KANSAS
1963
C23
9-7865
67
Jaw Musculature
Of the Mourning and White-winged Doves
BY
ROBERT L. MERZ
For some time many investigators have thought that the genus
Zenaida, which includes the White-winged and Zenaida doves, and
the genus Zenaidura, which includes the Mourning, Eared, and
Socorro doves (Peters, 1937:83-88), are closely related, perhaps
more closely than is indicated by separating the several species
into two genera. It is the purpose of this paper to report investi-
gations on the musculature of the jaw of doves with the hope that,
together with the results of other studies, the relationships of the
genera Zenaida and Zenaidura can be elucidated.
METHODS AND MATERIALS
In order to determine in each species the normal pattern of musculature
of the jaws, heads of 18 specimens of doves were dissected (all material is in
the Museum of Natural History of The University of Kansas): White-winged
Doves (Zenaida asiatica), 40323, 40324, 40328, 40392, 40393; Zenaida Doves
(Z. aurita), 40399, 40400; Mourning Doves (Zenaidura macroura), 40326,
40394, 40395, 40396, 40397, 40398.
Thirty-seven skulls from the collection of the Museum of Natural History
of The University of Kansas and two skulls from the United States National
Museum were measured. The measurements are on file in the Library of The
University of Kansas in a dissertation deposited there by me in 1963 in partial
fulfillment of requirements for the degree of Master of Arts in Zoology. Speci-
mens used were: White-winged Doves, KU 19141, 19142, 19148, 19144,
19145, 19146, 19147, 23188, 23139, 24387, 24889, 24841, 23592, 23593,
24840, 31025, 31276; Mourning Doves, KU 14018, 14781, 15347, 15533,
15547, 15550, 15662, 15778, 15872, 16466, 17782, 17786, 17788, 17795,
19153, 19242, 20321, 21669, 22394, 22715; Eared Doves (Zenaidura auricu-
lata), USNM 227496, 318381. Additionally, the skulls of the Zenaida Doves
mentioned above were measured. All measurements were made with a dial
caliper and read to tenths of a millimeter.
ACKNOWLEDGMENTS
My appreciation is extended to Professor Richard F. Johnston, who advised
me during the course of this study, and to Professors A. Byron Leonard and
Theodore H. Eaton for critically reading the manuscript.
I would like also to acknowledge the assistance of Dr, Robert M. Mengel
and Mr. Jon C. Barlow for suggestions on procedure, and Mr. William C.
Stanley, who contributed specimens of Mourning Doves for study. Mr. Thomas
(523)
gener ai Co
UNIVERSITY
524 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
H. Swearingen offered considerable advice on production of drawings and
Professor E. Raymond Hall suggested the proper layout of the same and gave
editorial assistance otherwise, as also did Professor Johnston.
MYOLOGY
The jaw musculature of doves is not an imposing system. The
eating habits impose no considerable stress on the muscles; the
mandibles are not used for crushing seeds, spearing, drilling, gaping,
or probing as are the mandibles of many other kinds of birds.
Doves use their mandibles to procure loose seeds and grains, which
constitute the major part of their diet (Leopold, 1943; Kiel and
Harris, 1956: 877; Knappen, 1938; Jackson, 1941), and to gather
twigs for construction of nests. Both activities require but limited
gripping action of mandibles. The crushing habit of a bird such as
the Hawfinch (Coccothraustes coccothraustes), on the other hand,
involves extremely powerful gripping (see, for example, Sims,
1955 ); the contrast is apparent in the development of the jaw muscu-
lature in the two types. Consequently, it is not surprising to find a
relatively weak muscle mass in the jaw of doves, and because the
musculature is weak there are few pronounced osseous fossae,
cristae and tubercles. As a result, the bones, in addition to being
small in absolute size, are relatively weaker when compared to
skulls of birds having more distinctive feeding habits which require
more powerful musculature.
The jaw muscles of the species dissected for this study are, in
gross form, nearly identical from one species to another. Thus, a
description of the pertinent myology of each species is unnecessary;
one basic description is hereby furnished, with remarks on the
variability observed between the species.
The terminology adopted by me for the jaw musculature is in
boldfaced italic type. Synonyms are in italic type and are the
names most often used by several other writers.
M. pterygoideus ventralis: part of Mm. pterygoidei, Gadow, 1891:323-325,
table 26, figs. 1, 2, 3 and 4, and table 27, fig. 3—part of M. pterygoideus
internus, Shufeldt, 1890:20, figs. 8, 5, 6, 7 and 11—part of M. adductor
mandibulae internus, Edgeworth, 1935:58, figs. 605c and 607—part of
M. pterygoideus anterior, Adams, 1919:101, pl. 8, figs. 2 and 8.
M. pterygoideus dorsalis: part of Mm. pterygoidei, Gadow, 1891:823-325,
table 26, fig. 7 and table 27, figs. 1 and 3—part of M. pterygoideus
internus, Shufeldt, 1890:20—-part of M. adductor mandibulae internus,
Edgeworth, 1935:58, fig. 605c—? part of M. pterygoideus anterior,
Adams, 1919:101, pl. 8, figs. 2 and 3.
M. adductor mandibulae externus: a) pars superficialis: parts 1 and 2 of
M. temporalis, Gadow, 1891:320-321—part of M. temporal, Shufeldt,
1890:16, figs. 5 and 7—part of M. adductor mandibulae externus, Edge-
Jaw MuscuLATuRE OF Doves 525
worth, 1935:58-60—-M. capiti-mandibularis medius and profundus,
Adams, 1919:101, pl. 8, fig. 1.
b) pars medialis: ? parts 1, 2 and 3 of M. temporalis, Gadow, 1891:320-
322—part of M. masseter and ? part of M. temporal, Shufeldt, 1890:
16-18, figs. 5, 6, 7 and 11—part of M. adductor mandibulae externus,
Edgeworth, 1935:58-60—M. capiti-mandibularis superficialis, first part,
Adams, 1919:100-101, pl. 8, fig. 1.
c) pars profundus: part 2 of M. temporalis, Gadow, 1891:321, table 27,
fig. 2—part of M. temporal and ? part of M. masseter, Shufeldt, 1890:
16-18—part of M. adductor mandibulae externus, Edgeworth, 1935:
58-60—? part of M. capiti-mandibularis medius and all of pars super-
ficialis, second part, Adams, 1919:100-101.
M. pseudotemporalis profundus: M. quadrato-maxillaris, Gadow, 1891:
322-328—M. pterygoideus externus, Shufeldt, 1890:20-21, figs. 3, 5 and
11—part of M. adductor mandibulae medius, Edgeworth, 1935:58-59—
? part of M. pterygoideus posterior, Adams, 1919:101, pl. 8, figs. 2 and 8.
M. protractor pterygoidei: part 4b of M. temporalis, Gadow, 1891: 322-
323, table 27, fig. 4—part of M. entotympanious, Shufeldt, 1890:19-20,
figs. 8 and 11—part of M. spheno-pterygo-quadratus, Edgeworth, 1935:
oT
M. depressor mandibulae: M. digastricus s. depressor mandibulae, Gadow,
oe biventer maxillae, Shufeldt, 1890:18-19, figs. 8, 4,
5, 6, 7 and 11.
M. pseudotemporalis superficialis: M. spheno-maxillaris, Gadow, .1891:323
—part of M. temporal, Shufeldt, 1890:16—part of M. pseudotemporalis,
Hofer, 1950:468-477—part of M. adductor mandibulae medius, Edge-
worth, 1935:277.
M. adductor mandibulae posterior: ? part of M. temporal, Shufeldt, 1890:
16—part of M. adductor mandibulae medius, Edgeworth, 1935:58-59—
? part of M. pterygoideus posterior, Adams, 1919:101, pl. 8, figs. 2 and 8.
M. protractor quadrati: part 4a of M. temporalis, Gadow, 1891:322-323,
table 27, fig. 4—part of M. entotympanicus, Shufeldt, 1890:19-20, figs. 8
and 11—part of M. spheno-pterygo-quadratus, Edgeworth, 1935:57.
The terminology adopted by me is that of Lakjar (1926) except that the
divisions of M. depressor mandibulae are designated by the Latinized equiva-
lents of the names used by Rooth (1953:261-262).
M. pterygoideus ventralis lateralis—The origin is fleshy and by aponeurosis
on the ventral side of the palatine anterior to the palatine fossa. The insertion
is fleshy on the ventromedial surface of the lower mandible and continues
along the anteromedial surface of the internal angular process to its distal tip.
A few fibers leave pars lateralis and insert on an aponeurosis which receives
also all the fibers of M. pterygoideus dorsalis lateralis. The latter fact may
have prompted Rooth (1953:257) to make the statement that the fibers
originating on the dorsal part of the palatine inserted more laterally than those
originating on the ventral side. Rooth worked with Columba palumbus, the
Woodpigeon, and his description concerned M. adductor mandibulae internus
pterygoideus, which is composed of Mm. pterygoideus ventralis et dorsalis
of Lakjar (1926). His assertion that ventral fibers, that is to say, fibers arising
on the ventral surface of the palatine, insert medially does not appear to be
completely true for doves.
Aponeuroses cover most of the lower surface of the muscle and one or two
nerves extend into the substance of the muscle. The nerves run from the
526 University OF Kansas Pusuxs., Mus. Nat. Hist.
anterior edge of M. pterygoideus dorsalis medialis and farther posteriorly from
a separation in the muscle.
M. pterygoideus ventralis medialis—The origin is by aponeurosis from
the ventral surface of the palatine and fleshy from the palatine fossa. The
aponeurosis is the same that gives origin to the fibers of pars lateralis. Part
of the aponeurosis becomes tendonlike in the middle of M. pterygoideus ven-
tralis and separates its two divisions. The insertion is fleshy on the lower
one-third of the anterior surface of the internal angular process of the lower
mandible, and by two tendons on the distal tip of that process. Many of the
fibers of pars medialis insert on the tendons. The fibers at their insertion are
not distinctly separate from those of pars lateralis and there is considerable
mingling of the fibers. Consequently, the medial part of M. pterygoideus
ventralis cannot be removed as a part distinct from the lateral part (figs. 1,
4, 10, 21 and 22).
Ordinarily M. pterygoideus ventralis does not cross the ventral edge of the
lower mandible, but in one white-wing the muscle was slightly expanded on
the right side and it could be seen in lateral view. The homologous muscle
in Columba palumbus apparently is consistently visible in lateral view. (See
Rooth, 1953, fig. 6.)
M. pterygoideus dorsalis medialis—The origin is fleshy on the dorsolateral
surface of the palatine immediately anterior to the pterygoid and also on the
anterior, dorsolateral, posterior and ventromedial surfaces of the pterygoid.
The insertion is fleshy on the ventromedial surface of the lower mandible and
the anterior surface of the internal angular process immediately dorsal to the
insertion of M. pterygoideus ventralis lateralis.
M. pterygoideus dorsalis lateralis—The origin is fleshy from the dorso-
lateral surface of the palatine, anterior to the origin of pars medialis and the
insertion is by means of an aponeurosis on the medial surface of the lower
mandible, lateral to the insertion of M. pterygoideus ventralis lateralis. The
aponeurosis crosses the medial side of the insertion of M. pterygoideus dorsalis
medialis. The fibers run in a posteroventrolateral direction and insert on the
ventromedial side of the aponeurosis (figs. 1, 6, 8, 9, 18-22).
In one individual, a Mourning Dove, the origin of pars lateralis of M. ptery-
goideus dorsalis extended to the pterygoid. With this one exception the
muscle was uniform throughout the several species.
M. adductor mandibulae externus—This is the most complex muscle in
the jaw owing to its system of tendons and aponeuroses. Three divisions of
this muscle were described by Lakjar (1926:45-46) and the divisions appear
to be distinguishable in the doves, but there is no clear line of demarcation
for any of the parts and the following description is based upon my own at-
tempts to delineate the muscle.
M. adductor mandibulae externus superficialis—The origin is fleshy from
the most lateral area of the temporal fossa. Dorsally the origin is bounded by
the base of the postorbital process and ventrally by the temporal process. The
fibers converge upon a tendon that passes beneath the postorbital ligament
and runs anteriorly among the fibers of pars profundus. The insertion is
tendinous on the dorsal surface of the lower mandible in common with the
dorsal aponeurosis of pars profundus. The insertion is immediately anterior
Jaw MuscuLatureE OF Doves 527
to the ventral aponeurosis of pars profundus near the medial edge of the
dorsal surface on a tubercle at the posterior end of the dorsal ridge of the
lower mandible.
M. adducior mandibulae externus medialis—The origin is by a flat, heavy
tendon from the temporal process. The tendon is attached almost vertically
on the temporal process. It twists approximately 180° as it runs anteriorly,
and becomes a thin aponeurosis, which gives rise on its dorsal and ventral
surfaces to many fibers that insert in a fan-shaped area on the mandibular
fossa. Fibers from the dorsal and dorsomedial sides of the heavy tendon run
rostrad and insert on the ventral surface of the dorsal aponeurosis of pars
profundus. From the ventral surface the most posterior fibers converge on
an aponeurosis that inserts on a transverse crista on the dorsal surface of the
mandible immediately lateral to the ventral aponeurosis of pars profundus and
dorsal to the insertion of M. adductor mandibulae posterior. The more an-
terior fibers insert fleshily on the mandibular fossa. The tendon of origin is
actually one with the ventral aponeurosis of pars profundus, which is situated
in a horizontal plane. The insertion is primarily a fleshy attachment on the
mandibular fossa. Some of the fibers that arise on the dorsomedial and
lateral surfaces of the tendon of origin attach to another tendon, which inserts
in the midline of the mandibular fossa on a small tubercle near the anterior
end. Also, there is insertion by an aponeurosis anterior to M. adductor man-
dibular posterior as stated above. Fibers attach to the dorsal and ventral side
of the aponeurosis.
M. adductor mandibulae externus profundus—The origin is fleshy from
the medial surface of the temporal fossa, the posterior wall of the orbit and
the otic process of the quadrate. The origin is bounded laterally by the
origin of pars superficialis and medially by the origin of M. pseudotemporalis
superficialis. Ventrally the muscle lies against its own ventral aponeurosis,
which originates on the posterior wall of the orbit immediately above the articu-
lation of the otic process of the quadrate, and which also receives many fibers
from the surface of the quadrate. The insertion is primarily by means of
two aponeuroses. The most dorsal aponeurosis inserts on a tubercle at the
posterior tip of the dorsal edge of the mandible. The lateral tendon of
M. pseudotemporalis superficialis converges with the aponeurosis. It is super-
ficial and there are no fibers on its dorsal surface. The ventral aponeurosis
inserts on a crista immediately below the insertion of the dorsal aponeurosis.
It receives fibers on its ventral surface from the otic process of the quadrate,
and on its dorsal surface gives rise to fibers that insert on the dorsal apo-
neurosis (figs. 2, 3, 5, 9, 10, 11, 18-18).
The tendon of insertion of pars medialis of M. adductor mandibulae externus
does not become a superficial aponeurosis posteriorly in the Zenaida Dove as
it does in the Mourning and White-winged doves.
M. pseudotemporalis profundus.—The origin is fleshy from the medial and
partially from the dorsal surface of the lower mandible. The origin is almost
completely anterior to and partly dorsal and ventral to the medial (most
anterior) insertion of M. pseudotemporalis superficialis. The anterior margin
of the origin is at the point where the mandibular ramus of the trigeminal
nerve enters the mandible. Posteriorly the origin is bounded by the insertion
528 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
of M. adductor mandibulae posterior, and ventrally by a ridge that is situated
about halfway down the medial side of the mandible. The insertion is by
aponeurosis on the tip of the orbital process of the quadrate and fleshily on
the anterior surface of the same process. ‘The aponeurosis extends about
three-fifths of the distance along the muscle and it is dorsal or superficial
to all of the fibers. Many fibers insert on the ventral side of the aponeurosis
(figs. 1, 5, 18, 14, 15, 16, 21 and 22).
This muscle is the most variable of all the jaw muscles. In the Mourning
Dove the muscle appears rather slender in dorsal view and in the White-
winged Dove has an enlarged lateral belly that gives the appearance of a
thicker muscle. In the Zenaida Dove M. pseudotemporalis profundus is inter-
mediate in shape between those of the other two species. This muscle will be
discussed in detail later.
M. protractor pterygoidei.—The origin is fleshy from the junction of the
sphenoidal rostrum and the interorbital septum. Fibers converge on the
pterygoid in anteroventrolateral and posteroventrolateral directions. The pos-
terior edge of the muscle is in contact with M. protractor quadrati with which
its fibers mingle. The insertion is fleshy on the posterior surface of the lateral
half of the pterygoid to its articulation with the body of the quadrate (figs.
6, 8, 9, 11, 18-20).
M. depressor mandibulae superficialis medialis—The origin is fleshy from
the lateral edge of the basioccipital where the muscle is attached to Liga-
mentum depressor mandibulae and extends in a lateral direction to a point
where the structures involved turn dorsad. The insertion is by fibers and a
light aponeurosis on the crista that is situated on the posteroventromedial edge
of the lower mandible.
M. depressor mandibulae superficialis lateralis—The origin is fleshy from
the squamosal region, slightly posteroventral to the origin of M. adductor man-
dibulae externus superficialis. A thin aponeurosis lies medial to the muscle
fibers. The insertion is by means of an aponeurosis that becomes tendonlike
along the posteroventrolateral crista and the posteriormost part of the ventral
edge of the lower mandible.
M. depressor mandibulae medialis——The origin is fleshy from the lateral
and ventral surfaces of Ligamentum depressor mandibulae. The insertion is
fleshy on the posterior surface of the lower mandible, posterodorsal to the
insertions of partes superficialis medialis et lateralis (figs. 4, 9, 10, 13 and 14).
The parts of M. depressor mandibulae are difficult to distinguish from one
another because of considerable intermingling of fibers.
M. pseudotemporalis superficialis—The origin is fleshy from the posterior
wall of the orbit, dorsal to the foramen of the trigeminal nerve, lateral to the
origin of M. protractor quadrati and medial to M. adductor mandibulae ex-
ternus profundus. The insertion is by means of an aponeurosis that bifurcates
at the point of contact with the mandibular ramus of the trigeminal nerve,
which is at the level of the orbital process of the quadrate (except in the
Mourning Dove where the division is more anterior), and which inserts as two
tendons on the dorsomedial edge of the lower mandible posterior to the
insertion of M. pseudotemporalis profundus. The lateral tendon is superficial
to the dorsomedial edge of M. adductor mandibulae externus, and converges
with the aponeurosis of pars profundus of that muscle and inserts with it on
Jaw MuscuLaTurE OF Doves 529
a tubercle near the dorsomedial edge of the mandible anterior to the insertion
of M. adductor mandibulae posterior as mentioned before. The anterior half
of the medial tendon lies ventral to the lateral edge of M. pseudotemporalis
profundus and the mandibular ramus of the trigeminal nerve. All of the
fibers of the muscle insert on the posteroventral surface of the aponeurosis
before it divides. Part of M. pseudotemporalis profundus also lies ventral to
the medial tendon of M. pseudotemporalis superficialis and, in effect, the
tendon is imbedded in the substance of M. pseudotemporalis profundus as it
proceeds anteriorly. The trigeminal nerve leaves a slight impression on the
ventral surface of the muscle near its origin (figs. 1, 8, 11, 18, 14, 15 and 16).
M. adductor mandibulae posterior—The origin is fleshy from the antero-
dorsal and anterior surfaces of the quadrate body, from the anterodorsolateral,
medial and anterior surfaces of the orbital process of the quadrate. The
muscle also has an origin from the otic process of the quadrate, partly fleshy
and partly by a slight aponeurosis. The insertion is fleshy on the dorsal and
lateral surfaces of the mandible immediately anterior to the articulating
surface. This muscle also has extensive insertion on the medial side of the
lower mandible dorsal to the insertion of M. pterygoideus dorsalis medialis and
posterior to the origin of M. pseudotemporalis profundus (figs. 1, 8, 5, 17,
18, 19 and 20).
The fibers of M. pseudotemporalis profundus can be distinguished from
the fibers of M. adductor mandibulae posterior because the pterygoideus nerve
passes between the two (Lakjar, 1926:55). Rooth (1953:255-256) considers
as part of this muscle the ventral aponeurosis of pars profundus of M. adductor
mandibulae externus and all the fibers ventral to it. But I could not justify
the inclusion of that aponeurosis as part of M. adductor mandibulae posterior
in the doves because none of the fibers of M. adductor mandibulae posterior
as I have described it were attached to that particular aponeurosis.
M. protractor quadrati—tThe origin is fleshy from the posterior wall of
the orbit medial to the foramen of the trigeminal nerve and also medial to the
origin of M. pseudotemporalis superficialis. The origin describes an arc in
the horizontal plane until it reaches the interorbital septum and the optic
nerve. The insertion is fleshy on the posteromedial edge of the body of the
quadrate and the orbital process of the quadrate and on the otic process of
the quadrate. The muscle also inserts on the ventromedial surface of the
orbital process of the quadrate and the adjacent area of the body of the
quadrate (figs. 5, 7,9, 11, 18-18).
M. protractor quadrati possesses many fibers that arise from M. protractor
pterygoidei. Consequently, it is difficult to determine the exact extent of the
origin or the insertion of either muscle.
ACTION OF JAW MUSCLES
M. pterygoideus ventralis—Contraction of this muscle retracts the upper
mandible by moving the palatine posteriorly, and simultaneously adducts the
lower mandible.
M. pterygoideus dorsalis—This muscle functions in essentially the same
manner as M. pterygoideus ventralis. The result of having a part of its origin
on the pterygoid as well as on the palatine is to facilitate retraction of the
upper mandible.
530 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
M. adductor mandibulae.—This is the chief adductor of the lower mandi-
ble and the muscle functions solely in that capacity. In birds having great
crushing ability, this muscle is much larger and more powerful and the skull
is reinforced behind the quadrate in order to withstand the pressure of the
lower mandible against the quadrate during adduction (Sims, 1955:374 and
Bowman, 1961:219-222).
M. pseudotemporalis profundus.—With origin and insertion on _ highly
movable bones, this muscle, when it contracts, retracts the upper mandible
and adducts the lower mandible. Like the pterygoid muscles, this muscle,
by itself, would allow the bird to grasp objects by means of its mandibles.
However, M. pseudotemporalis profundus could produce a more powerful grip
because it takes origin farther anteriorly on the lower mandible.
M. protractor pterygoidei—Contraction of M. protractor pterygoidei pulls
the pterygoid anteromedially and causes it to slide forward along the sphenoidal
rostrum. This action aids in protraction of the upper mandible.
M. depressor mandibulae.—The depressor of the lower mandible is the
sole muscle other than M. geniohyoideus involved in the function of abducting
the lower jaw of doves. Its size can be correlated especially well with feeding
habits of the bird. Other birds that force their closed mandibles into fruit,
wood or the earth and then forcibly open them, belong to groups possessing
enlarged depressors. Contraction of the muscle pulls the postarticular (retro-
articular) process upward with the resultant downward movement of that
part of the mandible which is anterior to the articulation. Since there is no
“gaping” in doves the muscle is only large enough to overcome the inherent
tone of the relaxed adductor muscles.
In some non-passerine species as well as in certain passerines the muscle
also serves to raise the upper jaw by acting on the quadrate, which is capable
of rotating vertically on its otic process. Especially in the gapers, where
resistance is offered near the tip of the lower mandible, contraction of the
muscle pulls the entire mandible dorsad thus forcing the jugal and palatal struts
forward (Zusi, 1959:537-539). The action supplements that of Mm. pro-
tractor pterygoidei et quadrati and is enhanced by anterior migration of the
origin of M. depressor mandibulae.
There is no lifting action involved in contraction of the depressor muscle
in doves for two reasons—(A) the origin of the muscle is situated much too
far posteriorly on the skull, and, more important, (B) the quadrate is not
hinged for vertical movement. As will be discussed later, it moves only in
a horizontal plane.
M. pseudotemporalis superficialis—Like M. adductor mandibulae, this
muscle performs only the one function of adducting the lower mandible, and
like M. pseudotemporalis profundus it is a synergist of that muscle.
M. adductor mandibulae posterior—Although this muscle undoubtedly
acts as an adductor of the lower mandible, I believe that, because of its dis-
advantageous insertion so near the articulation, its main function must be
concerned with firming the mandible against the quadrate. This is to say
that its function is partially that of a ligament.
M. protractor quadrati—When M. protractor quadrati contracts, the quad-
rate bone is swung medially. This action, as mentioned previously, results in
jaw MuscuLaTurReE OF Doves 531
protraction of the upper jaw, and, as a consequence, its action supplements the
action of M. protractor pterygoidei.
CRANIAL OSTEOLOGY
The ability of most birds to protract the upper mandible, and
the structure of the skull which enables them to do so are responsible
for common reference to the skull as “kinetic” (Beecher, 1951a:412;
Fisher, 1955:175). The movement is effected by muscular action
on a series of movable bones that exert their forward force on the
upper mandible, which in turn swings on a horizontal hinge, the
“naso-frontal hinge,” at the base of the beak. The bone initiating
the movement is the quadrate, which is hinged posteriorly by its
otic process and which ordinarily swings up or down depending
on the muscle or muscles being contracted at any given moment.
The upward swing of the quadrate pushes the jugal bar, which is
attached to its lateral tip, along its longitudinal axis, in an antero-
dorsal direction, and the force is transferred to the upper mandible,
which is thereby elevated. A synergetic mechanism is simultane-
ously initiated by the same bone—the quadrate. Since the quadrate
body articulates with the pterygoid, the upward movement forces
the pterygoid to slide along a ridge in the ventral midline of the
cranium, the sphenoidal rostrum, thus pushing the palatine forward
and exerting an upward push on the upper mandible.
In the columbids the quadrate has a bifurcated otic process that
functions as the hinge. The posterior tips of the forks are situated
almost vertically (one above the other) and the movement of the
quadrate is not so much up and down, or vertical, as it is horizontal
(fig. 12). When the quadrate moves medially the upper mandible
is protracted; a lateral movement results in retraction. There is a
slight, almost negligible, upward movement of the quadrate. The
movements of the various bony elements were observed on a skull
that had been made flexible by boiling in water for a minute as
suggested by Beecher (1951a:412).
Also in the columbids the naso-frontal hinge is not constructed
in the same manner as it is in many other birds as there is not a
simple hinge across the entire base of the beak. In fact, there is
no true hinge at all in the area of the nasals, but those bones are
extremely thin and they bend or flex under pressure. Actually,
the hinge is double or divided. One part is on either side of the
nasals. The hinges are situated at the posterodorsal tips of two
thin processes of the maxillary bones and the appearance is not
unlike that of half a span of a suspension bridge having the hinges
532 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
at the tops of the towers. Several other species of birds share
this type of hinge construction with columbids.
The movement of the lower jaw is, of course, the primary opera-
tion involved in opening the mouth. The lower jaw possesses a
deep fossa at its posterior end, or on its posterodorsal surface,
which articulates with the body of the quadrate bone. The length
of that part of the mandible extending behind the articulation is
directly correlated with the resistance offered the mandible in
opening, since it is on the posterior extension that the depressor
of the lower mandible inserts. The larger the muscle the more
surface is needed for attachment. Also the added length of the
mandible posterior to the articulation serves as a lever in opening
the mandible, and the fulcrum is moved relatively farther forward.
In birds lacking resistance to abduction of the lower mandible, as
in doves, it is nevertheless necessary for a slight postarticular
process to remain for the insertion of a small depressor muscle
which, as mentioned previously, is necessary to counteract the re-
laxed adductor muscles of the lower jaw.
There are many exceptions to the rule that birds have kinetic
skulls, and usually a secondary fusion and reinforcement of bones
around the hinge has limited or eliminated all movement. Sims
(1955) describes the Hawfinch’s immobile upper jaw, which is
used as a powerful press in cracking the stones of fresh fruit.
Skulls of woodpeckers have been modified somewhat in the same
manner as a result of their foraging and nesting habits ( Burt, 1930).
The two most distantly related members of the genera under
investigation are the White-winged Dove, Zenaida asiatica, and the
Mourning Dove, Zenaidura macroura. They were chosen to demon-
strate differences and likenesses in proportions of members of the
genera.
Ten measurements were taken on each skull, but simple observa-
tion reveals that, in relation to total length of the skull, the beak of
the White-winged Dove is longer than that of the Mourning Dove.
Tip of upper mandible to base of beak averaged 48.6 and 42.9
per cent of the total length of the skull in the White-winged Dove
and Mourning Dove, respectively. The position of the jugal bar
has remained about the same with respect to the cranial part of
the skull, and the entire cranial part of the skull is almost the same
shape in the species studied.
Likewise, in the White-winged Dove the distance from the
anterior tip of the lower mandible to the anterior part of M. adduc-
Jaw MuscuLatTurE OF Doves 533
tor mandibulae externus is relatively longer in relation to the length
of the lower mandible than in the Mourning Dove. Finally, the
position of the jugal with respect to the naso-frontal hinge is about
the same in the two species.
Measurements and calculations indicate that the longer beak of
the White-winged Dove as compared with the Mourning Dove is
a function of the beak itself, not of differences in other parts of the
skull. Measurements of skulls of Eared and Zenaida doves support
this view.
OTHER MORPHOLOGICAL FEATURES
In the species dissected, the only variable muscle that I consider
significant in revealing relationships is M. pseudotemporalis pro-
fundus. It is markedly enlarged in the White-winged Dove in
relation to the homologous muscle in the Mourning Dove. The
muscle is enlarged in such a manner that a lateral expansion of its
mass is apparent in superficial or dorsal view (compare figures 15
and 16). This, of course, indicates a muscle with powerful con-
traction, which has been unable to enlarge its circumference sym-
metrically because the eye is immediately dorsal to the muscle.
Therefore it has expanded laterally. Ventral expansion is blocked
by the presence of other muscles, and medially there is no surface
for the insertion of additional fibers on the orbital process of the
quadrate.
The jaw musculature has been known for some time to be highly
adaptive (Beecher, 195la and b, 1953; Bowman, 1961; Burt, 1980;
Engels, 1940 and Goodman and Fisher, 1962) and it would not be
unreasonable, I think, to expect the jaw muscles of closely related
species with similar habits to be similar. The beak of the White-
winged Dove is longer in proportion to the length and height of
the skull (exclusive of the beak) than is the beak of the Mourning
Dove. The lengthened beak is probably an adaptation for nectar-
feeding, which has been documented by McGregor, Alcorn and
Olin (1962:263-264) while investigating pollinating agents of the
Saguaro Cactus (Cereus giganteus), and by Gilman (1911:53)
who observed the birds thrusting their bills into the flowers of the
plant. Gilman indicated, however, that he could not be sure if
the birds were seeking insects, pollen, or nectar. In any event
the lengthened bill probably facilitates getting food by birds that
probe parts of flowers. Hensley (1954:202) noted that both
Mourning and White-winged doves were “exceptionally fond of
534 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist.
this source of nourishment.” But he also points out an “interesting
correlation” between the presence of the white-wings in the desert
and the flowering of the saguaro. During his studies the appear-
ance of the first white-wing preceded the opening of the first
saguaro flower by two days. The flowering and fruiting season
lasted until August, the month of termination of the white-wing
breeding season.
Since Hensley makes the correlation solely with the white-wings,
I assume that there is no other obvious correlation between plants
and birds among the remainder of the avifauna of the desert.
Probably the Mourning Dove has failed to adapt to nectar-feeding
as yet, and the White-winged Dove is the primary exploiter of this
food niche. It should be noted, also, that the head of the Mourning
Dove is smaller than the white-wing’s, and perhaps there is no
need for an elongated beak for reaching deeply into the flowers.
The lengthened bill should produce no difficulties in protraction
of the upper mandible and depression of the lower for the reason
that in the dove there is no known resistance offered to these move-
ments. The genus Icterus furnishes an example wherein resistance
is met in the process of opening the mandibles; individuals of this
genus thrust their closed bill into certain fruits and forcibly open
their mandibles against the resistance of the pulp by strong pro-
traction and depression, thus permitting the juices of the fruit to
lake and ultimately to be consumed (Beecher, 1950:53). Beecher
refers to the technique used in fruit-eating as “gaping.” The result
of gaping in Icterus should be the presence of a more massive set
of muscles concerned with protraction and depression than is
found in non-gaping groups. Beecher found the situation to be
exactly as expected in that genus and in other genera which also
gape. Meadowlarks (Sturnella) and caciques (Archiplanus) gape
and pry in soil and wood respectively (Beecher, 1951a:422 and 426).
The lengthened beak would be a problem when the White-
winged Dove attempted to pick up objects such as seeds, which
do in fact constitute the largest percentage of its diet in spite of its
nectar-feeding habit. A similar situation exists in the genus Icterus,
which is primarily adapted for gaping even though it shows a
preference for insects when they are abundant (Beecher, 1950:53).
The lengthened beak could be compensated for by (A) migration
of the anterior end of the jugal bar toward the rostral tip of the
bill and away from the fronto-nasal hinge with a simultaneous
enlargement of the adductor muscles of the lower mandible, or
Jaw MuscuLaTuRE OF Doves 535
(B) enlargement of the one muscle that functions simultaneously
as an efficient retractor of the upper mandible and adductor of the
lower mandible, namely M. pseudotemporalis profundus. Mm.
pierygoideus dorsalis et lateralis perform the same function, but
because of their position on the lower mandible they, apparently,
are stronger retractors of the upper mandible than they are adduc-
tors of the lower.
It will be recalled that the jugal bar bears the same, or nearly
the same, relationship to the cranium in the white-wing as it does
in the Mourning Dove and that the heads, excluding the beaks of
both species, are of nearly the same proportions. Also, Mm. ad-
ductor mandibulae externus and pseudotemporalis superficialis, the
chief adductor muscles of the lower mandible, were not noticeably
enlarged in the white-wing. It is also important to note that other
combinations of migration of bone and/or enlargement of muscles
could successfully solve the problem of providing sufficient lever-
age for the proper functioning of the lengthened mandibles, but
it is my thesis that the second alternative sufficed for seed-eating
habits and that that is the adaptation that was established; it is,
in fact, the only one present in the White-winged Dove.
It is unlikely that this enlarged muscle and beak are the remains
of another series of jaw muscles that have converged toward the
condition in Mourning Doves. Columbids are almost unquestion-
ably monophyletic, and two lines would have had to diverge and
then converge. There is no evidence for such an evolutionary
occurrence.
GENERIC RELATIONSHIP
An attempt will be made here to summarize all the available
evidence, direct or indirect, which bears on the problem of relation-
ship of these genera. The original dissections which are discussed
in this report are only valuable as one more bit of evidence con-
cerning one characteristic that aids in clarification of generic rela-
tionship, and it is only in conjunction with other evidence that any
satisfactory conclusion may be forthcoming.
Morphology
My dissections demonstrated that, in relation to the size of the
doves, the jaw musculature of all the specimens investigated was
so nearly alike that only one major difference was detected. M.
pseudotemporalis profundus appeared to be enlarged in the White-
winged Dove. This might have been predicted, since the white-
536 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
wing was also shown to possess an elongated beak, presumably an
adaptation for nectar-feeding, which would necessitate additional
muscle development in order to compensate for the added length.
Measurements recorded from several skulls indicated that the
heads of the birds (excluding the beak) are nearly proportional.
Perhaps plumage patterns are the most widely used characters
for determining generic relationships of birds. Ridgway (1916:
339-385 ) followed the columbid classification of Salvadori (1893)
using plumage patterns and body proportions to distinguish be-
tween the genera. In the genus Zenaidura he included the unique
specimen Zenaidura yucatanensis, and he placed auriculata in
Zenaida. The White-winged Dove was referred to a separate genus,
Melopelia. He described the genus Zenaidura in the following
manner:
“Plumage of head, neck and under parts soft and blended; bare orbital
space moderate, broadest beneath eyes. Coloration plain, the proximal second-
aries (sometimes adjacent wing-coverts and scapulars also) spotted with
black; rectrices (except middle pair) with a black band across postmedian
portion, the apical portion paler gray than basal portion, sometimes white; a
small black subauricular spot; adult males with head, neck and anterior under
parts more or less vinaceous and sides of neck glossed with metallic purple.”
He noted that the plumage of Zenaida was almost precisely as
described for Zenaidura. Also, although all members of Zenaida
reputedly possessed twelve rectrices, a characteristic of the genus,
it was later found that auriculata possessed fourteen rectrices. The
species was promptly placed in the genus Zenaidura by Peters
(1934:213-215). In plumage and coloration, Melopia was described
as similar to Zenaida and Zenaidura but without black spots on the
wings.
The White-winged Dove also has twelve rectrices, but Bond
(1940:53) and Goodwin (1958:330-334) considered the number
and shape of rectrices to be of minor importance when compared
to the homologous markings of the plumage. Goodwin stated that
his conclusion was emphasized by the fact that the tail of auriculata
is intermediate in length and shape between those of macroura and
aurita. In summary Goodwin “lumped” the genera Zenaida and
Zenaidura under the genus Zenaida.
Nidification
It has been adequately documented that members of these
genera closely resemble one another in their nesting and egg-laying
habits. Bent (1932:407, 417), Davie (1889:157), Goss (1891:242)
Jaw MuscuLaturE OF Doves 537
and Nice (1922:466) have described the two, white eggs of the
clutch of the Mourning Dove. They have also noted that their
nests are composed mainly of twigs and may be constructed in
trees, shrubs or on the ground. The Eared Dove has nearly identi-
cal habits (Bond, 1961:104), and a similar situation exists with the
Zenaida Dove (Audubon, 1834:356; Bent, 1932:418-419).
Like the other species, White-winged Doves lay two white or
buffy eggs per clutch and build frail nests of sticks (Bent, 1932:
431; Wetmore, 1920:141; Baird, Brewer and Ridgway, 1905:877).
The point to be made here is simply this: If the species in
question are to be considered congeneric then it might reasonably
be expected that they would display some similarity in nidification
and egg-laying. If their habits varied considerably it would not
necessarily mean that their relationship was more distant, but
similarities can usually be considered indicative of affinities because
they are the phenotypic expression of the partially unaltered geno-
type of the common ancestor.
Interbreeding
Intergeneric crosses of columbids in captivity are common, but
in nature there is little evidence that even interspecific crosses
occur. Only one apparent hybrid between members of the genus
Zenaida and genus Zenaidura has ever been discovered. The indi-
vidual was taken on the Yucatan peninsula of Mexico, and was de-
scribed and named as a new species (Zenaidura yucatanensis).
Salvadori (1893:373), Ridgway (1916:353) and Peters (1934:
213-215) agree that Zenaidura yucatanensis Lawrence is a hybrid
between Zenaidura macroura marginella and Zenaida aurita yuca-
tanensis. Ridgway (1916:355), however, notes that“. . . If Zen-
aidura yucatanensis Lawrence should prove to be really a distinct
species, and not a hybrid . . . unquestionably Zenaida and
Zenaidura can not be separated generically, since the former is in
every way exactly intermediate between the two groups.” In the
event that the unique type is a hybrid, the very fact of its existence
supports the hypothesis that the genera are more closely related
than is currently recognized.
Serology
There have been no investigations having as their sole purpose
the clarification of the relationship of the genera Zenaida and Zen-
aidura. But some work has involved the comparison of the anti-
genic content of individual columbids with the antigenic content
of a member of another species of the same family.
538 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
Irwin and Miller (1961) tested, along with other columbids,
members of Zenaida and Zenaidura for presence of, 1) species-
specific antigens of Columba guinea (in relation to Columba livia)
which are designated A, B, C and E, and, 2) species-specific an-
tigens of C. livia (in relation to C. guinea) which are designated
A@ Bi Goandtiz
In the first test all five species of Zenaida and Zenaidura possessed
antigens A and C, and all but auriculata possessed E. None of the
species gave evidence of the presence of the B antigen of C. guinea
in their blood. In the latter test only macroura had A’, only asiatica
had B’ (aurita was not tested for B’), and none had C’ or E’.
These results would indicate that the five species are similar
regarding antigenic content of the blood, and the variation is not
consistent within one or the other genus as presently known.
SUMMARY AND CONCLUSION
The avian genus Zenaida is currently considered to be distinct
from the genus Zenaidura by most columbid taxonomists. The jaw
muscles of six Mourning Doves (Zenaidura) and five White-winged
Doves (Zenaida) were investigated as to differences and similarities
that might clarify the relationships of the genera. The sizes and
proportions of skulls were also considered in 37 Mourning and
White-winged doves and two Eared Doves. Larger size of M.
pseudotemporalis profundus, the muscle that functions simultane-
ously as an adductor of the lower jaw and retractor of the upper
jaw, in the White-winged Dove was the character found in the jaw
musculature that could be used to support the contention that
Zenaidura and Zenaida represent distinct genera. Larger size of
this muscle in the white-wing seems to be related to its elongated
beak. The long beak apparently is used for nectar-feeding in
flowers of the Saguaro Cactus.
Excluding the beak, skulls of the white-wing and Mourning
doves are of nearly the same shape. Previous investigators have
shown that in Zenaida and Zenaidura plumage patterns are similar,
nesting habits and eggs are nearly identical, blood proteins are
similar, and one “intergeneric” hybridization in nature is known.
Consequently, it is concluded that species of the two alleged
genera are congeneric, and I agree with Goodwin (1958) that the
genus Zenaida (Bonaparte, 1888:41) should include the Mourning
Dove, Eared Dove, Socorro Dove, Zenaida Dove, and White-winged
Jaw Muscu.Lature OF Doves 539
Dove. Their Latin binomina are Zenaida macroura, Zenaida auricu-
lata, Zenaida graysoni, Zenaida aurita, and Zenaida asiatica, re-
spectively.
pterygoideus dorsalis medialis (insertion)
‘ adductor mandibulae posterior
\ lateral tendon
pseudotemporalis superficialis (insertion)
medial tendon
pseudotemporalis profundus (origin)
pterygoideus dorsalis lateralis (insertion)
pterygoideus ventralis lateralis (insertion)
adductor mandibulae extenus profundus (insertion)
adductor mandibulae externus medialis (insertion)
Fic. 1. Medial view of left ramus of lower mandible of Mourning Dove. x 2%.
Fic. 2. Lateral view of right ramus of lower mandible of Mourning Dove. x 2%.
540 UnIvErsITy OF Kansas Pusts., Mus. Nat. Hist.
so adductor mandibulae posterior
i adductor mandibulae externus profundus (insertion)
adductor mandibulae externus superficialis (insertion)
fe)
medial tendon
\ pseudotemporalis superficialis
lateral tendon (insertion)
depressor mandibulae medialis (insertion)
/-—Pterygoideus ventralis lateralis (insertion)
depressor mandibulae superficialis medialis
(insertion)
A
depressor mandibulae superficialis lateralis
(insertion)
Fic. 8. Dorsal view of lower mandible of Mourning Dove. > 2%.
Fic. 4. Ventral view of lower mandible of Mourning Dove. X 2%.
Fic.
Fic.
Fic.
Fic.
CD
Jaw MuscuLature oF Doves
protractor quadrati (insertion)
pseudotemporalis profundus (insertion)
adductor mandibulae posterior
protractor quadrati (insertion)
Dorsal view of right quadrate of Mourning Dove.
Dorsal view of right pterygoid of Mourning Dove.
Ventral view of right quadrate of Mourning Dove.
Ventral view of right pterygoid of Mourning Dove.
pterygoideus dorsalis
medialis (origin)
adductor mandibulae externus profundus (origin)
protractor pterygoidei (insertion)
pterygoideus dorsalis medialis (origin)
~
xX oO.
x 5.
~
sab:
SD:
541
542 University OF Kansas Pusus., Mus. Nat. Hist.
adductor mandibulae externus superficialis (origin)
adductor mandibulae externus profundus (origin)
adductor mandibulae externus medialis (origin)
protractor quadrati (origin)
protractor pterygoidei (origin)
pterygoideus dorsalis
9 / ! | lateralis (origin)
/ / ee enyaodene dorsalis medialis (origin)
(“depressor mandibulae superficialis medialis (origin)
depressor mandibulae superficialis lateralis (origin)
depressor mandibulae superficialis medialis
(origin)
pterygoideus ventralis lateralis
(origin)
/ [oS wieryaoideus ventralis medialis (origin)
4_adductor mandibulae externus medialis (origin)
Fic. 9. Right lateral view of skull of Mourning Dove. xX 2%.
Fic. 10. Ventral view of skull of Mourning Dove. X 22.
Jaw MuscuLatTurRE OF Doves 543
interorbital septum
optic foramen
protractor pterygoidei
(origin)
protractor quadrati
(origin)
pseudotemporalis
Sees superficialis (origin)
trigeminal foramen
ets rostrum
pterygoid
quadrate
adductor mandibulae externus profundus (origin)
Fic. 11. Cross section of skull of Mourning Dove; anterior
view. xX 2.
Fic. 12. Dorsal view of right quadrate of Mourning Dove
showing movement which protracts the upper mandible
(broken line). > 5.
544 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
adductor mandibulae externus profundus pseudotemporalis profundus
pseudotemporalis superficialis pterygoideus dorsalis medialis
protractor quadrati
I4
adductor mandibulae mandible
externus superficialis pterygoideus dorsalis lateralis
adductor mandibulae externus medialis
depressor mandibulae superficialis medialis
depressor mandibulae superficialis lateralis
Fic. 13. Right lateral view of the jaw musculature of the White-winged Dove;
superficial layer. x 5.
Fic. 14. Right lateral view of the jaw musculature of the Mourning Dove;
superficial layer. x 5
Jaw Muscu.aTurE oF Doves 545
pseudotemporalis superficialis
| fsa ee dorsalis medialis
= pseudotemporalis profundus
|
pterygoid ————_, he | | r—Ppterygoideus dorsalis lateralis
|
= - 1s |
protractor pterygoidei | | |
protractor quadrati—
quadrate
mandible
ZZ adductor mandibulae externus medialis
adductor mandibulae externus profundus
Fic. 15. Dorsal view of the jaw musculature of the White-winged Dove
(right side); superficial layer. x 5
Fic. 16. Dorsal view of the jaw musculature of the Mourning Dove (right
side); superficial layer. x 5.
546 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hisr.
pterygoid emai adductor mandibulae posterior
protractor pterygoidei
protractor quadrati
I8
pterygoideus dorsalis medialis
pterygoideus dorsalis lateralis
quadrate
trigeminal foramen — \\ mandible
ae: = LEGER astarce mandibulae externus medialis
adductor mandibulae extemus profundus
Fic. 17. Dorsal view of the jaw musculature of the White-winged Dove (right
side); middle layer. x 5.
Fic. 18. Dorsal view of the jaw musculature of the Mourning Dove (right
side); middle layer. » 5.
Fic. 19. Dorsal view of the jaw musculature of the White-winged Dove
(right side); deep layer. x 5.
Fic. 20. Dorsal view of the jaw musculature of the Morning Dove
(right side); deep layer. x 5.
548 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
pseudotemporalis profundus
pterygoideus dorsalis lateralis
pterygoideus dorsalis medialis
Fic. 21. Ventral view of the jaw musculature of the White-winged Dove
(M. depressor mandibulae not shown). x 5
Fic, 22. Ventral view of the jaw musculature of the Mourning Dove (M.
depressor mandibulae not shown). X 5
Jaw MuscuLaTuRE OF DovEs 549
LITERATURE CITED
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1919. A memoir on the phylogeny of the jaw muscles in recent and fossil
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AUDUBON, J. J.
1834. Ornithological biography. Vol. II. Adam & Charles Black, Edin-
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Bairp, S. F., BREwER, T. M., and Rmeway, R.
1905. The land birds of North America. Little, Brown, and Company,
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BEECHER, W. J.
1950. Convergent evolution in the American orioles. Wilson Bull., 62:
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1951b. Convergence in the Coerebidae, Wilson Bull., 63:274-287.
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BENT, A. C.
1932. Life histories of North American gallinaceous birds. Bull. U. S.
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BonaPaBkTE, C. L.
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OND, J.
1961. Birds of the West Indies. Houghton Mifflin Company, Boston, 256
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Bowmaw, R. I.
1961. Morphological differentiation and adapiation in the Galapagos
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Davi, O.
1889. Nests and eggs of North American birds. Hann & Adair, Colum-
bus, 455 + xii pp., 13 pls.
EpGEworTH, F. H.
1935. The cranial muscles of vertebrates. Cambridge Univ. Press, viii +
498 pp., 841 figs.
ENGELS, W. L.
1940. Structural adaptations in thrashers (Mimidae: genus Toxostoma)
with comments on interspecific relationships. Univ. California
Publ. Zool., 42:341-400, 24 figs., 11 tables.
Fisuer, H. I.
1955. Some aspects of the kinetics in the jaws of birds, Wilson Bull.,
67:175-188, 4 figs., 6 tables.
Gavow, H.
1891. Vogel: I. Anatomischer Theil. Bronn’s Klassen und Ordnungen des
Thier-Reichs. C. F. Winter, Leipzig, 6:1-1,008, many figs., 59 pls.
GiLmaNn, M. F.
1911. Doves on the Pima Reservation. Condor, 18:51-56.
GoopMan, D. C., and Fisuer, H. I.
1962. Functional anatomy of the feeding apparatus in waterfowl. South-
ern Illinois Univ. Press, Carbondale, xii +- 193 pp.
550 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hisr.
Goopwin, D.
1958. Remarks on the taxonomy of some American doves. Auk, 75:330-
334.
Goss, N. S.
1891. History of the birds of Kansas. Geo. W. Crane & Co., Topeka,
692 pp., 35 pls.
HENSLEY, M. M.
1954. Ecological relations of the breeding bird populations of the desert
biome of Arizona. Ecol. Monographs, 24:185-207.
Horer, H.
1950. Zur Morphologie der Kiefermuskulatur der Vogel. Zool. Jahrb.
Jena (Anat.), 70:427-556, 44 figs.
Irwin, M. R., and MmILLer, W. J.
1961. Interrelationships and evolutionary patterns of cellular antigens in
columbidae. Evolution, 15:30-48.
Jackson, A. S.
1941. The mourning dove in Throckmorton County, Texas, Unpubl.
manuscript (Abstract).
Kieu, W. H., Jn., and Harris, J. T.
1956. Status of the white-winged dove in Texas. Trans, 21st N. Amer.
Wildl. Conf., pp. 376-388.
KNAPPEN, P.
1938. Preliminary report on some of the important foods of the mourning
dove in the southeastern U. S. Trans. 3rd N. Amer. Wildl. Conf.,
pp. 776-781.
Laxjer, T.
1926. Studien Uber die Trigeminus-versorgte Kaumuskulatur der Sau-
pops C. A. Reitzel Buchhandlung, Kopenhagen, 154 pp., 26
pls.
Lropotp, A. S.
1943. Autumn feeding and flocking habits of the mourning dove in
southern Missouri. Wilson Bull., 55:151-154.
McGrecor, S. E., ALcorn, S. M., and Ou, G.
1962. Pollination and pollinating agents of the saguaro. Ecology, 43:
259-267.
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1922. A study of the nesting of mourning doves. Auk, 39:457-474;
40:87-58.
Peters, J. L.
1934. The classification of some American pigeons. Condor, 36:213-215.
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Rootn, J.
19538. On the correlation between the jaw muscles and the structure of
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SALVADORI, T.
1893. Catalogue of birds in the British Museum, 21:xvii + 676 pp., 15
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SHUFELDT, R. W.
1890. The myology of the raven. MacMillan & Co., London, xix + 348
pp., 76 figs.
Jaw MuscuLaTurE OF Doves 551
Sms, R. W.
1955. The morphology of the head of the hawfinch. Bull. Brit. Mus.
(Nat. Hist.) Zool., 2:369-393.
WETMORE, A.
1920. Observations of the habits of the white-winged dove. Condor,
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1959. The function of the depressor mandibulae muscle in certain pas-
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Transmitted June 3, 1963.
O
29-7865
wee ery Faatiamieiel car Tl ae ee
lie —_ LIBRARY
JUL 21 {564
HARV AZ PD
UNIVERSITY,
UNIVERSITY OF KANSAS PUBLICATIONS
MuSsEUM OF NATURAL HISTORY
Volume 12, No. 18, pp. 553-573, 7 figs.
— March, 2, 1964
Thoracic and Coracoid Arteries
In Two Families of Birds,
Columbidae and Hirundinidae
BY
MARION ANNE JENKINSON
UNIVERSITY OF KANSAS
LAWRENCE
1964
UNIVERSITY OF KAnsAsS PUBLICATIONS, MusEUM OF NATURAL History
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 12, No. 13, pp. 553-573, 7 figs.
Published March 2, 1964
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY THE STATE PRINTER
TOPEKA, KANSAS
1964
LIBRARY
Jur ZL io4
HARVARD
Thoracic and Coracoid Arteries®!" Y
In Two Families of Birds,
Columbidae and Hirundinidae
BY
MARION ANNE JENKINSON
CONTENTS
PAGE
TNTRODUCTION rte fies So cesses Soy ok sk Scat Shins es asp mie shel eee eee 555
METHODSEAND MATERTALS) ogc a cuss 6s og ese oe an ee See 556
MyoLocy AND ANGIOLOGY: HIRUNDINIDAE ....................00000: 557
INGVOIOEM Gee eas Glade Gabe ccuiwn sage ebb ka Rae We oa oe Oe 557
ADPIOIOG | do $2 tae cee eae oh od Uaitls SEM as wae beaks oa eee 558
MYOLOGY AND ANGIOLOGY: COUUMBIDAE <2. .4:)...0)..0. 0.25 eee 560
IMEVOlO Gy Me... Gary tee co eer oa aan Sees owe SST. Sl tore eee 560
AM@IOIO RY. ons ss ales de Ese nk wk oes Dae Smeal he ia Pee 560
SUMMARY OF ARTERIAL ARRANGEMENT ................000 050000 eee 562
DISCUSSION ZAND" GONCLUSIONS (25) 0.22 + foc.e dys Bin v1 Ste chs 2d eee 562
Individual: Variation) (7:52. «2 Sk ous. oee Pee he brs: £| coalesce Saeeanaee 562
dnitratamilialy DitterenCes® - 5. 1s «oe ahiee oud eee hed bas dnd pane seen 563
Interfamilial” Differences... 6 ek ee eee cease 3 ioe ee 565
SUNENEAT aud a, been Mins iets hee ar brea Sparcis, sistent ae Wanner 567
LEVRERA TURE | GLUED ptt ac: suis hier ac Gishs- sas ope alee opie eee aus oan ee 573
INTRODUCTION
Most descriptions of the circulatory system of birds, largely the
work of Glenny, have dealt with arteries of the neck and thorax
in a wide variety of species. As a result of his work, Glenny offered
several hypotheses concerning the phylogenetic, hence taxonomic,
significance of differences in some of these vessels. He also de-
scribed six types of thoracic arterial arrangements and stated that
these categories might represent various levels of evolution (Glenny,
1955:548-544 ).
The families Columbidae (pigeons) and Hirundinidae (swal-
lows) have two nearly extreme arterial types described by Glenny,
and are universally acknowledged as monophyletic. Differences
within the families, therefore, can be considered as valid intra-
familial differences. I have investigated the thoracic and coracoid
arteries and their branches in members of these two families to de-
termine the degree of individual variability of the vessels, and the
possible causes of interspecific and intrafamilial differences.
(555)
556 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
METHODS AND MATERIALS
All specimens studied are in The University of Kansas Museum of Natural
History. They were preserved in alcohol and their blood vessels were not
injected. Dissections were made with the aid of a binocular microscope at
magnifications of 10 x and 20 x.
Following is a list of the species studied, the number of individuals of each
species dissected, and the catalogue numbers of the specimens. The nomen-
clature and classification are those of the American Omithologists’ Union’s
Check-List of North American Birds, fifth edition (1957).
Family Columbidae
Zenaidura macroura (Linnaeus), Mouming Dove 2: 40325, 40826.
Zenaida asiatica (Linnaeus), White-winged Dove 1: 40328.
Scardafella inca (Lesson), Inca Dove 5: 34894, 34896, 34902, 34906, 34907.
Columba livia Gmelin, Rock Dove (domestic pigeon) 1: 40321.
Family Hirundinidae
Iridoprocne bicolor (Vieillot), Tree Swallow 1: 38101.
Progne subis (Linnaeus), Purple Martin 5: 87711, 38794, 38796, 38798,
88804,
Stelgidopteryx ruficollis (Vieillot), Rough-winged Swallow 1: 38277.
Riparia riparia (Linnaeus), Bank Swallow 2: 38784, 38785.
Hirundo rustica Linnaeus, Barn Swallow 1: 38839.
The following descriptions are of Progne subis and Scardafella inca. Differ-
ences in the vascular system in other members of the families represented by
P. subis and S. inca are mentioned at the appropriate places. The muscles
briefly described for each of these two species are those that are supplied by
the thoracic or coracoid arteries or by branches of the same, and muscles that,
by their origin, location, or insertion, seem to affect the course or origin of one
of these arteries.
The following sources have been particularly useful for the terminology of
muscles and of skeletal features: Ashley (1941), Beddard (1898), Coues
(1903), Howard (1929), Howell (1937), and Hudson and Lanzillotti (1955).
The names used for most arteries are those in common usage for vertebrates.
I have not used the terms “internal mammary” and “intercostal” artery as sub-
stitutes for “thoracic” artery, except when referring to the work of others.
The vessel’s homology with the internal mammary artery of mammals has been
denied (Glenny, 1955:541), and the name “mammary” is certainly not useful
descriptively in birds. The term “intercostal” is less objectionable, except that
such a name may call to mind segmental vessels arising from the dorsal aorta.
The term “thoracic” seems best, as it is reasonably descriptive, and has been
used by Glenny in the majority of his descriptions covering a wide variety of
birds. The name “sternoclavicular” has been used by others as a synonym fo:
the “coracoid” artery. I have arbitrarily chosen to use the latter.
ACKNOWLEDGMENTS
I gratefully acknowledge many valuable suggestions in my research and the
preparation of this manuscript from Professors Theodore H. Eaton, A. Byron
Leonard, Richard F. Johnston, Robert M. Mengel, and E. Raymond Hall.
Mr. Abbot S. Gaunt and Miss Sandra Lovett assisted in collecting specimens.
Final drafts of the illustrations were prepared by Mr. Thomas Swearingen.
ARTERIES IN Two FAMILIES OF Birps 557
MYOLOGY AND ANGIOLOGY: HIRUNDINIDAE
Figs. 1, 2, 3, and 4 illustrate the following muscles and arteries
described for Progne subis.
Myology
M. pectoralis thoracica, Fig. 1. The origin is from slightly less than the
posterior half of the sternum, from the ventral half of the keel, almost the
entire length of the posterolateral surface of the clavicle and adjacent portion
of the sterno-coraco-clavicular membrane, and tendinously from the ventral
thoracic ribs. This massive muscle covers the entire ventral surface of the
thorax and converges to insert on the ventral side of the humerus on the
pectoral surface.
M. supracoracoideus, Fig. 1. The origin is from the dorsal portion of the
keel and medial portion of the sternum, and is bordered ventrally by the origin
of M. pectoralis thoracica, and laterally by M. coracobrachialis posterior. The
origin is also from the manubrium and the anterolateral portion of the proximal
half of the coracoid and to a slight extent from the sterno-coraco-clavicular
membrane adjacent to the manubrium. This large pinnate muscle converges,
passes through the foramen triosseum, and inserts by a tendon on the external
tuberosity of the humerus, immediately proximal to the insertion of M. pec-
toralis thoracica.
M. coracobrachialis posterior, Figs. 1 and 3. The origin is from the dorso-
lateral half of the coracoid, anterolateral portion of the sternum (where the
area of origin is bordered medially by M. supracoracoideus, posteriorly by
M. pectoralis thoracica, and laterally by M. sternocoracoideus), and also to a
slight extent from the area of attachment of the thoracic ribs to the sternum.
The muscle fibers converge along the lateral edge of the coracoid and insert
on the median crest of the humerus immediately proximal to the pneumatic
foramen. In passing from the origin on the sternum to the insertion on the
humerus, the belly of the muscle bridges the angle formed by the costal process
of the sternum and the coracoid.
M. sternocoracoideus, Figs. 2 and 3. The origin is from the entire external
surface of the costal process of the sternum, and to a small extent from the
extreme proximal ends of the thoracic ribs where they articulate with the costal
process. The muscle inserts on a triangular area on the dorsomedial surface
of the coracoid. Like M. coracobrachialis posterior, this muscle bridges the
angle formed by the costal process and the coracoid.
M. subcoracoideus (ventral head), Figs. 2 and 3. The origin is from the
dorsomedial edge of the coracoid at its extreme proximal end, and to a slight
extent from the adjacent portion of the manubrium. The origin is medial to
the insertion of M. sternocoracoideus. The ventral head passes anterodorsally
along the medial edge of the coracoid and joins the dorsal head (not here
described). The combined muscle then inserts by a tendon onto the internal
tuberosity of the humerus.
M. costi-sternalis, Figs. 1, 2, and 3. The origin is from the anterior edge
of the sternal portion of the first four thoracic ribs. This triangular muscle
narrows and inserts on the posterior edge of the apex of the costal process.
The portion arising from the first rib may share slips with M. sternocoracoideus.
558 UNIVERSITY OF KAnsaAs Pus3s., Mus. Nat. Hist.
M. costi-sternalis anterior, Figs. 1, 2, and 8. This muscle is variously
developed, and originates from a small area on the ventral end of the vertebral
portion of the last cervical rib. The insertion is on the apex of the costal
process, immediately anterior to the insertion of M. costi-sternalis.
Mm. intercostales externus, Fig. 1. These muscles extend posteroventrally
between the vertebral portions of successive thoracic ribs, and between the
last cervical and first thoracic ribs. In the more posterior intercostal spaces
these muscles are poorly developed, but they become progressively better de-
veloped anteriorly, and are fully represented in the most anterior intercostal
spaces.
Mm. intercostales internus, Fig. 3. These muscles resemble the external
intercostal muscles, but extend anteroventrally, with the muscles being most
fully developed posteriorly, and progressively less so anteriorly.
Costopulmonary muscles, Fig. 3. This diagonal series of muscle slips from
the thoracic ribs attaches to the aponeurosis covering the lungs.
Angiology
Figs. 8 and 4 show all arteries discussed for this family. The numbers fol-
lowing the names or descriptions of arteries in the text refer to numbered
arteries in one or both of these figures.
The right and left innominate or brachiocephalic arteries arise from the
aortic trunk and give rise to the common carotid arteries (14). The major
vessel continuing across the thoracic cavity is the subclavian artery. Classically
the subclavian is considered as continuing into the anterior appendage as the
axillary artery. However, in the species studied, the axillary artery can best
be described as a branch from the subclavian; the pectoral stem forms a more
direct continuation of the subclavian. In traversing the thoracic cavity, the
subclavian gives rise to the thoracic, coracoid, and axillary arteries, and leaves
the thoracic cavity as the pectoral trunk, dorsal to the area where Mm. coraco-
brachialis posterior and sternocoracoideus span the agle formed by the coracoid
and costal process.
The pectoral trunk bifurcates into two main pectoral arteries (9), which
penetrate M. pectoralis thoracica. Neither the axillary artery nor these pectoral
arteries were traced in my study.
The coracoid artery (2) arises from the ventral face of the subclavian (1),
either opposite the base of, or medial to, the axillary artery (10). The coracoid
artery passes ventrad between the medial edge of the coracoid and the ventral
head of M. subcoracoideus, and an artery (7) is given off to supply that
muscle. The main vessel then penetrates M. supracoracoideus and bifurcates
or ramifies into several vessels (12).
Between the origin of the coracoid artery from the subclavian, and the
point where the coracoid artery passes the medial edge of the coracoid, several
branches are given off. These vessels are highly variable in origin, as described
below, and not all were always found. Along with the coracoid artery, they
are termed a “coracoid complex.”
The first artery (11) of this complex arises from any one of several places:
from the lateral face of the coracoid artery at its base; independently from the
subclavian immediately lateral to the origin of the coracoid artery; and from
the thoracic artery near its origin. This vessel travels laterad, parallel to the
ARTERIES IN Two FAMILIES OF Birps 559
subclavian, and penetrates M. coracobrachialis posterior at the same point that
the pectoral artery passes dorsal to that muscle.
Another vessel (common stem of 4 and 5) of the coracoid complex in most
specimens arises from the anterior face of the coracoid artery and branches
into several vessels, some of which (5) supply M. subcoracoideus, and some
of which (4) feed M. coracobrachialis posterior. The vessel occasionally shares
a common stem with the main vessel (11) to M. coracobrachialis posterior, and
in some specimens arises independently from the subclavian, immediately
anterior to the origin of the coracoid artery. The branch (4) to M. coraco-
brachialis posterior was also seen to arise independently from any of the above-
mentioned positions.
Two remaining vessels (6 and 8) are often found as branches from the
coracoid artery. They were small and often were collapsed in the individuals
I dissected, but were most clearly seen in Iridoprocne bicolor. The vessels
occasionally had a common base, and in some specimens only one vessel was
found. The first artery (6) passes mediad into M. sternocoracoideus, or con-
tinues across that muscle onto the inner face of the sternum. The second vessel
(8) also supplies M. sternocoracoideus or the inner surface of the sternum, and
often a large branch continues across the dorsal surface of the coracoid to M-
coracobrachialis posterior. Fig. 3 shows a composite of these vessels; not all
branches were seen in any one specimen. In the specimen of I. bicolor a fora-
men existed on the lateral edge of the coracoid where the branch (of 8) to
M. coracobrachialis posterior passed. An examination of skeletons of five to
10 individuals each of the five species for which dissections were made, and of
Petrochelidon pyrrhonota (Cliff Swallow) and Tachycineta thalassina (Violet-
green Swallow), in the University of Kansas collection, showed that most
coracoids of these seven species of swallows had a small notch (as shown in
Fig. 4) or a complete foramen there.
The thoracic artery (3) arises from the subclavian opposite the base of the
coracoid artery, or from the base of the coracoid artery. Of the five specimens
of P. subis dissected, one individual had the former arrangement on both sides,
and one had the latter on both sides, whereas in the remaining three the
thoracic artery arose from the coracoid artery on one side and from the sub-
clavian on the other side. The distance between these two possible sites of
origin is slight.
The thoracic artery usually passes ventral to M. costi-sternalis anterior. Oc-
casionally a small artery (13) could be traced from the main trunk of the
thoracic artery to that muscle. The main thoracic artery bifurcates near the
insertion of M. costi-sternalis, the branches traveling posteriad on both sides
of the muscle. On one side of one specimen this artery bifurcated immediately
after leaving the subclavian, the dorsal trunk passing dorsal to M. costi-sternalis
anterior, and the ventral trunk ventral to the muscle. On the other side of the
same individual the artery passed dorsal to M. costi-sternalis anterior, bifurcat-
ing at the normal point.
From the ventral trunk of the thoracic artery a variable number of small
vessels arises to supply the costosternal articulations. The main ventral trunk
bifurcates into two branches, one of which passes onto the inner face of the
sternum, and one of which supplies the posterior two intercostal spaces.
The dorsal thoracic trunk supplies M. costi-sternalis, several dorsal inter-
costal areas, and the costopulmonary muscles. Minor variations in all of the
smaller branches of the thoracic artery were common,
560 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
MYOLOGY AND ANGIOLOGY: COLUMBIDAE
Figs. 5, 6, and 7 illustrate the following muscles and arteries de-
scribed for Scardafella inca.
Myology
M. pectoralis thoracica, Fig. 5. The origin is from approximately the
ventral third of the keel, the lateral and anterior portion of the clavicle and
the adjacent sterno-coraco-clavicular membrane, and from the lateral portion
of the sternum and the fascia overlying the thoracic ribs. This massive muscle
covers the entire ventral surface of the thorax, converges, and inserts on the
pectoral surface on the ventral side of the humerus.
M. supracoracoideus, Fig. 5. The origin is from the dorsal two-thirds of
the keel and medial half of the sternum (where the origin is bordered ven-
trally, posteriorly, and laterally by the origin of M. pectoralis thoracica) and
from the sterno-coraco-clavicular membrane adjacent to the coracoid. This
large pinnate muscle converges, passes through the foramen triosseum, and
inserts by means of a strong tendon on the dorsal surface of the humerus on
the deltoid ridge.
M. coracobrachialis posierior, Fig. 5. The origin is from a prominent
lateral wing on the posterolateral portion of the coracoid, and from the lateral
surface of the proximal two-thirds of the coracoid. The insertion is by means
of a tendon on the internal tuberosity of the humerus. Of the musles de-
scribed here, this one differs most strikingly from the homologous muscle in
P. subis. The difference can be seen by comparing Figs. 1 and 5.
M. sternocoracoideus, Figs. 5, 6, and 7. The origin is from the external,
and to a slight extent from the internal, surface of the costal process. The
insertion is on a posterolateral triangular area on the dorsal surface of the
coracoid.
M. costi-sternalis, Figs. 5 and 6. The origin is from the anterior edge of
the sternal portion of the first three thoracic ribs. The muscle converges and
inserts on the apex of the costal process.
M. subcoracoideus (ventral head), Fig. 6. The origin is from the manu-
brium and from approximately the posterior half of the coracoid and on the
medial and dorsal surface of that bone, and the medial side of the sterno-
coraco-clavicular membrane adjacent to the coracoid. The ventral head passes
anterodorsally to join with the dorsal head (not here described), and the
combined muscle inserts by a tendon on the internal tuberosity of the humerus.
Mm. intercostales externus, Fig. 5. These muscles extend posteroventrally
between successive thoracic ribs and between the last cervical and first thoracic
ribs.
Mm. intercostales internus, Fig. 7. These muscles extend anteroventrally
between the last three thoracic ribs.
Costopulmonary muscles, Fig. 7. This series of muscle slips from the
thoracic ribs attaches to the aponeurosis covering the lungs.
Angiology
Figs. 5, 6, and 7 show all arteries discussed for this family. The numbers
following names or descriptions of arteries in the text refer to numbered arteries
ARTERIES IN Two FAMILIES OF Breps 561
in one of these figures. Insofar as possible, the numbers used for these arteries
are the same numbers used for the homologous vessels in swallows.
The right and left innominate arteries arise from the aortic trunk and give
rise to the common carotid (14) and subclavian (1) arteries. The latter
continues across the thoracic cavity, giving rise to the coracoid (2) and
axillary (10) arteries, and becoming the pectoral trunk. That trunk swings
posteriorly and leaves the thoracic cavity near the apex of the costal process,
as shown in Fig. 7. Where the trunk passes under M. sternocoracoideus, the
thoracic artery (3) is given off.
The various branches of the coracoid artery, again referred to as a “coracoid
complex,” are as follows: The first branch, from the posterior face of the
coracoid artery, is a relatively large vessel (6) here termed the sternal artery;
it passes mediad across M. sternocoracoideus, sending off a branch (6a) to that
muscle, The right sternal artery continues posteriorly on the midline of the
inner surface of the sternum, and appears to send branches into the various
pneumatic foramina of the sternum, but these vessels are minute and exceed-
ingly difficult to trace accurately. The corresponding left vessel is smaller and
ramifies on the anteromedial surface of the stemum, Variations found in these
vessels were the following: In one specimen of S. inca the sternal artery had, on
both sides, an independent origin from the subclavian, lateral to the origin of
the coracoid artery. In Zenaidura macroura both right and left sternal arteries
were similar to the left vessel described above, no median longitudinal vessel
being seen. In Columba livia no vessel corresponding to the sternal artery was
seen. In Zenaida asiatica these arteries penetrated M. sternocoracoideus; no
branch to the sternum was seen,
A small complex of vessels (4 and 4a) arises from the lateral face of the
coracoid artery and feeds M. coracobrachialis posterior, and occasionally M.
sternocoracoideus, One branch (4a) passes under the coracoid and travels
along the lateral side of that bone, supplying small branches to M. coraco-
brachialis posterior, and finally ramifying on the head of the coracoid. In C.
livia, Zenaidura macroura, and Zenaida asiatica this complex usually arises in-
dependently from the subclavian, and in one case it arose from the axillary
artery.
Two other branches from the coracoid artery were regularly seen. The
first (8) passes across M, sternocoracoideus and appears to supply the area of
the coracoid articulation with the sternum; the second (7) supplies M. sub-
coracoideus as the main vessel passes between that muscle and the coracoid
and penetrates M. suparacoracoideus. A small notch on the medial side of
the coracoid (shown in Figs. 6 and 7) often marks the passage of the coracoid
artery.
All vessels of the coracoid complex are exceedingly variable, in number, size,
and site of origin.
A prominent vessel (15) is given off from the posterior pectoral artery, out-
side the thoracic cavity, passes ventrad, and sends two branches into M. supra-
coracoideus. No corresponding artery was seen in the swallows dissected.
The thoracic artery (3), arising from the pectoral stem, characteristically
bifurcates at the anterior end of M. costi-sternalis. The dorsal, and larger,
branch passes posteriorly, sends several small branches to M. costi-sternalis,
and continues to the most posterior rib. The ventral trunk bifurcates, one
branch passing along the edge of, and supplying, M. costi-sternalis, the other
562 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
branch passing onto the surface of the stemum. In some specimens two such
branches to the sternum were seen.
SUMMARY OF ARTERIAL ARRANGEMENT
In both families the vessels that are relatively constant in ap-
pearance are: a subclavian giving rise to the carotid and axillary
arteries, and becoming the pectoral trunk; the thoracic artery arising
variously, and passing posteriorly to the rib cage; and the coracoid
complex of vessels. The coracoid complex includes the coracoid
artery, the vessels to Mm. sternocoracoideus and coracobrachialis
posterior, and the sternal artery, which is variously present, and
more extensive in some species than in others.
DISCUSSION AND CONCLUSIONS
In the vessels studied individual variation is marked, but the
arterial arrangement within both families is relatively constant.
Interfamilial differences probably represent responses of the arteries
to adaptive structural differences of other systems of the body.
Individual Variation
The term “individual variation” is used here to mean “continuous
non-sex-associated variation” (see Mayr, Linsley, and Usinger,
1953:93) found between members of the same species or between
the two sides of the same individual. It is hazardous to define in-
dividual variation (and also interspecific differences, as discussed
later) in the origin of one vessel by relating its location to other
vessels, because these may likewise vary in origin. But, by neces-
sity, certain vessels that are probably less variable (axillary, carotid,
and pectoral arteries) have been considered here as being constant
in origin. If these three vessels are accepted as reference points,
individual variants, as well as interspecific differences, can easily be
described in the thoracic and coracoid arteries and in their various
branches.
The thoracic artery in P. subis arose either from the subclavian
artery, or from the coracoid artery. Likewise in other swallows,
both of these origins were found. In doves the thoracic artery
arose consistently from the pectoral stem, lateral to the origin of
the axillary artery.
The coracoid artery in P. subis and other swallows arose from the
subclavian artery, either opposite the base of the axillary artery, or
medial to that vessei. In all doves studied the coracoid artery arose
from the subclavian medial to the axillary artery. I observed much
ARTERIES IN Two FAMILIES OF BIRDS 563
individual variation in the branches of the coracoid artery (that is
to say, in the vessels of the coracoid complex). In S. inca the
sternal artery arose either from the coracoid artery, or independently
from the subclavian. As mentioned earlier, in members of both
families the vessels to Mm. coracobrachialis posterior and sub-
coracoideus are highly variable, arising in swallows from the cora-
coid artery or from the subclavian artery, and in doves from either
of these two sites or from the axillary artery. The distribution of
these arteries after their origin is also diverse.
Individual variation in the arteries of the thorax has been re-
corded previously. Bhaduri, Biswas, and Das (1957:2) state that,
in the domestic pigeon, “the origin and course of various smaller
arteries . . . show noticeable variation,” although they do not
specifically state to which vessels they are referring. Fisher (1955:
287-288) found variability in the Whooping Crane, Grus americana,
of the axillary, coracoid, thoracic, and pectoral arteries. In one
specimen he found these vessels arising on the right side from the
subclavian, in the sequence just listed, and on the left side all arose
from the same point. Berger (1956:439-440) strongly emphasized
the variability of the vascular system, calling it the most variable in
the body. As he stated, this high degree of individual variation
seems to be due to the embryological development of the system,
wherein many of the adult channels of circulation are derived from
embryonic plexuses.
Intrafamilial Differences
In spite of the rather extensive amount of individual variability
in some vessels, I found the over-all pattern of arteries to be rel-
atively constant within the family Columbidae and within the family
Hirundinidae. There are, nevertheless, several intrafamilial dif-
ferences needing some further discussion and clarification.
Others have reported the occasional presence of more than one
coracoid artery on each side in some columbids, these arteries be-
ing described as arising from various sites and being variously
named. Bhaduri and Biswas (1954) described the arterial situation
in seven species of the family Columbidae (Columba livia, Strep-
topelia tranquebarica, S. chinensis, S. senegalensis, Chalcophaps
indica, Treron bicincta, and T. phoenicoptera) and stated (op.
cit.; 348) that “The sterno-clavicular [= coracoid] artery is similar
in all the species, but the domestic pigeon seems to be unique in
that it has, in addition, a small vessel, the accessory sternoclavic-
ular.” This artery was described later, in the domestic pigeon, as
564 UNIVERSITY OF KAnsAS Pus3s., Mus. Nat. Hist.
follows (Bhaduri, Biswas, and Das, 1957:5): “A minute and insig-
nificant vessel which has been termed the accessory sternoclavicular
artery . . . is given off close to the origin of the sternoclavic-
ular. It passes antero-ventrally to supply the adjacent muscles.”
Glenny (1955:577) described the arterial pattern characteristic of
members of the family Columbidae (more than 30 species studied
by him) and stated that “three pairs of coracoid arteries are found
in Otidiphaps nobilis, normally one or two pairs may be found.” As
suggested by Bhaduri and Biswas (1954:348), the “accessory” vessel
probably corresponds to a vessel previously described by Glenny
(1940) in Streptopelia chinensis and referred to as the “coracoid
minor.”
Bhaduri and Biswas (1954:348) have suggested that “the ac-
cessory sterno-clavicular artery occurring sporadically as it does in
some species of diverse groups may not have any phylogenetic
value.”
In no case did I find more than one coracoid artery on a side.
When one of the highly variable arteries feeding Mm. coraco-
brachialis posterior and sternocoracoideus (arteries 4 and 4a, Fig.
7) arises from the subclavian or axillary artery instead of from the
coracoid artery, that vessel may have been interpreted by others as
a second (accessory or minor) coracoid artery. If so, this artery
probably does not “occur sporadically.” Rather, its origin from the
subclavian, axillary, or thoracic artery may be sporadic, subject to
individual variation, and it may have been overlooked when it arose
from the coracoid artery.
Of the vessels described here, the only one that differed distinctly
in one species was the sternal artery. In Scardafella inca the right
sternal vessel was long, extending down the mid-line of the inner
surface of the sternum, whereas in other columbids the right and
left arteries ramified on the anterior part of the inner surface of the
sternum, or were altogether lacking. I am unable to account for the
differential development of this artery in S. inca.
In describing the arterial arrangement in the seven species of
Indian columbids named earlier, Bhaduri and Biswas (1954:348)
state that all species except Treron phoenicoptera have two “internal
mammary’ arteries on each side “showing variable sites of origin.”
These arteries were later described (Bhaduri, Biswas, and Das,
1957:4-5) as “a slender (outer) internal mammary artery ‘
to the outer wall of the thoracic cavity” and “a slender (inner) in-
ternal mammary artery . . . to supply the inner wall of the
chest cavity.” From this description, the question arises as to
~
ARTERIES IN Two FAMILIES OF Birps 565
whether the “outer” one of these arteries should properly be called
an external instead of internal mammary artery. In any case, I saw
no specimen possessing two thoracic arteries on a side.
Interfamilial Differences
As shown above, there is a high degree of individual variation in
the vessels being considered, while at the same time, few inter-
specific differences were noted within the families. On the other
hand, the vascular arrangement of swallows consistently differed
from that of pigeons in the species studied. The differences are
most easily described by discussing the resulting change in the site
of origin of the thoracic artery. In swallows the thoracic artery
arises between the carotid and axillary arteries, either from the
stem of the coracoid artery or independently from the subclavian,
but in pigeons the thoracic artery arises from the pectoral stem, a
site of attachment that is relatively more lateral than in swallows.
This difference, in my opinion, demonstrates well the topological
relationships of various systems of the body, here especially of the
skeletal, muscular, and vascular systems. The location of the
thoracic artery seems to be determined by the particular configura-
tion of skeletal and muscular elements, although even within the
bounds set by these elements, individual variation in the precise
origin of the artery is possible. In all swallows dissected Mm.
coracobrachialis posterior and sternocoracoideus bridge the angle
formed by the costal process and the coracoid. This arrangement
makes it necessary for the subclavian to leave the thoracic cavity
dorsal to the costal process, although it does pass immediately
anterior to that process. The thoracic artery arises from the vessel
next to the apex of the costal process, hence from the subclavian,
between the axillary and carotid arteries.
In pigeons, the wing of the coracoid extends farther laterally
than does the costal process, and the apex of the latter is displaced
farther posteriorly than it is in swallows. M. coracobrachialis
posterior does not arise from the sternum, and only part of the
costal process serves as a point of origin for M. sternocoracoideus.
Consequently, this region differs from that of swallows; the area
between the costal process and coracoid is not entirely bridged by
muscle, and the space between the two skeletal elements is of a
different shape and size. It seems that these differences have re-
sulted, in pigeons, in the subclavian assuming a more anterior posi-
tion with reference to the costal process. The subclavian in these
birds leads into the pectoral artery, which runs posteriad, passing
566 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist.
under M. sternocoracoideus and leaving the thoracic cavity ap-
proximately opposite the apex of the costal process. The thoracic
artery arises immediately opposite the apex of the costal process
from the main artery in the area, as it does in swallows, except
that in this case the adjacent artery from which it arises is the
pectoral stem.
The thoracic area seems to be most “efficiently” arranged when
the thoracic artery arises opposite the apex of the costal process,
from whatever main artery is closest to that site. This arrangement
existed in all species studied. Considering the differences in skeletal
and muscular structures, between pigeons and swallows, it would
be much more remarkable if an alternative were the case, that is to
say if the thoracic artery had the same site of attachment on the
subclavian in both groups.
A comparison of these suggestions with statements made previ-
ously about these arteries seems necessary. When Glenny (1955)
summarized his accumulative findings, concerning the main arteries
in the region of the heart, based on individuals representing more
than 750 avian species of 27 orders and 120 families, he described
five types of thoracic arteries that were distinguished by differences
in the site of their origin, and one type in which there were two
thoracic arteries on each side. His statements regarding these dif-
ferences were as follows (Glenny, 1955:543-544):
“The thoracic, intercostal, or internal mammary artery of birds . . . is
found to arise at slightly different relative positions—from a point at the base
of the inferior pectoral artery to a point near the base of the coracoid or sterno-
clavicular artery, and in some instances both of these vessels have a common
root from the subclavian artery. Such differences are found to be of common
occurrence within several orders of birds. In the Galliformes and the Passeri-
formes there appears to be a graded series in the sites of attachment of the
thoracic artery from a lateral to a medial position. As a result of these observa-
tions, numerical values can be assigned to the site of attachment of the inter-
costal or thoracic artery, and these values may come to be used as an index
in specific levels of evolution.
“The medial migration of the thoricte artery appears to have some phy-
logenetic significance as yet not understood.”
The six types of thoracic arteries described in Glenny’s classifica-
tion were distinguished as follows (Glenny, 1955:544):
“Type 1: attachment to the pectoral stem lateral to the axillary.
“Type 2: attachment to the subclavian between the axillary and coracoid.
“Type 3: attachment to the subclavian at the base of the coracoid.
“Type 4: attachment to the subclavian, but with a common root for both
the coracoid and thoracic.
“Type 5: attachment to the subclavian medial to both the axillary and
coracoid.
ARTERIES IN Two FAMILIES OF Birps 567
“Type 6: two separate thoracic arteries are present; the primary thoracic
is the same as type 1 above, while the secondary thoracic is the same as type 3
or type 4 above.”
Possibly the thoracic artery has undergone migration but ap-
parent differences in its origin might well be due to differences
in other vessels of the thoracic area. Additionally, there seems to
be no reason to assume that the lateral position of the thoracic
artery is the primitive one, or that the medial is the derived position,
as is implied by the phrase “medial migration.” Although the
lateral site of attachment (type 1) is predominant in the lower
orders of birds, and the medial attachment is found primarily in
Passeriformes, a fact which may indicate that type 1 is the more
primitive, it must nevertheless be kept in mind that a sequence of
a single morphological character does not necessarily represent the
phylogenetic sequence of the character itself (see Mayr, 1955:41).
Also, a given arterial arrangement might be independently de-
rived more than once. If such has been the case, similarities in
arterial arrangements in different taxa would sometimes be “chance
similarities,” that is to say, “resemblance in characteristics developed
in separate taxa by independent causes and without causal relation-
ship involving the similarity as such” (Simpson, 1961:79).
The particular arrangement of the arteries of the thoracic area
also seems to be of limited value as a clue to taxonomic relation-
ships. If the origin of any artery is determined by skeletal and
muscular features, as I suggest, the artery perhaps ought not be
considered as a separate character, but as part of a “character com-
plex” that varies as a unit (see Mayr, Linsley, and Usinger, 1953:
123). The skeleton offers a potential fossil record for consideration.
Changes in the skeleton and muscles, great enough to affect the
blood vessels, would probably be detected more easily than would
the resulting vascular changes. Also, I did not find as much in-
dividual variation in the skeleton and muscles in the area studied
as I did in the vascular system. In other words, within the bounds
established by the skeletal and muscular features, the artery still
exhibited individual variation in exact origin.
SUMMARY
The origin, distribution, and individual variation of the thoracic
and coracoid arteries, and their branches, have been studied in four
species of the family Columbidae (pigeons) and in five species of
the family Hirundinidae (swallows). These arteries are described
for Scardafella inca (Inca Dove) and Progne subis (Purple Martin).
Muscles that are supplied by these vessels, and muscles the partic-
568 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
ular configuration of which seems to effect the arrangement of the
arteries have also been described. Correlation of the arteries ob-
served with those named and described by other workers has been
attempted.
In most of the vessels studied there is a high degree of individual
variation, but few interspecific differences were noticed within
either family. Differences in the arteries of the thorax between
the two families are described by discussing the resulting different
origins of the thoracic artery. In swallows the thoracic artery arises
from either the subclavian artery or the coracoid artery, whereas
in pigeons it arises from the pectoral trunk. This difference in site
of attachment seems to be a result of differences between the two
families in muscular and skeletal elements of the thorax.
The particular site of attachment of the thoracic artery is of
limited value as a taxonomic character. Several considerations in-
fluenced this conclusion. (1) If the location of the artery is de-
termined by skeletal and muscular elements, these associated
structures must be considered taxonomically as a “character com-
plex” (a set of characters varying as a unit). (2) Even within the
bounds established by the skeleton and muscles, the artery dis-
plays a high degree of individual variation in exact origin. (8) A
given arterial arrangement could have been derived independently
many times. (4) Because differences are defined relative to other
likewise variable vessels, supposed similarities or differences in the
one artery may be artifacts of the system of description.
My findings and interpretations do not support previous sug-
gestions that the thoracic artery has undergone a mediad migration,
and that the various sites of attachment of that vessel may come to
represent various levels of evolution. The primitive site of attach-
ment of the vessel is unknown, and it seems to me that it has not
been sufficiently demonstrated that the vessel has undergone any
“migration.”
ARTERIES IN Two FAMILIES OF Brrps 569
M. supracoracoideus
M. coracobrachialis posterior
M. costi-sternalis anterior
Mm. intercostales extemmus
Fig. 1 M. costi-sternalis
Fic. 1. Progne subis. Lateral view of left half of thorax. M. pectoralis
thoracica (area of insertion indicated by dotted line) has been removed.
Muscles not described in text are not shown. (> 1.5.)
M. subcoracoideus
=
Ti
. Sx . sternocoracoideus
M. costi-sternalis anterior
eer M. costi-sternalis
Se
Fig. 2
Fic. 2. Progne subis. Lateral view of left half of thorax. Same view as shown
in Fig. 1, but with Mm. supracoracoideus, coracobrachialis posterior, and inter-
costales externus removed. (> 1.5.)
570 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
M. sternocoracoideus
M. costi-sternclis_/
anterior
Costo-pulmonary muscle_J
Mm. intercostales internus
seS:
Fic. 8. Progne subis. Medial view of left half of thorax. Not all
muscles shown. See Fig. 4 for identification of arteries. ( x 1.5.)
Fic. 4. Progne subis. Lateral view of left half of thorax. (x 1.5.)
(Applies also to Fig. 8.) 8. (Unnamed.) Supplies M. sternoco-
1. Subclavian artery. racoideus, M. coracobrachialis pos-
2. Coracoid artery. terior, and sternum.
3. Thoracic artery. 9. Pectoral artery.
4, (Unnamed.) Supplies M. 10. Axilliary artery.
coracobrachialis posterior. 11. (Unnamed.) Supplies M.
5. (Unnamed.) Supplies M. coracobrachialis posterior.
subcoracoideus. 12. (Unnamed.) Supplies M.
6. (Unnamed.) Supplies M. supracoracoideus.
sternocoracoideus and sternum. 18. (Unnamed.) Supplies M.
7. (Unnamed.) Supplies M. costi-sternalis anterior.
subcoracoideus. 14. Carotid artery.
ARTERIES IN Two FAMILIES OF Birps Sif
M. supracoracoideus
ic ts{__M. coracobrachialis posterior
M. sternocoracoideus
Fic. 5. Scardafella inca. Lateral view of left half of thorax. M.
pectoralis thoracica (area of insertion indicated by dotted line)
has been removed. Muscles not described in text are not shown.
See legend for Fig. 7 for identification of arteries. (xX 1.)
Mm. intercostales
externus
M. costi-sternalis
Fig. 6
( \ M. subcoracoideus
oe M. sternocoracoideus
T.S.
Fic. 6. Scardafella inca. Lateral view of left half of thorax. See
legend for Fig. 7 for identification of arteries. (x 1.)
572
UNIVERSITY OF Kansas Pus.s., Mus. Nat. Hist.
M. sternocoracoideus
M. costi-sternalis
Costo-pulmonary muscle
SSA = :
Zig” coe
gy
Fic. 7. Scardafella inca, Medial view of left half of thorax. (x 1.)
Key
(Applies also to Figs. 5 and 6.) Numerals not used are those used for Progne
CONG Ox Poor
i)
~»
subis for which no homologous artery occurs in Scardafella inca.
Subclavian artery.
Coracoid artery.
Thoracic artery.
(Unnamed.) Supplies Mm. coracobrachialis posterior and sternocoracoideus.
. (Unnamed.) Supplies M. coracobrachialis posterior.
Sternal artery. (Shown as it appears on right side. Left sternal artery not
so extensive. )
. (Unnamed.) Supplies M. sternocoracoideus.
(Unnamed.) Supplies M. subcoracoideus.
(Unnamed.) Supplies coracoid-sternal articulation.
Pectoral artery.
Axillary artery.
(Unnamed.) Supplies M. supracoracoideus.
Carotid artery.
(Unnamed.) Supplies M. supracoracoideus.
ARTERIES IN Two FAMILIES OF Brrps ae
LITERATURE CITED
AMERICAN ORNITHOLOGISTS’ UNION
1957. Check-List of North American birds. Baltimore, Maryland, Amer.
Ornith. Union, xiv + 691 pp.
ASHLEY, J. F.
1941. A study of the structure of the humerus in the Corvidae. Condor,
43:184-195.
BEDDARD, F. E.
1898. The structure and classification of birds. London, Longmans,
Green, & Co., xx + 548 pp.
BrErGer, A. J.
1956. Anatomical variation and avian anatomy. Condor, 58:433-441.
Buavurr, J. L., and Biswas, B.
1954. The main cervical and thoracic arteries of birds. Series 2, Colum-
biformes, Columbidae, part 1. Anat. Anz., 100:337-350.
Buapurl, J. L., Biswas, B., and Das, S. K.
1957. The arterial system of the domestic pigeon (Columba livia Gmelin).
Anat. Anz., 104:1-14.
Cougs, E.
1903. Key to North American birds. Vol. I, Fifth edit. Boston, The Page
Company, xlii + 535 + [55] pp.
Fisuer, H. I.
1955. Major arteries near the heart in the Whooping Crane. Condor,
57:286-289.
Genny, F. H.
1940. The main arteries in the region of the heart of three species of
doves. Bull. Fan Mem. Inst. Biol., Zool. ser., vol. 10, pt. 4, 271-278.
( Not seen. )
1955. Modifications of pattern in the aortic arch system of birds and their
phylogenetic significance. Proc. U. S. Nat. Mus., 104:525-621.
Howarp, H.
1929. The avifauna of Emeryville Shellmound. Univ. Calif. Publs. Zool.,
32:301-394.
HowE1., A. B.
1937. Morphogenesis of the shoulder architecture: Aves. Auk, 54:364-375.
Hupson, G. E., and LANZILLOTTI, P. J.
1955. Gross anatomy of the wing muscles in the family Corvidae. Amer.
Midl. Nat., 53:1-44.
Mayr, E.
1955. Comments on some recent studies of song bird phylogeny. Wilson
Bull., 67:33-44.
Mayr, E., Linstrey, E. G., and Usincerr, R. L.
1958. Methods and principles of systematic zoology. New York, McGraw-
Hill Book Co., x + 336 pp.
Smepson, G. G.
1961. Principles of animal taxonomy. New York, Columbia Univ. Press,
xiv -+ 247 pp.
Transmitted June 24, 1963.
29-8531
TOP NY DURAN oo
MUS. COMP. ZOOL
LIBRARY
tt]
kL ‘
UNIVERSITY OF KANSAS PUBLICATIONS
HARVARO
Museum oF Natura History jj pyensipy
Volume 12, No. 14, pp. 575-655, 10 figs.
May 18, 1964
The Breeding Birds of Kansas
BY
RICHARD F. JOHNSTON
UNIVERSITY OF KANSAS
LAWRENCE
ee" 1964
UNIVERSITY OF KANSAS PUBLICATIONS
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figures in’'text. December 10, 1955.
2. Additional records and extension of ranges: of mammals from. Utah. By
Stephen, D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80.
December 10, 1955.
8. 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. 125-835, 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. 387-346,
1 figure in text, I table. August 15,1956.
9. Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp.
847-351. August 15, 1956.
10. A new ‘bat (Genus Leptonycteris) from Coahuila. By Howard J. Stains.
Pp. 853-356.) 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. 863-387, 7 figures-in text. February. 21, 1958.
13. New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones,
Jr. Pp. 385-3888. May 12, 1958.
14. Pleistocene bats from San Josecito Cave, Nuevo Leén, México. 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 Sydney Anderson. Pp. 415-511, 12 figures in text, 2 tables.
August 1, 1959.
18. Conspecificity of two pocket mice, Perognathus goldmani and P. artus. - By
E. Raymond Hall and Marilyn Bailey Ogilvie.. Pp. 513-518, 1 map. Janu-
ary 14,.1960.
19. Records of harvest mice, Reithrodontomys, from Central America, with de-
scription of a new subspecies from Nicaragua. By Sydney Anderson and
J. Knox Jones, Jr.. Pp. 519-529. January 14, 1960.
20. Small carnivores from, San Josecito Cave (Pleistocene),. Nuevo Leédn, México.
By E..Raymond Hall: Pp. 531-538, 1 figure in text. January 14, 1960.
21. Pleistocene pocket gophers from San Josecito Cave, Nuevo Leén, México.
By Robert J. Russell. Pp. 589-548, 1 figure in text. January 14, 1960.
(Continued on inside of back cover)
MUS. COMP. ZOOL
LIBRARY,
HARVARD
UNIVERSITY OF KANSAS PUBLICATIONS...
MuSEUM OF NATURAL HIsTORY
Volume 12, No. 14, pp. 575-655, 10 figs.
May 18, 1964
The Breeding Birds of Kansas
BY
RICHARD F. JOHNSTON
UNIVERSITY OF KANSAS
LAWRENCE
1964
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HisToRY
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 12, No. 14, pp. 575-655, 10 figs.
Published May 18, 1964
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY
HARRY (BUD) TIMBERLAKE, STATE PRINTER
TOPEKA, KANSAS
1964
30-1476
The Breeding Birds of Kansas
BY
RICHARD F. JOHNSTON
CONTENTS
PAGE
ENPRODUCGTIONIE 4), ere. te icosee tcc Gis Sk Ae ae noe eae iid)
IDISTRIBUTION-OE, DIRDS IN, ICANSAS: ©. fossa 2 a ee 579
Avianvhabitatsum™ Kansas. 048. her onie Aes. oie ae 581
Species reaching distributional limits in Kansas ............ 584
IBREEDENG SEASONS) of uct, cities cocks «bad dere he tpn eee 588
REPOGUCHOMNG fied nck hocks cee ae ee eee 588
WarlatiOn inMbECeEGIN? SCASOMNS 2 4.0.5 sacs as oe see 589
POOCCOLTAPHIC CALE LOTICS, fs a j2e:, 8 Bact ii ee ae Re 593
Significance of phylogeny to breeding schedules ........... 595
Regulation’ of breeding schedules! 0). 47 5-h.<:0 490 ee eee 598
INGCOUNTES OR OPECUES§ 2000 cto 88, 25 chac 3 soul Sas caste oy IE 601
INCKNOWEEDG MENTS be 80 8. oP ot asc ses Huatie ss ¢ Sega oe ee 652
(ETRERA TURE MERI 2220 couse, Ahoy, athty. o,f chcoaaihacey fae ee 652
INTRODUCTION
The breeding avifauna of Kansas has received intermittent at-
tention from zoologists for about 75 years. Summary statements,
usually concerning ail birds of the state, have been published by
Goss (1891), Long (1940), Goodrich (1941), Tordoff (1956) and
Johnston (1960). All but the first dealt with the breeding birds
chiefly in passing, and none was concerned primarily with habitat
distributions and temporal characteristics of Kansan birds. The
present work treats mainly certain temporal relationships of breed-
ing birds in Kansas, but also geographic distribution, habitat prefer-
ences, and zoogeographic relationships to the extent necessary for
a useful discussion of temporal breeding phenomena.
Information on breeding of some of the 176 species of birds
known to breed in Kansas is relatively good, on a few is almost non-
existent, and on most is variously incomplete. It is nevertheless
possible to make meaningful statements about many aspects of the
breeding biology and distribution of most species of Kansan birds;
(577)
578 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
we can take stock, as it were, of available information and assess
the outstanding avenues of profitable future work. In the accounts
of species below, the information given is for the species as it
occurs in Kansas, unless it is otherwise stated. For the various
subsections analyzing biology and distribution, only information
taken in Kansas is used, and for this reason the analyses are made
on about half the species breeding in the state. An enormous
amount of observational effort has been expended by several dozen
people in order that suitable data about breeding birds of Kansas
be available; all persons who have contributed in any way are
listed in the section on acknowledgments, following the accounts
of species.
Kansas has been described topographically, climatically, and
otherwise ecologically many times in the recent past; the reader is
referred to the excellent account by Cockrum (1952), which treats
these matters from the viewpoint of a zoologist. For present pur-
poses it will suffice to mention the following characteristics of
Kansas as a place lived in by birds.
Topographically, Kansas is an inclined plane having an elevation
of about 4100 feet in the northwest and about 700 feet in the south-
east. West of approximately 97° W longitude, the topography is
gently rolling, low hills or flat plain; to the east the Flint Hills
extend in a nearly north to south direction, and to the east of these
heavily weathered, grassy hills is a lower-lying but more heavily
dissected country, hills of which show no great differences in ele-
vation from surrounding flatland.
The vegetation of eastern Kansas comingles with that of the
western edge of the North American deciduous forest; a mosaic
of true forest, woodland remnants, and tall-grass prairie occupies
this area east of the Flint Hills. From these hills west the prairie
grassland today has riparian woodland along watercourses; the
prairie is composed of proportionally more and more short-grass
elements to the west and tall-grass elements to the east.
Climate has a dominating influence on the vegetational elements
sketched above. Mean annual rainfall is 20 inches or less in western
sectors and increases to about 40 inches in the extreme eastern
border areas. Mean monthly temperatures run from 25°F. or 30°F.
in winter to 80°F. or 90°F. in summer. The northwestern edges of
Caribbean Gulf warm air masses regularly reach northward only
to the vicinity of Doniphan County, in northeastern Kansas, and
extend southwestward into west-central Oklahoma; these wet frontal
systems are usually dissipated along the line indicated by masses of
THE BREEDING Birps OF KANSAS 579
arctic air, sometimes in spectacular fashion. The regular recur-
rence of warm gulf air is responsible for the characteristically high
relative humidity in summer over eastern Kansas and it has an
ameliorating effect on winter climate in this region. Almost im-
mediately to the north in Nebraska and to the west in the high
plains, summers are dryer and winters are notably more severe.
The breeding distributions of some species of birds fairly closely
approximate the distribution of these warm air masses; these
examples are noted where appropriate below.
DISTRIBUTION OF BIRDS IN KANSAS
Birds breeding in Kansas are taxonomically, ecologically, and
distributionally diverse. Such diversity is to be expected, in view of
the midcontinental position of the State. Characteristics of insular-
ity, owing to barriers to dispersal and movement, tend to be lacking
in the makeup of the avifauna here. The State is not, of course,
uniformly inhabited by all 176 species (Table 1) of breeding birds;
most species vary in numbers from one place to another, and some
are restricted to a fraction of the State. Variations in numbers and
in absolute occurrence are chiefly a reflection of restriction or
absence of certain plant formations, which is to say habitats; the
analysis to follow is thus organized mainly around an examination
of gross habitat-types and the birds found in them in Kansas.
TABLE ].—THE BREEDING Binns oF KANSAS
Woodland Species
Elanoides forficatus N* Caprimulgus carolinensis N
Ictinia misisippiensis U C. vociferus U
Accipiter striatus U Phalaenoptilus nuttallii N
A. cooperii U Chaetura pelagica U
Buteo jamaicensis O Archilochus colubris N
B. lineatus N Colaptes auratus N
B. platypterus N C. cafer N
Aquila chrysaétos O Dryocopus pileatus O
Falco sparverius U Centurus carolinus N
Colinus virginianus N Melanerpes erythrocephalus N
Phasianus colchicus O Dendrocopos villosus O
Meleagris gallopavo N D. pubescens O
Philohela minor U Tyrannus tyrannus S
Zenaidura macroura N T. vociferans S
Ectopistes migratorius N Muscivora forficata S
Conuropsis carolinensis U Myiarchus crinitus S
Coccyzus americanus N Sayornis phoebe S
C. erythropthalmus N
Otus asio U
Bubo virginianus O
Strix varia U
Asio otus U
Aegolius acadicus U
Empidonax virescens S
Contopus virens S
Iridoprocne bicolor N
Progne subis N
Cyanocitta cristata N
Pica pica O
580 UNIVERSITY OF KAnsAs Pustis., Mus. Nat. Hist.
Corvus brachyrhynchos O Dendroica aestiva N
C. cryptoleucus O D. discolor N
Parus atricapillus O Seiurus motacilla N
P. carolinensis O Oporornis formosus N
P. bicolor O Icteria virens N
Sitta carolinensis O Wilsonia citrina N
Troglodytes aedon N Setophaga ruticilla N
Thryomanes bewickii N Passer domesticus O
Thryothorus ludovicianus N Icterus spurius N
Mimus polyglottos N I. galbula N
Dumetella carolinensis N I. bullockii N
Toxostoma rufum N Quiscalus quiscula N
Turdus migratorius O Molothrus ater N
Hylocichla mustelina N Piranga olivacea N
Sialia sialis O P. rubra N
Bombycilla cedrorum N Richmondena cardinalis S
Lanius ludovicianus O Pheucticus melanocephala §
Sturnus vulgaris O P. ludoviciana S
Vireo atricapillus N Guiraca caerulea S
V. griseus N Passerina ciris §
V. bellii N P. cyanea §
V. flavifrons N P. amoena §
V. olivaceus N Spinus pinus O
V. gilvus N S. tristis O
Mniotilta varia N Loxia curvirostra O
Protonotaria citrea N Pipilo erythrophthalmus N
Parula americana N Chondestes grammacus N
Spizella passerina N
Limnic Species
Podilymbus podiceps U Butorides virescens U
Phalacrocorax auritus U Florida caerulea U
Ardea herodias U Casmerodius albus U
Leucophoyx thula U Porzana carolina U
Nycticorax nycticorax U Laterallus jamaicensis U
Nyctanassa violacea U Gallinula chloropus U
Ixobrychus exilis U Fulica americana U
Botaurus lentiginosus U Charadrius alexandrinus U
Plegadis chihi U Actitis macularia U
Branta canadensis U Steganopus tricolor U
Anas platyrhynchos U Sterna albifrons U
A. acuta U Chlidonias niger U
A. discors U Telmatodytes palustris N
A. clypeata U Cistothorus platensis N
Aix sponsa U Geothlypis trichas N
Aythya americana U Xanthocephalus xanthocephalus N
Oxyura jamaicensis U Agelaius phoeniceus N
Rallus elegans U Rallus limicola U
Grassland Species
Buteo swainsonii N Asio flammeus U
B. regalis U Sayornis saya S
Circus cyaneus O Eremophila alpestris O
Tympanuchus cupido N Dolichonyx oryzivorus N
T. pallidicinctus N Sturnella magna N
Pedioecetes phasianellus N S. neglecta N
Charadrius vociferus U Spiza americana N
Eupoda montana U Calamospiza melanocorys N
Numenius americanus U Ammodramus savannarum N
Bartramia longicauda U Passerherbulus henslowii N
Speotyto cunicularia U Aimophila cassinii N
Spizella pusilla N
THE BREEDING Birps oF KANSAS 581
Xeric Scrub Species
Callipepla squamata N Geococcyx californianus N
Salpinctes obsoletus N
Unanalyzed Species
Cathartes aura N Chordeiles minor U
Coragyps atratus N Megaceryle alcyon U
Falco peregrinus U Riparia riparia O
Columba livia O Stelgidopteryx ruficollis N
Tyto alba U Hirundo rustica O
Petrochelidon pyrrhonota U
® The letter following each name refers to presumed zoogeographic derivation of the
species, modified after Mayr (1946). N = North American evolutionary stock; S = South
American stock; O = Eurasian stock; U = unanalyzed.
Avian Habitats in Kansas
Four major habitat-types can be seen in looking at the distribu-
tion of the breeding avifauna of Kansas. These are woodland,
grassland, limnic, and xeric scrub plant formations. A little more
than half the breeding birds of Kansas live in woodland habitats,
about one-fifth in limnic habitats, about one-eighth in grassland
habitats, and less than two per cent in scrub habitats; this leaves
some 6.4 per cent of the breeding avifauna unanalyzed (Table 2).
Woodland Habitats
One hundred one species of Kansan birds are woodland species
(tables 1 and 2). The analysis of Udvardy (1958) showed wood-
land birds to be the largest single avifaunal element in North
America, with 38 per cent of North American birds relegated to it.
It is likewise the largest element in the Kansan avifauna, represent-
ing 58 per cent of Kansan birds. Although woodland makes up
TABLE 2.—ANALYSIS OF THE BREEDING AVIFAUNA OF KANSAS BY HABITAT-TYPES
Percentage of the Avifauna of
HABITAT-TYPE |
North Stated
Kansas America habitat
Woodlands 101 species:.. ... 25 5-26-27 58 16.7 44.4
ImMMICsySOISPCCIES rete we) es sees eee 21 6.0 38.5
Grasslandcs25 SpeClesste. Ac ys. ce esti 13 3.8 (Ales)
DRCTIC STUDS: SISPECIES). dia his 2 s.cta/i< wisiees) 3 = 2 0.5 10.2
Unanalyzeds Miispectes'.2 2 =... ot. ces siers 6 2.0 55.0
Totals WA species. 0.2 ciad5 1s asa a: 100 29.0 43.2
1. Does not include the Canvasback (Aythya valisineria), the Forster Tern (Sterna
forsteri), and the Black Tern (Chlidonias niger), all recently added to the breeding avi-
fauna of Kansas.
582 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
a relatively small fraction of the vegetational complexes in Kansas,
a large number of habitats exist in what woodland is present. An
even larger number of possible woodland habitats is clearly missing,
however, because the 101 Kansan species actually represent but 44
per cent of all woodland birds in North America, according to
Udvardy’s analysis. Broad-leaved, deciduous woodlands in Kansas
are of restricted horizontal and vertical stratification. More com-
plex deciduous forest associations and all coniferous forest associa-
tions are absent from the State.
Using Mayr’s (1946) breakdown of geographical origin of the
North American bird fauna, about 53 per cent of the woodland
passerine birds in Kansas are of “North American” origin, 22 per
cent are of “Eurasian” origin, and 14 per cent are of “South Ameri-
can” origin (Table 3). These figures for Kansas are commensurate
with those found for other geographic districts at the same latitude
TABLE 3.—ANALYSIS OF EcoLocic Groups OF Birps BY STATUS OF RESIDENCY
AND AREA OF ORIGIN
3
aS} 8
Be ee al ee) ar eae
S as] > = & a 5
Sa ene: ate = Es
= om Ay S) Zi wh =)
Me OTSA Te erties 60% | 29% | 11% | 22% | 58% | 14% | 11%
Limnic species wate 94% | 0 | 6%] 0 114%] 0 | 86%
Grassland species on ane 61% | 26% | 13% | 9% | 56% | 3% | 30%
Bit fy)
Xeric Sor species........ 33% | 66% 0 0 {100% 0 0
‘270
Unanalyzed species........ 64% | 27% | 9% | 26% | 26% 0 48%
11s (a)
in North America (Mayr, 1946:28). Other characteristics of wood-
land birds are summarized in tables 4 and 5.
Limnic Habitats
Of Kansan birds, 36 species (20 per cent) prefer limnic habitats
(Table 1). Udvardy found this group to represent 15 per cent
of the North American avifauna. Kansas is not notably satisfactory
for limnic species, and only 38 per cent of the total North American
limnic avifauna is present in the State.
Thirty-one species of limnic birds belong to families that Mayr
(1946) considered to be unanalyzable as to their geographic origin;
of the five remaining species, all seem to be of North American
THE BREEDING Birnps OF KANSAS 583
origin. Other characteristics of limnic birds are summarized in
tables 4 and 5.
Grassland Habitats
Twenty-three species of our total can be called grassland species
(Table 1). The subtotal is less than one-fifth of the Kansan avi-
TABLE 4.—ANALYSIS BY HABITAT-TYPE AND RESIDENCY STATUS OF HISTORIC
AVIAN STOCKS IN KANSAS
—
:
2 a 5
SS re} = a —
E So iiag ss (RSE eee
=! = S wR a a o >
S SI z 2 = z sn eee
5 g 3 B S oh a &
S 42 a o =) ‘= oe 3
eS 4 ) ra =) = font a
Old World Element..} 80% 0 8% 0 12%) 1% | (8G ALG
27:16%
North American
Element........ 69% | 6% | 17% | 4% | 4% | 72% | 14% | 14%
77:44%
South American
eae ii Os ee 93% 0 7% 0 0 93% | 7% 0
23°70
Unanalyzed Origin. .| 22% | 56% | 138% 0 9% | 79% | 16% | 5%
53:32%
fauna, but it represents 72 per cent of the grassland birds of North
America; grassland habitats abound in Kansas. Only 5.3 per cent
of all North American birds are grassland species (Udvardy, 1958).
About 56 per cent of these birds are of North American stocks,
nine per cent of Eurasian stocks, and three per cent of South Ameri-
can stocks. The percentage of North American species is the
greatest for any habitat group here considered. Other character-
istics of grassland birds are summarized in tables 4 and 5.
Xeric-Scrub Habitats
Three species of Kansan birds can be placed in this category
(Table 1). This is less than one per cent of the North American
avifauna, two per cent of the Kansan avifauna, and ten per cent
of the birds of xeric scrub habitats in North America. The three
species are considered to be of North American origin.
Unanalyzed as to Habitat
Eleven species of Kansan birds could not be assigned to any of
the habitat-types mentioned above. The total represents two per
084 UNIVERSITY OF KANSAS Pusts., Mus. Nar. Hist.
TaBLe 5.—ANALYsSIS BY EcoLocic STATUS AND AREA OF ORIGIN OF MIGRANT
AND RESIDENT Birps
Woodland
North America
Grassland
Xeric Scrub
Unanal. Hab.
Old World
South America
Unanalyzed
Migrant
: 9 ne
eS 52% | 29% | 12% 1% | 6% | 2% | 49% | 12% | 37%
Resident
piese ae 73% 0 15% | 5% | 7% | 51% | 26% | 2% | 21%
Partly
migrant. .| 64% | 11% | 17% 0 6% | 17% | 66% 0 17%
17:10%
cent of the North American avifauna, six per cent of the birds of
Kansas, and 55 per cent of the species reckoned by Udvardy (loc.
cit.) to be unanalyzable. Fifty-five per cent is a large fraction, but
only to be expected: species are considered unanalyzable if they
show a broad, indiscriminate use of more than one habitat-type,
and such birds tend to be widely distributed.
Species Reaching Distributional Limits in Kansas
The distributional limits of a species are useful in indicating
certain of its adaptive capacities and implying maintenance of or
shifts in characteristics of habitats. Although it is generally an
oversimplification to ignore abundance when treating of distribu-
tion, the present remarks of necessity do not pertain to abundance.
Western Limits Reached in Kansas
Thirty-one species (tables 6 and 7) reach the western limits of
their distribution somewhere in Kansas. Most of these limits are
in eastern Kansas, and coincide with the gradual disappearance of
TABLE 6—BREEDING Braps REACHING DisTRIBUTIONAL Limits IN KANSAS
Species reaching northern distributional limits
Florida caerulea Geococcyx californianus
Leucophoyx thula Caprimulgus carolinensis
Coragyps atratus Muscivora forficata
Elanoides forficatus Parus carolinensis
Ictinia misisippiensis Vireo atricapillus
Tympanuchus pallidicinctus Passerina ciris
Callipepla squamata Aimophila cassinii
THE BREEDING Birps of KANSAS 585
Species reaching southern distributional limits
Aythya americana
Parus atricapillus
Bombycilla cedrorum
Dolichonyx oryzivorus
Pedioecetes phasianellus
Empidonax minimus
Steganopus tricolor
Chlidonias niger
Coccyzus erythropthalmus
Species reaching eastern distributional limits
Eupoda montana
Numenius americanus
Phalaenoptilus nuttallii
Colaptes cafer
Tyrannus verticalis
Sayornis saya
Corvus cryptoleucus
Salpinctes obsoletus
Icterus bullockii
Pheucticus melanocephalus
Passerina amoena
Species reaching western distributional limits
Aix sponsa Hylocichla mustelina
Buteo platypterus Vireo griseus
Philohela minor V. flavifrons
Ectopistes migratorius Mniotilta varia
Protonotaria citrea
Parula americana
Dendroica discolor
Seiurus motacilla
Oporornis formosus
Wilsonia citrina
Setophaga ruticilla
Conuropsis carolinensis
Chaetura pelagica
Archilochus colubris
Dryocopus pileatus
Centurus carolinus
Myiarchis crinitus
Empidonax virescens
E. traillii Sturnella magna
Parus bicolor Piranga olivacea
Thryothorus ludovicianus Pheucticus ludovicianus
Cistothorus platensis Pipilo erythrophthalmus
Passerherbulus henslowii
the eastern deciduous forest formation. Twenty-nine species are
woodland birds, and few of these seem to find satisfactory condi-
tions in the riparian woods extending out through western Kansas.
The Wood Thrush is the one woodland species that has been found
nesting in the west (Decatur County; Wolfe, 1961). Descriptively,
therefore, the dominant reason for the existence of distributional
limits in at least 28 of these birds is the lack of suitable woodland
in western Kansas; these 28 are the largest single group reaching
distributional limits in the State. Many other eastern woodland
birds occur in western Kansas along riparian woodlands, as is
mentioned below.
Two species showing western limits in Kansas are characteristic
of grassland habitats; the Eastern Meadowlark seems to disappear
with absence of moist or bottomland prairie grassland and the
Henslow Sparrow may be limited westerly by disappearance of
tall-grass prairie.
The Short-billed Marsh Wren, a marginal limnic species, reaches
its southwesterly mid-continental breeding limits in northeastern
586 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
Kansas. The species breeds in Kansas in two or three years of each
ten, in summers having unusually high humidity.
Northern Limits Reached in Kansas
Fourteen species (tables 6 and 7) reach their northern distribu-
tional limits in Kansas. Eight of these are birds of woodland habi-
tats, but of these only the Carolina Chickadee is a species of the
eastern deciduous woodlands; the other seven live in less mesic
woodland. Three of these species (Chuck-will’s-Widow, Scissor-
tailed Flycatcher and Painted Bunting) have breeding ranges that
suggest the northwesterly occurrences of summer humid warm air
masses (“gulf fronts”) and this environmental feature perhaps is
of major importance for these birds, as it is also for the vegetational
substratum in which the birds live.
The Lesser Prairie Chicken and the Cassin Sparrow are the two
birds of grasslands that are limited northerly in Kansas. Xeric,
sandy grassland is chiefly limited to the southwestern quarter of
Kansas, and this limitation is perhaps of major significance to these
TABLE 7.—ANALYSIS BY HABITAT-TYPE OF Birps REACHING DISTRIBUTIONAL
LIMITS IN KANSAS
Habitat-types
DIRECTIONAL ===
LIMIT .
Woodland | Grassland | Limnic oe Total
Western extent..... 28 2 1 0 31
Northern extent... . 8 2 2 2 14
Eastern extent... .. 6 4 0 1 11
Southern extent. ... 4 2 3 0 9
MRotalsse- a. esses 46 10 6 3 65
Per cent of the Spe-
cies in Stated
Habitathna ea: 46 43 14 100 By
two species. The Scaled Quail and Roadrunner tend to drop out
as the xeric “desert scrub” conditions of the southwest drop out
in Kansas.
Eastern Limits Reached in Kansas
Eleven species (tables 6 and 7) reach their eastern distributional
limits in Kansas. Six of these are woodland birds. Four of these
are members of well-known species-pairs: the Red-shafted Flicker,
Bullock Oriole, Black-headed Grosbeak, and Lazuli Bunting. Pres-
ence to the east of complementary species has much to do with the
THE BREEDING Birps OF KANSAS 587
absence of these species in eastern Kansas. Four of the eleven are
birds of grasslands, and they drop out as the short-grass prairie
is restricted easterly.
The Rock Wren may be considered characteristic of xeric scrub
in Kansas, and it is not found to the east in the absence of such
scrub.
Southern Limits Reached in Kansas
Eight species (tables 6 and 7) reach their southern distributional
limits in Kansas. Half of these birds are of woodland habitats,
and of these four, the Black-capped Chickadee and Cedar Waxwing
are chiefly of sub-boreal distribution. The Black-capped Chickadee
also finds its niche partly pre-empted in southern Kansas by the
Carolina Chickadee.
The Bobolink and Sharp-tailed Grouse are grassland species that
are seemingly adapted to cooler, dryer grassland than is found in
most of Kansas.
The Redhead, Wilson Phalarope, and Black Tern are limnic
species, perhaps limited southerly by high summer temperatures;
the three species are entirely marginal anywhere in Kansas.
Influence of Riparian Woodland
Although the largest single element of the Kansan avifauna that
reaches distributional limits in Kansas is made up of birds of the
eastern deciduous forest, several species of the eastern woodlands
are present in Kansas along the east-west river drainages in riparian
woodland; the species are listed in Table 8. Twenty-one kinds are
involved if we include the Cooper Hawk, Yellow-billed Cuckoo,
Orchard Oriole, Summer Tanager, Rufous-sided Towhee, and Chip-
ping Sparrow, all of which breed farther to the west but are present
in western Kansas only along river drainages. This leaves 15 species
of eastern deciduous woodlands that occur west in Kansas along
TABLE 8.—Brimaps OF THE EASTERN Decipuous ForEsT FOUND IN WESTERN
KANSAS IN RIPARIAN WOODLAND
Accipiter cooperii * Toxostoma rufum
Coccyzus americanus * Sialia sialis
Centurus carolinus Vireo olivaceus
Melanerpes erythrocephalus Icterus spurius *
Tyrannus tyrannus Icterus galbula
Myiarchus crinitus Quiscula quiscalus
Contopus virens Piranga rubra *
Sayornis phoebe Passerina cyanea
Cyanocitta cristata Richmondena cardinalis
Dumetella carolinensis Pipilo erythrophthalmus *
Spizella passerina *
® Breeds farther west in North America in other types of vegetation.
588 UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist.
riparian woodland (versus 30 species that drop out chiefly where
eastern woodland drops out). These 15 species are about one-third
of all woodland birds in western Kansas. Riparian woodland does
not seem to afford first-rate habitat for most of the eastern woodland
species that do occur; breeding density seems to be much lower
than in well-situated eastern woodland.
The importance of these linear woodlands as avenues for gene-
flow between eastern and western populations, especially of species-
pairs (grosbeaks, flickers, orioles, and buntings), is obviously great.
Likewise significant is the existence of these alleys for dispersal
from the west of certain species (for instance, the Black-billed
Magpie and the Scrub Jay) into new but potentially suitable areas.
BREEDING SEASONS
introduction
An examination of breeding seasons or schedules is properly
undertaken at several levels. The fundamental description of varia-
tion in breeding schedules must itself be detailed in several ways
and beyond this there are causal factors needing examination. The
material below is a summary of the information on breeding sched-
ules of birds in Kansas, treated descriptively and analytically in
ways now thought to be of use.
Almost any event in actual reproductive activity has been used in
the following report; nestbuilding, egg-laying, incubation, brooding
of young, feeding of young out of the nest are considered to be of
equal status. To any such event days are added or subtracted from
the date of observation so as to yield the date when the clutch
under consideration was completed.
Such corrected dates can be used in making histograms that show
the time of primary breeding activity, or the “egg-season.” All such
schedules are generalizations; data are used for a species from any
year of observation, whether 50 years ago or less than one year ago.
One advantage of such procedure is that averages and modes are
thus more nearly representative of the basic temporal adaptations
of the species involved, as is explained below.
When information on the schedule of a species from one year
is lumped with information from another year or other years, two
(and ordinarily more than two) frequency distributions are used
to make one frequency distribution. The great advantage here is
that the frequency distribution composed of two or more frequency
distributions is more stable than any one of its components. Second,
the peak of the season, the mode of egg-laying, is represented more
THE BREEDING Birps oF KANSAS 589
broadly than it would have been for any one year alone. Third,
the extremes of breeding activity are fairly shown as of minute
frequency and thus of limited importance, which would not be true
if just one year were graphed. All these considerations combine
to support the idea that general schedules in fact represent the
basic temporal adaptations of a species much better than schedules
for one year only.
Variation in Breeding Seasons
In the chronology of breeding seasons of birds, there are three
basic variables: time at which seasons begin, time at which seasons
end, and time in which the major breeding effort occurs. These
variables have been examined in one population through time
(Lack, 1947; Snow, 1955; Johnston, 1956) in several populations
of many species over wide geographic ranges (Baker, 1938; Moreau,
1950; Davis, 1953), and in several populations of one species (Lack,
loc. cit.; Paynter, 1954; Johnston, 1954). The analysis below is
concerned with breeding of many kinds of birds of an arbitrarily
defined area and with the influence of certain ecologic and zoo-
geographic factors on the breeding seasons for those several species.
THE INFLUENCE OF SEASONAL StaTus.—Here we are interested in
whether a species is broadly resident or migrant in Kansas; 70
species are available for analysis.
Resident Species
Twenty-four species, furnishing 875 records of breeding, are here
considered to be resident birds in northeastern Kansas. These
species are Cooper Hawk, Red-tailed Hawk, Prairie Chicken, Bob-
white, Rock Dove, Great Horned Owl, Red-bellied Woodpecker,
Hairy Woodpecker, Downy Woodpecker, Horned Lark, Blue Jay,
Common Crow, Black-billed Magpie, Black-capped Chickadee,
Tufted Titmouse, Carolina Wren, Bewick Wren, Mockingbird,
Eastern Bluebird, Loggerhead Shrike, Starling, House Sparrow,
Eastern Meadowlark, and Cardinal. The distribution of completed
clutches (Fig. 1) runs from mid-January to mid-September, with a
modal period in the first third of May. Conspicuous breeding
activity occurs from mid-April to the first third of June.
Migrant Species
Forty-six species, furnishing 2,522 records of breeding, are con-
sidered to be migrant in northeastern Kansas. These species are
Great Blue Heron, Green Heron, Swainson Hawk, American Coot,
590 UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist.
Killdeer, Upland Plover, American Avocet, Least Tern, Yellow-
billed Cuckoo, Black-billed Cuckoo, Burrowing Owl, Common
Nighthawk, Chimney Swift, Red-headed Woodpecker, Eastern
Kingbird, Western Kingbird, Scissor-tailed Flycatcher, Great
Crested Flycatcher, Eastern Phoebe, Eastern Wood Pewee, Bank
Swallow, Rough-winged Swallow, Barn Swallow, Purple Martin,
Brown Thrasher, Catbird, House Wren, Robin, Wood Thrush, Blue-
gray Gnatcatcher, Bell Vireo, Warbling Vireo, Prothonotary
Warbler, Yellow Warbler, Chat, Western Meadowlark, Red-winged
Blackbird, Orchard Oriole, Baltimore Oriole, Common Grackle,
Black-headed Grosbeak, Indigo Bunting, Dickcissel, Lark Sparrow,
and Field Sparrow. The distribution of completed clutches runs
from mid-March to the first third of September, with a modal period
of egg-laying in the first third of June (Fig. 1). Conspicuous breed-
ing activity occurs from the first third of May to the last third of
June.
THE INFLUENCE OF DOMINANT ForRAGING ADAPTATION.—Five cate-
gories here considered reflect broad foraging adaptation: woodland
species, taking invertebrate foods in the breeding season from
woody vegetation or the soil within wooded habitats; grassland
species, taking invertebrate foods in the breeding season from
within grassland situations; limnic species, foraging within marshy
or aquatic habitats; aerial species, foraging on aerial arthropods;
raptors, feeding on vertebrates or large insects.
Raptors
Six species, furnishing 174 records of breeding, are here con-
sidered, as follows: Cooper Hawk, Red-tailed Hawk, Swainson
Hawk, Great Horned Owl, Burrowing Owl, and Loggerhead Shrike.
The distribution of clutches (Fig. 1) runs from mid-January to the
first third of July and is bimodal. One period of egg-laying occurs
in mid-February and a second in the last third of April. Such a
distribution indicates that two basically independent groups of
birds are being considered. The first peak of laying reflects activi-
ties of the large raptors, and the second peak is that of the insectiv-
orus Burrowing Owl and Loggerhead Shrike. The peak for these
two birds is most nearly coincident with that for grassland species,
a category to which the Burrowing Owl might well be relegated.
Limnic Species
Six species, the Great Blue Heron, Green Heron, American Coot,
American Avocet, Least Tern and Red-winged Blackbird, furnish
THE BREEDING Birps OF KANSAS
wan. Feb. Mch. Apr. May, June July jAu gSep. Oct. jNov. Dec.,
Raptorial Species
174
Limnic Species
264
Grassland Species
404
Aerial Insectivores
287
Woodland Species
i882
Migrant Species
2022
Resident Species
879
Old World Species
306
North American Species
12335
30 w
Fa} ;
20y 5 South American Species
Ju
.-¢
Oo. 308
oOo
o sx temperature
80 en ee 7, precipitation
_— OS
60 °F DF i en,
a Ta
40 A ES
Jan. Feb. Mch. Apr. May June July Aug. Sep. Oct. Nov.Dec.
591
inches
Fic. 1.—Histograms representing breeding schedules of tea categories
of Kansan birds. Heights of columns indicate percentage of total of
clutches of eggs, and widths indicate ten-day intervals of time, with the
5th, 15th, and 25th of each month as medians. The occurrences of
monthly means of temperature and precipitation are indicated at the
bottom of the figure.
2—1476
592 UNIVERSITY OF KANSAS PusLs., Mus. Nar. Hist.
264 records of breeding. The distribution of clutches (Fig. 1)
runs from mid-March to the last third of July and is bimodal. This
is another heterogeneous assemblage of birds; the Great Blue Heron
is responsible for the first peak, in the first third of April. The other
five species, however, show fair consistency and their peak of egg-
laying almost coincides with peaks for aerial foragers, woodland
species, and migrants, considered elsewhere in this section.
Grassland Species
Ten species, Greater Prairie Chicken, Bobwhite, Killdeer, Upland
Plover, Horned Lark, Starling, Eastern Meadowlark, Western Mead-
owlark, Common Grackle, and Dickcissel, furnish 404 records of
breeding activity. The distribution of clutches (Fig. 1) runs from
the first of March to mid-September. The peak of egg-laying occurs
in the first third of May. This is coincident with the peak for
resident species, perhaps a reflection of the fact that half the species
in the present category are residents in northeastern Kansas.
Woodland Species
In this category are included species characteristic of woodland
edge. Thirty-four species, furnishing 1,882 records of breeding,
are here treated: Yellow-billed Cuckoo, Black-billed Cuckoo,
“flicker” (includes birds thought to be relatively pure red-shafted,
pure yellow-shafted, as well as clear hybrids), Red-bellied Wood-
pecker, Red-headed Woodpecker, Hairy Woodpecker, Downy
Woodpecker, Blue Jay, Black-billed Magpie, Common Crow, Black-
capped Chickadee, Tufted Titmouse, Carolina Wren, Bewick Wren,
House Wren, Brown Thrasher, Catbird, Mockingbird, Robin, Wood
Thrush, Eastern Bluebird, Blue-gray Gnatcatcher, Bell Vireo,
Warbling Vireo, Prothonotary Warbler, Yellow Warbler, Chat,
Orchard Oriole, Baltimore Oriole, Cardinal, Black-headed Grosbeak,
Indigo Bunting, Lark Sparrow, and Field Sparrow. The distribu-
tion of clutches runs from the first third of March to mid-September
(Fig. 1). The modal period for completed clutches is the first
third of June. Conspicuous breeding activity occurs from the first
third of May to mid-June. The distribution of the season in time
is almost identical with that for migrant species, reflecting the large
number of migrant species in woodland habitats in Kansas.
Aerial Foragers
Twelve species, Common Nighthawk, Chimney Swift, Eastern
Kingbird, Western Kingbird, Scissor-tailed Flycatcher, Great
Crested Flycatcher, Eastern Phoebe, Eastern Wood Pewee, Bank
Tue BREEDING Birps OF KANSAS 593
Swallow, Rough-winged Swallow, Barn Swallow, and Purple Martin,
furnish 587 records of breeding. The distribution of clutches
(Fig. 1) extends from the last third of March to the first third of
August, and the modal date of clutches is in the first third of June.
Conspicuous breeding activity occurs from the end of May to the
end of June. The peak of nesting essentially coincides with that
characteristic of migrants.
Zoogeographic Categories
Three categories of Mayr (1946) are of use in analyzing trends
in breeding schedules of birds in Kansas. These categories of pre-
sumed ultimate evolutionary origin are the “Old World Element,” the
“North American Element,” and the “South American Element.”
Not always have I agreed with Mayr’s assignments of species to
these categories, and such differences are noted. There is some
obvious overlap between these categories and those discussed
previously.
Old World Element
Eighteen species, Red-tailed Hawk, Rock Dove, Great Horned
Owl, Hairy Woodpecker, Downy Woodpecker, Black-billed Magpie,
Common Crow, Black-capped Chickadee, Tufted Titmouse, Robin,
Loggerhead Shrike, Starling, House Sparrow, Bank Swallow, Barn
Swallow, and Blue-gray Gnatcatcher, furnish 969 records of breed-
ing (Fig. 1). Species for which I have records but which are not
here listed are the Blue Jay and the Wood Thrush, both of which I
consider to be better placed with the North American Element. The
distribution of completed clutches runs from mid-January to the
first third of August, and shows a tendency toward bimodality. The
second, smaller peak is due to the inclusion of relatively large sam-
ples of three migrant species (Robin, Bank Swallow, and Barn
Swallow). The timing of the breeding seasons of these three species
is in every respect like that of most other migrants; if they are re-
moved from the present sample the bimodality disappears, indicat-
ing an increase in homogeneity of the unit.
North American Element
Twenty-six species, Greater Prairie Chicken, Bobwhite, “flicker,”
Rough-winged Swallow, Purple Martin, Blue Jay, Carolina Wren,
Bewick Wren, House Wren, Mockingbird, Catbird, Brown Thrasher,
Wood Thrush, Bell Vireo, Warbling Vireo, Prothonotary Warbler,
Yellow Warbler, Chat, Eastern Meadowlark, Western Meadowlark,
Red-winged Blackbird, Orchard Oriole, Baltimore Oriole, Common
594 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Grackle, Lark Sparrow, and Field Sparrow, furnish 1,233 records
of breeding (Fig. 1). The distribution of completed clutches runs
from the first third of April to the first third of September. The
modal date for completion of clutches is June 1.
South American Element
Twelve species, Eastern Kingbird, Western Kingbird, Scissor-
tailed Flycatcher, Great Crested Flycatcher, Yellow-bellied Fly-
catcher, Traill Flycatcher, Eastern Wood Pewee, Eastern Phoebe,
Cardinal, Black-headed Grosbeak, Rose-breasted Grosbeak, and
Indigo Bunting, furnish 552 records of breeding (Fig. 1). The
curve representing this summary schedule is bimodal, wholly as a
result of including the Eastern Phoebe and the Cardinal with this
sample.
Relationship of Schedules to Temperature and Precipitation
In outlining the ten categories above, attention has been given to
certain similarities and differences in the frequency distributions. A
slightly more refined way of comparing the frequency distributions
is to relate them to other, seasonally variable phenomena. Figure 1
shows the frequency distributions of egg-laying of these ten cate-
gories of birds in terms of the regular changes in mean temperature
and mean precipitation characteristic of the environments in which
these birds live in the breeding season.
Table 9 shows that there are two basic groups of birds according
to peak of egg-laying and incidence of precipitation; raptors, birds
of Eurasian origin, resident birds, and birds of grassland habitats
tend to have their peaks of egg-laying prior to the peak of spring-
summer rains, and the other six categories tend to have their peaks
of egg-laying occur in the time of spring-summer rains. Regarding
temperature, there are four categories of birds; these are evident in
the table.
Some of the correspondences deserve comment. Residents and
grassland species both breed before the rains come and before mean
temperatures reach 70°F., and this correspondence probably results
from most of the grassland species being residents. Contrariwise,
most birds of Eurasian stocks are residents, but not all residents are
of such stocks; the two groups are discrete when mean temperature
at breeding is considered. Woodland birds, aerial foragers, and
birds of South American evolutionary stocks breed after tempera-
tures surpass 70°F. on the average. Almost all such species are mi-
grants, but many migrants have different temporal characteristics,
and the categories thus are shown to be discrete on the basis of
temperature at time of breeding.
THE BREEDING Birps OF KANSAS 595
The change through spring and summer of temperature and
precipitation delineates the inception and waxing of the growing
season of vegetation and of the subsequent arthropod populations,
on which most of the birds feed in the breeding season. The tem-
poral characteristics of growing seasons in North America have been
treated by Hopkins (1938) and have been related to timing of breed-
ing seasons in Song Sparrows (Passerella melodia) of the Pacific
coast of North America (Johnston, 1954).
Significance of Phylogeny to Breeding Schedules
Evidence from a variety of sources demonstrates that timing of
breeding seasons is either broadly or specifically genetically-de-
termined. For some species in some situations major environmental
variables are paramount in regulating timing of breeding, but in
others the innate, regulatory “clock” is less closely tied to conspicu-
ous exogenous stimuli. The work by Miller (1955a, 1955b, 1960)
with several species of Zonotrichia strongly indicates that endoge-
nous timing is most important for these birds, and there is ecological
evidence for Song Sparrows that supports the same point (Johnston,
1954, 1956). It is, in any event, possible to treat breeding schedules
as species-specific characters, for any one geographic area.
In an attempt to relate a breeding schedule to previous ancestral
modes, that is by extension to phylogeny, it is necessary to know
how often ancestral adaptations can persist in the face of necessity
to adapt to present environmental conditions. It is necessary to
know how conservative or how immediately plastic breeding sched-
ules can be. The disadvantage of using available information about
configurations of breeding seasons (as shown in Figs. 3 to 9) is
that it is extremely difficult to compare visually at one time more
than six or eight histograms as to the trenchant similarities and
differences regarding times of inception and cessation of breeding,
and time of peak egg-laying. It is possible, however, to reduce
these three variables to one variable (as described below), which
allows the necessary comparisons to be made more easily; this
variable may be called the breeding index.
Calculation of Breeding Index
The chronological year is broken roughly into ten-day intervals
numbered 1 to 36. The histogram describing the temporal occur-
rence of the breeding season of a species in our area usually will
lie within intervals 7 to 25. The modal date for completion of
clutches is given a value corresponding to the number of ten-day
396 UNIVERSITY OF KANsAsS Pusis., Mus. Nat. Hist.
intervals beyond interval 7 (March 1-10); this describes the modal
variable. The date of completion of 83 per cent of all clutches is
given a value corresponding to the number of ten-day intervals it
lies from interval 11 (April 11-20); this describes the 83 per cent
variable (and is a measure of the length of the season in terms of
its inception). The breeding index can then be calculated as
follows:
= Xie tXaas
where: I is the breeding index,
Xm is the modal variable, and
Xsa is the 83 per cent variable.
This is obviously an arbitrary scheme to gain a simple measure of
beginning, peak, and end of a breeding season. Other schemes
could be devised whereby different absolute values would be in-
volved, but the relative nature of the results would be preserved.
The values under the present system for 73 species of Kansan birds
run from —5 to + 22; early modal dates and cessation to breeding
give low values, late dates high values.
Within this framework there are other, presumably subordinate,
factors that influence the values of breeding indices, as follows:
1. Migratory habit. Any migrant tends to arrive on breeding
grounds relatively late, hence migrants ordinarily have higher
index values than do residents.
2. Colonial breeding. The strong synchrony of colonially-breed-
ing species tends to move the modal egg-date toward the time of
inception of breeding; as a result colonially-breeding species prob-
ably have lower index values than they would have if not colonial.
3. Single-broodedness. Species having only one brood per
season tend to have shorter seasons than double-brooded species,
and their index values tend to be lower than those of double-
brooded species.
Migratory habit unquestionably has considerable influence on
index values in some species. It is not, however, as important as
other matters, such as the condition of the food substratum or
sensitivity of the pituitary-gonadal mechanism, in determining
timing and mode of breeding activity. The schedule of the Purple
Martin is the extreme example showing that time of spring arrival
on breeding grounds is not necessarily related to time of inception
of breeding. It should be emphasized that the factors leading to
northward migratory movement may be involved in timing of the
annual gonadal and reproductive cycle.
Figure 2 presents a graphic summary of values of breeding
THE BREEDING Binps OF KANSAS 597
Breeding Index
ONE 4G SENIOL 2 a4 Borlss2O eZ
PO
Ardeidae SSS
Accipitridae Se
Galli ae Daa
Charadrii ——_—_
Columbidae See Se er
Cuculidae ae
Strigidae SSS
Picidae ee
Tyrannidae Sees bo aS
Hirundinidae —-
Corvidae Sarre eT
Paridae ia
Mimidae eaclied
Troglodytidae a:
Turdidae elt oe
Vireonidae Hi
Porulidae fare
Icteridae a ta
Richmondeninae ae are ay
Emberizinae oc
ies 2c ls lee ip a ee ee eee
Or? 4°68 lONI2 ie l6r3e.2Or2e
Breeding Index
Fic. 2.—Breeding indices for Kansan birds. Vertical hash-marks indi-
cate the value of breeding index for a given species; horizontal lines show
the range of values of breeding index for families and orders.
indices for many groups of Kansan birds. The values for species
of a given family have been linked by a horizontal line. The length
of this line is proportional to the degree to which the index values
for the species concerned resemble one another. Note that the
plottings for the Picidae, Corvidae, Turdidae, Tyrannidae, and
Icteridae each contain one point that is well-removed from a cluster
of points. This can be interpreted as a measure of the frequency
of adaptive plasticity versus adaptive conservatism; five of the 24
plottings show a plastic character, 19 a conservative. There are 26
plottings that show temporal consistency, all of which may be taken
as evidence of adaptive (or relictual) conservatism of the species
in question.
598 UNIVERSITY OF KANSAs PuBLs., Mus. Nat. Hist.
Conclusion
Such patterns of breeding chronology support the idea that
seasonal response to the necessities of breeding is conservative more
often than plastic. Most students of breeding schedules believe
that since these are highly adaptive, they must also be capable of
flexibility to meet variable environments within the range of the
species. Such thinking receives support when different geographic
localities are considered for one species (Johnston, 1954), or when
specific features of a special environment are considered (see Miller,
1960; Johnston, 1956).
Yet, if one, relatively restricted locality is considered, as in the
present study, evidence of a conservative characteristic in breeding
schedules can be detected. This conservatism may result from the
historic genetic “burden” of the species; that is to say, previous
adaptive peaks may in part be evident in the matrix of contemporary
adaptation. Adaptive relicts of morphological nature have been
many times documented, but characteristics associated with season-
ality and timing schedules have not.
In any event, genetic relationships are evident in the configuration
of breeding seasons of many species here treated. Thus, any con-
sideration of variation in breeding schedules must be sensitive to
the limits, whether broad or restricting, that the heritage of a
species sets on its present chronological adaptation.
Regulation of Breeding Schedules
Regulation of breeding schedules in birds always involves some
exogenous, environmental timing or triggering mechanism. Broad
limits to functional reproductive activity seem to be set by the
photoperiod-neuroendocrine system. This basic, predominately
extra-equatorial, regulator can be ignored by temperate-zone species
only if they possess chronological adaptation to special, aperiodic
environmental conditions, as does the Red Crossbill (Loxia curvi-
rostra; see McCabe and McCabe, 1933; H. B. Tordoff, ms.), for
which the chief consideration seems to be availability of conifer
seeds. Environmental phonomena otherwise known to trigger
breeding activity include rainfall (Davis, 1953; Williamson, 1956),
presence of suitable nesting material (Marshall and Disney, 1957;
Lehrman, 1958), temperature (Nice, 1987), and presence of a mate
(Lehrman, Brody, and Wortis, 1961). Such regulators, or environ-
mental oscillators, are the “phasing factors” of the physiologic clock
that dictate the temporal occurrence of primary reproductive ac-
tivity.
THE BREEDING Birps OF KANSAS 599
TABLE 9.—RELATIONSHIP BETWEEN ENVIRONMENTAL FACTORS AND TIMING OF
BREEDING IN Briaps OF KANSAS
Occurrence of Peak of Egg-laying
When Precipi- When Mean
tation is: Temperature (F.) is:
Light | Heavy | < 55° | < 70° | = 70° | > 70°
IRAPUOIS <6 sees <5 so x
O.W. Element....... x X
Residents: ...)......:<(- «). x
Grassland species... . x
Marshland species... .
N. Amer. Element....
Migrants® ascii e:
Woodland species... .
Aerial foragers.......
S. Amer. Element....
pa a a a
Am mM
mm
None of the regulators mentioned above has been specifically
investigated for any Kansan bird, but it is reasonable to suppose
that, in these temperate-zone species, the photoperiod is the most
important general phasing factor in seasonal breeding. Although
gonadal response and seasonal restriction of breeding are set by
the photoperiod, specific temporal relationships are dictated by more
immediate environmental variables.
Table 9, as already noted, shows the gross relationships between
certain groups of birds, certain arbitrary indicators of seasonal
temperature-humidity conditions bearing significantly on the grow-
ing season, and occurrence in time of peak of egg-laying by the
birds involved. Some species and groups of Kansan birds breed
chiefly under cool-dry environmental conditions, and some under
warm-wet environmental conditions. Within each of these cate-
gories some variation occurs. Thus, raptors and boreally-adapted
species (the Eurasian zoogeographic element) breed under cool
conditions prior to rains, and residents and grassland species breed
under slightly warmer conditions prior to rains; limnic species,
species derived from North American evolutionary stocks, and mi-
grants tend to breed in the cooler segment of the warm-wet period,
and woodland birds, aerial foragers, and species derived from South
American evolutionary stocks tend to breed in the warmer segment
of the warm-wet period.
So much, then, for relationships between birds and their environ-
ments at a descriptive level. It would be useful at this point to
600 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
examine how environmental variables relate to timing of breeding.
Certain independent lines of investigation indicate that birds have
a well-developed internal timing device; most convincing is the
work of Schmidt-Koenig (1960) and the others who have shown
that the endogenous clock of birds can be shifted in its periodicity
forward or backward in time. This and much other evidence (see
Brown, 1960) indicate that many fundamental periodic regulators
are extrinsic to the animal; it is thus permissible for present purposes
to consider any expression of variation in timing as dependent on
environmental oscillators. It is not hereby meant to ignore the fact
that differential responses to dominant environmental variables oc-
cur within a species, indicating endogenous control over timing of
breeding. The work by Miller (1960:518) with three populations
of the White-crowned Sparrow, revealing innately different re-
sponses to vernal photoperiodic increase, is especially important in
this regard. For the moment, however, we may consider exogenous
controls only.
Any exogenous control, or environmental variable, can be looked
on simply as a timing oscillator. Such variables show regular or ir-
regular periodic activity, and the independent actions as a whole
result in the more-or-less variable annual schedule of breeding for
any species at any one place. It would seem that some oscillators
are linked to one another, but there is a real question concerning the
over-all degree to which linkage is present. It is significant that fre-
quency distributions of breeding activity of various species and
groups of birds take on the shape of a skewed normal curve. The
more information is added to such distributions, the more nearly
they approach being wholly normal, with irregularities tending to
disappear. This kind of response itself is evidence that most of the
variables influencing the distribution are not mutually linked.
This conclusion is warranted if we examine what would happen
to frequency distributions if the variables or oscillators regulating
timing were linked. The frequency distribution of breeding activity
in birds is described by a nonlinear curve (a normal distribution is
nonlinear). Let us assume that each of the environmental variables
is a nonlinear oscillator, as is probable. A set of nonlinear oscillators
mutually entrained or coupled and operating with reference to a
given phenomenon would result in that phenomenon being described
by a frequency distribution much more stable than if it were regu-
lated by any one oscillator alone. However, the frequency distribu-
tion of a set of coupled nonlinear oscillators is non-normal (Wiener,
1958).
THE BREEDING Binps OF KANSAS 601
We do not obtain such distributions in describing breeding ac-
tivity, so we may say that the oscillators regulating such activity
are not coupled. Present distribution, habitat preference, residency
status, foraging adaptation, previous zoogeographic history, and
relicts of ancestral adaptation, all bear on the character of the
breeding schedule of any bird species. The emphasis above on
multiple regulation of breeding schedules conceivably reflects the
true picture, but any such emphasis is made at the expense of taking
one factor as basic, or reducing the many to one, in order to manu-
facture simplicity.
ACCOUNTS OF SPECIES
In each account below information is given concerning status,
habitat, geographic distribution, seasonal occurrence, schedule of
egg-laying, number of eggs laid, and sites of nests, as these pertain
to Kansas, unless otherwise stated. The ways in which some of
these points were elucidated are as follows.
1.—Breeding schedule. Frequency distributions of egg-laying in
time are calculated on the basis of dates of completed clutches, as
described earlier (p. 588). Any event in the series of actions of
nesting—nestbuilding, egg-laying incubation, brooding, feeding
young out of nests—can be manipulated by adding or subtracting
days to or from the date of record to yield the probable date of
completion of the clutch. The resulting data are grouped into class
intervals of ten days. Extreme dates here given for egg-laying
may be as much as nine days off in accuracy, but the error does
not often exceed five days. Extreme dates indicated here may be
taken as actual or predicted extremes. The raw data used are on
file at the Museum of Natural History and are available for use by
any qualified individual.
2.—Dates of occurrence. First and last annual occurrences in the
State for migrant species are indicated by both a range of dates
and a median date. Twenty to 30 dates of first observation in spring
are available for most of the common species, and 10 to 20 dates
of last observation in autumn are at hand for such species. The
median dates, earlier than and subsequent to which an equal
number of observations are available, are reliable indicators of the
dates on which a species is likely to be seen first in the State in an
average year.
3.—Clutch-size. Information on number of eggs is given for
each species according to the mode, followed by the mean, the
range, and the size of the sample.
4.—Distribution in Kansas. Information on distribution in the
602 Unrversitry OF Kansas Puszs., Mus. Nat. Hist.
breeding season within the borders of Kansas is given in accounts
below chiefly by reference to one or more counties of the State.
Location of counties can be made by referring to Figure 10.
Pied-billed Grebe: Podilymbus podiceps podiceps (Linnaeus).—This is a
common but local summer resident, in and on ponds, marshes, streams, ditches,
and lakes. The species can be seen in the State at any time, but usually ar-
rives in the period March 1 to April 18 (the median is March 21), and departs
southward in the period October 13 to November 18 (the median is October
24).
Breeding schedule——Nineteen records of breeding span the period May 1
to June 30; the modal date for egg-laying is May 15.
Number of eggs.—Clutch-size is 4 to 10 eggs.
Nests are floating masses of marsh vegetation (cattail, smartweed, duck-
weed, filamentous green algae, and the like), kept green on top by addition
of fresh material, in or at the edge of emergent marsh vegetation.
Double-crested Cormorant: Phalacrocorax auritus auritus (Lesson ).—This
is a transient, but has been found nesting on one occasion in Barton County
( Tordoff, 1956:311).
Breeding schedule—Eggs were laid in July and August in the one known
nesting effort.
Number of eggs.—Clutch-size is 2 to 4 eggs (Davie, 1898).
Great Blue Heron: Ardea herodias Linnaeus.—This common summer resi-
dent nests in tall trees along rivers, streams, and marshes. The sector of
greatest abundance is the Flint Hills. A. h. herodias Linnaeus occurs in ex-
treme northeastern Kansas, A. h. wardi Ridgway breeds in southeastern Kansas,
and A. h. treganzai Court breeds in western Kansas; specimens showing inter-
mediate morphology have been taken from the central part of the State.
Occurrence in time, exclusive of the few that overwinter in Kansas, is shown
in Table 10.
Breeding schedule——Seventy-seven records of breeding span the period
March 1 to April 30 (Fig. 3); the modal date of egg-laying is April 5.
Number of eggs.—Clutch-size is 4 eggs (4.4, 3-6; 36).
TABLE 10.—OcCURRENCE IN TIME OF SUMMER RESIDENT HERONS IN KANSAS
Arrival Departure
SPECIES
Range Median Range Median
Great Blue Heron..| Feb. 4-Apr. 8 | Mar. 20 | Oct. 10-Nov. 29 | Oct. 23
Green Heron...... Mar. 29-May 4 | Apr. 27 | Sept. 1-Oct. 30 | Sept. 9
Common Egret....} Apr. 8-May 12 | Apr. 2 | Sept. 4-Sept. 30 | Sept. 21
Black-crowned
Night Heron. ..} Mar. 27-May 18 | Apr. 25 | Sept. 10—-Nov. 11 | Sept. 25
Yellow-crowned
Night Heron: .,:|(Apr.. 15-MayalS-i Apr 20 tlic perce cc seat au| ae ceera
American Bittern..| Apr. 4-May 9] May 1 | Oct. 6—Dec. 12 | Oct. 16
Least Bittern. “| Apr. - 9-May 22 | Apri 8) i-Octy 240 1° ie sien
THE BREEDING Birps OF KANSAS 603
Nests are placed in crotches of sycamore, cottonwood, elm, hackberry, oak,
and walnut, from 30 to 60 feet high; the average height is about 40 feet.
Green Heron: Butorides virescens virescens (Linnaeus ).—This is a common
summer resident about streams, lakes, and marshes throughout the State.
Some characteristics of the temporal occurrence of this species are indicated
in Table 10.
Breeding schedule—Twenty-eight records of breeding span the period April
21 to June 20 (Fig. 3); the modal date of completion of clutches is May 5.
Number of eggs.—Clutch-size is 8 eggs (3.1, 3-5; 17).
Nests are placed about 10 feet high (two to 35 feet) in willow, cottonwood,
elm, and the like.
Little Blue Heron: Florida caerulea caerulea (Linnaeus).—This is chiefly
a postbreeding summer visitant, but there is one record of breeding in Finney
County (Tordoff, 1956:312).
Breeding schedule——There is no information on breeding schedule in Kansas
or in adjacent areas.
Number of eggs.—Clutch-size is 2 to 4 eggs (Davie, 1898).
Nests are placed in trees and bushes at various heights above the ground.
Common Egret: Casmerodius albus egretta (Gmelin).—This is a post-
breeding summer visitant, but has been found nesting once in Cowley County
(Johnston, 1960:10). Occurrence in time is listed in Table 10.
Breeding schedule.—There is no information on breeding schedule in Kansas.
Number of eggs.—Clutch-size is 2 to 4 eggs (Davie, 1898).
Nests are placed in trees, usually above 20 feet in height; the one instance
of nesting in the State was within a colony of Great Blue Herons.
Snowy Egret: Leucophoyx thula thula (Molina).—This postbreeding sum-
mer visitant has been found nesting once in Finney County (Tordoff,
1956:312).
Breeding schedule.—There is no information on breeding schedule in the
State.
Number of eggs.—Clutch-size is 2 to 5 eggs (Davie, 1898).
Nests in Kansas are placed among those of Great Blue Herons.
Black-crowned Night Heron: Nycticorax nycticorax hoactli (Gmelin).—
This is a locally common summer resident around marshes and riparian habitats.
Characteristics of the occurrence of the species in time are given in Table 10.
Breeding schedule——Eggs are laid in the period May 1 to August 10.
Number of eggs—Clutch-size is about 4 eggs.
Nests are placed at medium elevations in riparian trees, in Kansas chiefly
cottonwood, or in beds of emergent marsh vegetation.
Yellow-crowned Night Heron: Nyctanassa violacea violacea (Linnaeus ).—
This is a local summer resident in riparian habitats, chiefly in southeastern
Kansas. Specimens taken in the breeding season and records of nesting come
from Meade, Stafford, Doniphan, Douglas, Greenwood, Woodson, Labette,
and Cherokee counties. Characteristics of occurrence in time in Kansas are
shown in Table 10.
604 UNIVERSITY OF KANSAS PuBLS., Mus. Nat. Hist.
Breeding schedule —Eggs are laid in May and June.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are placed in riparian trees.
Least Bittern: Ixobrychus exilis exilis (Gmelin).—This is a local summer
resident in marshland. Characteristics of its occurrence in time are indicated
in Table 10.
Breeding schedule.—Eleven records of breeding span the period May 21
to July 20; the modal date of egg-laying seems to be in the first week of June.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are placed in dense emergent vegetation a few inches to a foot above
the surface of the water.
American Bittern: Botaurus lentiginosus (Rackett).—This is a local sum-
mer resident in marshes and heavy grassland. The species occurs temporally
according to characteristics as listed in Table 10.
Breeding schedule —Eggs are laid in May and probably in June.
Number of eggs.—Clutch-size is 3 or 4 eggs.
Nests are placed on the ground in heavy cover.
White-faced Ibis: Plegadis chihi (Vieillot).—This is a local summer resi-
dent in marshland; actual records of breeding come only from Barton County
(Nossaman, 1952:7; Zuvanich, 1963; M. Schwilling, personal communication,
July, 1962). The species has been recorded in the State from April 17 to
October 6.
Breeding schedule —Twenty-five breeding records are for June and early
July.
Number of eggs.—Clutch-size is about 4 eggs (38.9, 8-4; 24).
Nests are placed in emergent marsh vegetation near the surface of the
water, in Barton County in extensive cattail beds harboring also Black-crowned
Night Herons.
Mallard: Anas platyrhynchos platyrhynchos Linnaeus.—This is a local sum-
mer resident around marshes. The time of greatest abundance is October to
April, but most birds move north for breeding.
Breeding schedule.—Fifteen records of breeding span the period April 1 to
June 10; the modal date of egg-laying is in the first ten days of May.
Number of eggs.—Clutch-size varies widely; first clutches are of about 12
eggs. Brood sizes vary from 8 to 12 individuals in Kansas.
Nests are placed on the ground surface, in pasture grasses, marsh grasses,
cattail, sedge, and smartweed.
Pintail: Anas acuta Linnaeus.—This is a local summer resident in marsh-
land. The time of greatest abundance is from September to May, but most
birds move north for breeding.
Breeding schedule—Eleven records of breeding span the period April 21
to June 10; the peak of egg-laying seems to be in the period May 1 to 10.
Number of eggs.—Clutch-size is around 10 eggs. Brood sizes vary from 3
to 8 individuals in Kansas.
Nests are placed on the ground surface, in cover of marsh grass, cattail,
or sedge.
Tue BREEDING Birps OF KANSAS 605
Blue-winged Teal: Anas discors discors Linnaeus—This summer resident
is locally common around marshes and ponds. The species arrives in spring
in the period March 9 to April 5 (the median is March 23); birds are last
seen sometime between October 7 and November 26 (the median is October
20).
Breeding schedule-—Twenty-two records of breeding span the period May 1
to May 30; the peak of egg-laying is around May 15. It is doubtful that the
present data indicate the full extent of the egg-season in this duck.
Number of eggs.—Clutch-size is 8 to 12 eggs.
Nests are placed on the ground surface, in cover of grasses, cattail and
sedges.
Shoveler: Anas clypeata Linnaeus.—This is an irregular and local summer
resident, around marshes. Most individuals seen in the State are passage
migrants. Breeding records are from Barton and Finney counties.
Breeding schedule—Seasonal limits are unknown for the Shoveler in Kansas.
Number of eggs.—Clutch-size is about 8 eggs (Davie, 1898).
Nests are placed on the ground surface in cover of marsh vegetation.
Wood Duck: Aix sponsa (Linnaeus ).—This is an uncommon summer resi-
dent around wooded streams and ponds in eastern Kansas. Nesting records
and specimens taken in the breeding season come from east of stations in
Pottawatomie, Coffey, and Woodson counties. Most nesting records at present
come from the Marais des Cygnes Wildlife Refuge, Linn County. The species
is present in the State from March 5 to December 8.
Breeding schedule —Eleven records of breeding span the period March 21
to May 10; the peak of egg-laying is probably in mid-April. The present data
are inadequate for showing the full span of the breeding season.
Number of eggs.—Clutch-size is around 15 eggs, varying from 10 to 23 in
the sample at hand.
Nests are placed in crevices and hollows in trees near water, 10 to 70 feet
high.
Redhead: Aythya americana (Eyton).—This duck nested at Cheyenne
Bottoms, Barton County, 1962: 9 eggs found May 31 (M. Schwilling); also
reported to have nested at Cheyenne Bottoms about 1928 ( Tordoff, 1956:316).
Canvasback: Aythya valisineria (Wilson)—This duck nested at Cheyenne
Bottoms, Barton County, 1962: 14 eggs found June 20 (M. Schwilling).
Ruddy Duck: Oxyura jamaicensis rubida (Wilson).—This is a local summer
resident in marshland; numbers seem generally higher in western than in eastern
Kansas. The season of greatest abundance is March through November, but
numbers are conspicuously reduced in midsummer.
Breeding schedule.—Eggs are known to be laid in May and June.
Number of eggs.—Clutch-size is about 10 eggs (Davie, 1898).
Nests are placed near the edge of water, either in or on emergent marsh
vegetation; nests of other marshland birds, such as coots, are sometimes ap-
propriated (Davie, 1898).
Turkey Vulture: Cathartes aura teter Friedmann.—This summer resident
is common throughout Kansas. Occurrence in time is indicated in Table 11.
Breeding schedule—Fifteen records of breeding span the period April 21
606 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
to June 10; earlier records will doubtless be found, to judge from the frequency
distribution of the present sample. The peak of egg-laying is perhaps around
May 1.
Number of eggs.—Clutch-size is 2 eggs (1.8, 1-2; 12).
jFeb., Mch,Apr. ,May JuneJuly Aug. Sep,Oct.,
Ardea herodias
(MG
Butorides virescens
28
L
Buteo jamaicensis
36
Colinus virginianus
24
ASS,
a is Charadrius vociferus
29
Recurvirostra americana
26
WwW
re)
SCALE
% OF CLUTCHES
LRH PELL) FAULT aaa (P| Pease cial aera eR ete]
Feb. Mch. Apr. May June July Aug Sep. Oct.
Fic. 3.—Histograms representing breeding schedules of two
herons, the Red-tailed Hawk, Bobwhite, and two shore birds
in Kansas. See legend to Figure 1 for explanation of histo-
grams.
THE BREEDING Brirps OF KANSAS 607
Nests are placed in holes and crevices in trees and cliffs, on rocky ledges,
and the like.
Black Vulture: Coragyps atratus (Meyer).—This is possibly a summer
resident in the southeastern sector of Kansas. There is one nesting record,
for Labette County (Goss, 1891:245).
Breeding schedule.—There are no data for this species in Kansas.
Number of eggs—Clutch-size is 2 eggs (Davie, 1898).
Nests are placed in hollows (logs, stumps, etc.) on the ground surface.
Swallow-tailed Kite: Elanoides forficatus forficatus (Linnaeus ).—This kite
was formerly a summer resident in eastern Kansas; it no longer occurs as a
breeding species.
Breeding schedule-——In Kansas the season seemed to occur relatively late
in the year for a raptor; eggs were laid in May, so far as is known.
Number of eggs.—Clutch-size is about 2 eggs (Davie, 1898).
Nests are placed in tops of trees.
Mississippi Kite: Ictinia misisippiensis (Wilson).—This is a common sum-
mer resident in southern Kansas, west to Morton County. Specimens taken in
the breeding season and records of nesting come from south of stations in
Grant, Barton, Harvey, and Douglas counties; the present center of abundance
is in Meade, Clark, Comanche, Barber, and Harper counties.
Breeding schedule—Seven records of breeding span the period April 20 to
June 10; the peak of egg-laying seems to be in the first week of May.
Number of eggs.—Clutch-size is 2 eggs.
Nests are placed about 35 feet high (from 25 to 50 feet) in cottonwood,
willow, elm, black locust, and the like.
Sharp-shinned Hawk: Accipiter striatus velox (Wilson ).—This rare summer
resident apparently occurs only in the eastern part. The two nesting records
are from Cloud and Pottawatomie counties.
Breeding schedule—The information at hand suggests the birds lay in
April and May.
Number of eggs.—Clutch-size is about 4 eggs (Davie, 1898).
Nests are placed 20 or more feet high in coniferous or deciduous trees.
Cooper Hawk: Accipiter cooperii (Bonaparte).—This is an uncommon
resident. Specimens taken in the breeding season and actual records of nesting
come from east of stations in Cloud, Anderson, and Montgomery counties.
Breeding schedule.—Fourteen records of breeding span the period March 21
to May 30; the modal date of egg-laying is April 25.
Number of eggs.—Clutch-size is 4 eggs (3.8, 2-5; 5).
Nests are placed from 15 to 80 feet high, averaging 25 feet in elm, oak,
and other trees.
Red-tailed Hawk: Buteo jamaicensis borealis (Gmelin).—This is a common
resident east of the 100th meridian; to the west numbers are reduced, although
the species is by no means unusual in western Kansas. Red-tails probably
always were uncommon in western Kansas; Wolfe (1961) reports that they
were “very rare as a nesting species” in Decatur County shortly after the turn
of the 20th Century.
38—1476
608 UNIVERSITY OF KAnsAs Pus3s., Mus. Nat. Hist.
Breeding schedule.—Thirty-six records of breeding span the period February
21 to April 10 (Fig. 3); the modal date of egg-laying is March 5.
Number of eggs.—Clutch-size is 3 eggs (2.6, 2-3; 20).
Nests are placed about 40 feet high, ranging from 15 to 70 feet in cotton-
wood, honey locust, osage orange, sycamore, and walnut.
Red-shouldered Hawk: Buteo lineatus lineatus (Gmelin).—This is an un-
common summer resident in eastern Kansas, in riparian and bottomland timber.
Nesting records are available from Leavenworth, Woodson, and Linn counties,
and red-shoulders probably also nest in Doniphan County (Linsdale, 1928).
Breeding season.—Eggs are laid in March and April.
Number of eggs.—Clutch-size is about 3 eggs (Davie, 1898).
Nests are placed up to 70 feet high in elms and other streamside trees.
TABLE 11.—OcCURRENCE IN TIME OF THE SUMMER RESIDENT VULTURE AND
Hawks IN KANSAS
Arrival Departure
SPECIES
Range Median Range Median
Turkey Vulture....} Mar. 7—Mar. 30 | Mar. 15 | Sept. 24-Oct. 28 | Oct. 5
Red-shouldered
Hawke 42 2+. Beb, 10=Mar5i4 } Heb. (26% "Oct-=Dee.? °F 9S. t.8
Broad-winged
la wikis fuadiekee Apr. 4-Apr. 21 | Apr. 12 | Sept. 1-Oct. 20 }........
Swainson Hawk....| Mar. 24-Apr. 28 | Apr. 12 | Oct. 5-Nov. 2 |} Oct. 11
Broad-winged Hawk: Buteo platypterus platypterus (Vieillot).—This is an
uncommon summer resident in eastern Kansas, in swampy woodland. Speci-
mens taken in the breeding season and nesting records are from Shawnee,
Douglas, Leavenworth, and Johnson counties; there are several nesting records
from Missouri in the bottomlands just across the river from Wyandotte County
Kansas, Occurrence in time is listed in Table 11.
Breeding schedule——Four records of nesting span the period April 21 to
May 30, but it is likely that the egg-season is longer than this.
Number of eggs.—Clutch-size is about 3 eggs.
Nests are placed high in deciduous trees.
Swainson Hawk: Buteo swainsoni Bonaparte—This is a common summer
resident in prairie grassland with open groves and scattered trees. Records of
breeding are available from all parts of the State, but are least numerous from
the southeastern quarter. Occurrence in time is listed in Table 11.
Breeding schedule.—Sixteen records of breeding span the period April 11
to June 10; the modal date for completion of clutches is April 25.
Number of eggs.—Clutch-size is 2 eggs (2.4, 2-3; 5).
Nests are placed about 35 feet high, actually ranging from 12 to 75 feet,
in cottonwood, elm, willow, and honey locust. Occasionally nests are placed
on ledges in cliffs.
Tue BREEDING Brrps oF KANSAS 609
Ferruginous Hawk: Buteo regalis (Gray).—This is an uncommon resident
in western Kansas, in grassland with scattered trees. Records of nesting and
specimens taken in the breeding season come from Wallace, Hamilton, Gove,
Logan, and Finney counties.
Breeding schedule —Five records of breeding span the period March 11 to
April 30.
Number of eggs.—Clutch-size is about 3 eggs (3.3, 3-4; 4).
Nests are placed on the ground surface on small cliffs or promontories or
low (six to 10 feet) in small trees such as osage orange, cottonwood, and
mulberry.
Marsh Hawk: Circus cyaneus hudsonius (Linnaeus ).—This is a local resi-
dent in grassland throughout Kansas. Most records of breeding come from
east of the Flint Hills, but it is not certain that the few records from the west
actually reflect a low density of Marsh Hawks in that area.
Breeding schedule—Sixteen records of breeding span the period April 11
to May 20; the modal date for egg-laying is May 5.
Number of eggs—Clutch-size is 5 eggs (5.2, 3-7; 14).
Nests are placed on the ground surface in grassy cover.
Peregrine Falcon: Falco peregrinus anatum Bonaparte.—This falcon nested,
perhaps regularly but clearly in small numbers, in Kansas prior to the 20th
Century. The best documented breeding occurrence was at Neosho Falls,
Woodson County (Goss, 1891:283).
Breeding schedule—Eggs were recorded as being laid in February and
March.
Number of eggs.—Clutch-size is 3 or 4 eggs (Davie, 1898).
Nests are placed relatively high on cliffs and in trees; at Neosho Falls these
birds used open cavities 50 to 60 feet high in sycamores.
Sparrow Hawk: Falco sparverius sparverius Linnaeus.—This is a common
resident throughout Kansas, in parkland and woodland edge.
Breeding schedule——Thirteen records of egg-laying span the period March
21 to May 20; the modal date of laying is not evident in this sample but it
probably falls around April 10.
Number of eggs.—Clutch-size is 4 eggs (4.2, 3-5; 5).
Nests are placed in cavities about 16 feet high, actually 12 to 30 feet, in
cottonwood, ash, maple, Purple Martin “houses,” and human dwellings.
Greater Prairie Chicken: Tympanuchus cupido pinnatus (Brewster ).—This
is a locally common resident in eastern Kansas, in and about bluestem prairie
grassland, and is local in the northwest in undisturbed plains grassland.
Wolfe (1961) reports that the species was common in Decatur County shortly
after the turn of the Century, but that it became rare by 1914.
Breeding schedule.—Twenty-one records of breeding span the period May
1 to June 10 (Fig. 3); the modal date for laying is May 5. The sample indi-
cates an abrupt inception to laying of eggs, and this may be a reflection of
timing characteristic of behavior at leks, or booming grounds.
Number of eggs.—Clutch-size is 12 eggs (11.7, 9-15; 17).
Nests are placed on the surface of the ground in bluestem grassland or plains
bunchgrass, usually under cover of prairie grasses and forbs.
610 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Lesser Prairie Chicken: Tympanuchus pallidicinctus (Ridgway ).—This is
a local resident in sandy grassland in southwestern Kansas. Distribution is to
the west and south of Pawnee County.
Breeding schedule——There is no information on timing of the breeding sea-
son in Kansas.
Number of eggs.—Clutch-size is thought to be near that of the Greater
Prairie Chicken. Vic Housholder (MS) observed a hen with ten chicks ten
miles south of Dodge City, Ford County, on June 1, 1955.
Bobwhite: Colinus virginianus (Linnaeus).—This is a common resident in
the east, but is local in western Kansas; occurrence is in broken woodland and
other edge habitats. C. v. virginianus (Linnaeus) is found northeast of sta-
tions in Nemaha, Douglas, and Miami counties, and C. v. taylori Lincoln is
found in the remainder of the State.
Breeding schedule—Twenty-four records of breeding span the period May
1 to September 20 (Fig. 3); the modal date for first clutches is May 25. The
long period of egg-laying after May probably includes both renesting efforts
and true second nestings.
Number of eggs.—Clutch-size is about 13 eggs (12.8, 8-21; 22); in the
present sample 16 eggs was the most frequent number.
Nests are placed on the surface of the ground at bases of bunch grasses,
saplings, trees, or posts, under cover of prairie grasses, forbs, or small woody
plants.
Scaled Quail: Callipepla squamata pallida Brewster—This is a locally
common resident in southwestern Kansas, chiefly west of Clark County and
south of the Arkansas River; preferred habitat seems to be in open, sandy
prairie.
Breeding schedule-——Eggs are laid at least in May; the egg-season in Kansas
is unlikely to be so prolonged as that of the Bobwhite; among other factors
involved, the Scaled Quail in Kansas is at a northern extreme of its distribu-
tion, where suboptimal environmental conditions may occur relatively fre-
quently.
Number of eggs.—Clutch-size is around 10 to 12 eggs.
Nests are placed on the ground surface under woody or herbaceous cover.
Ring-necked Pheasant: Phasianus colchicus Linnaeus.—This introduced
resident is common in western Kansas, is local and uncommon in the east,
and is found in agricultural land with scattered woody vegetation.
Breeding schedule—Eggs are laid at least in May.
Number of eggs.—Clutch-size is 10 to 12 eggs.
Nests are placed on the surface of the ground in woody or herbaceous
cover.
Wild Turkey: Meleagris gallopavo Linnaeus.—Turkeys formerly occurred
as common residents in floodplain woodland in eastern Kansas, and _ their
distribution extended through the west in riparian woodland. Present popu-
lation in eastern and southern sectors are partly the result of introductions of
birds from Missouri by humans in the 1950s. Turkeys in southern Kansas are
also present owing to natural dispersal along the Arkansas and Medicine Lodge
rivers of birds native to and introduced into Oklahoma. No specimens of tur-
keys presently found in Kansas are available for examination but these birds
THE BREEDING Birps OF KANSAS 611
probably are referable to M. g. silvestris Vieillot, the trinomen applied to tur-
keys in Missouri and northeastern Oklahoma.
Turkeys from southern Texas recently have been liberated at several locali-
ties in southern Nebraska; turkeys seen in extreme northern Kansas are thus
probably of these stocks. The name M. g. intermedia Sennett is applicable to
these birds.
Breeding schedule.—No information is available on the egg-season in Kan-
sas; turkeys have nested in southern Kansas within recent years, however.
Number of eggs——Clutch-size is perhaps 12 eggs.
Nests are placed on the surface of the ground, usually well-concealed under
woody vegetation.
King Rail: Rallus elegans elegans Audubon.—This summer resident is
locally common in marshlands. Nesting records or adults taken in the breed-
ing season are from Cheyenne, Meade, Pratt, Stafford, Cloud, Riley, Douglas,
Anderson, and Allen counties. Dates of arrival in spring are recorded from
April 7 to April 28; the median date is April 18. Departure in autumn is pos-
sibly as early as September in the north, but four records are in the period Oc-
tober 12 to November 25. The species occasionally can be found in winter
(Douglas County, December 28, 1915).
Breeding schedule—Fourteen records of breeding span the period May 1
to July 20; the modal date for egg-laying is June 5.
Number of eggs—Clutch-size is about 10 eggs (9 to 12; 4 records).
Nests are placed on the surface of the ground, under grassy or woody cover.
Virginia Rail: Rallus limicola limicola Vieillot—This is an uncommon sum-
mer resident, presumably throughout the State. The one breeding record is
from Morton County (May 24, 1950; Graber and Graber, 1951). Dates of
spring arrival are from April 19 to May 18; dates of last observation in autumn
are within the period September 1 to October 30. A few birds overwinter in
the southern part of the State (Meade County, December and January).
Breeding season—Eggs are laid probably in May and June.
Number of eggs.—Six to 12 eggs are laid (Davie, 1898).
Nests are placed in emergent aquatic plants, near the surface of the water.
Sora: Porzana carolina (Linnaeus).—This is an uncommon summer resi-
dent in marshland. Nesting records or specimens taken in the breeding season
come from Finney, Barton, Jefferson, Douglas, and Miami counties. First
dates of observation in spring are from April 11 to May 9 (the median is May
1); dates when last observed in autumn are from September 30 to November
9 (the median is October 18).
Breeding schedule—The one dated record comes from August.
Number of eggs—Clutch-size is around 10 eggs (Davie, 1898).
Nests are on the ground in grassy or herbaceous cover.
Black Rail: Laterallus jamaicensis jamaicensis (Gmelin).—This is an un-
common summer resident in Kansas. Records of breeding and specimens taken
in the breeding season come from Finney, Meade, Riley, and Franklin coun-
ties. Seasonal occurrence is within the period March 18 to September 26.
Breeding schedule—Eggs are laid at least in June.
Number of eggs.—Clutch-size is about 8 eggs (6-10; 4).
612 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
Nests are on the ground under cover of marsh plants.
Common Gallinule: Gallinula chloropus cachinnans Bangs.—This is a local
summer resident in marshlands. Nesting records and specimens taken in the
breeding season come from Barton, Stafford, Shawnee, Douglas, and Coffey
counties. Occurrence in the State is from April through September.
Breeding schedule—Eggs are laid in May and June.
Number of eggs——Clutch-size is about 10 eggs.
Nests are in marsh grasses and other emergent vegetation, not necessarily
over water.
American Coot: Fulica americana americana Gmelin.—This is an uncom-
mon, local summer resident in wetlands in Kansas. Coots are at greatest
abundance in autumnal and spring migratory movements, but are present all
year. Nesting has been recorded from Barton, Stafford, Doniphan, and
Douglas counties.
Breeding schedule.—Thirty-eight records of breeding span the period May
11 to June 30; the mode to laying is May 25. Earlier breeding probably occurs
in the State.
Number of eggs——Clutch-size is 8 eggs (7.7, 5-12; 28).
Nests are made of marsh vegetation (arrowhead, cattail) and float on water.
Snowy Plover: Charadrius alexandrinus tenuirostris (Lawrence).—This
summer resident is fairly common on the saline flats of central and south-
central Kansas. Breeding records are from Barton, Stafford, Meade, Clark,
and Comanche counties.
Breeding schedule.—Fifteen records show that eggs are laid in the period
May 25 to June 20; the peak of laying seems to be around June 10.
Number of eggs.—Clutch-size is 3 eggs.
Eggs are deposited on bare sand.
Killdeer: Charadrius vociferus vociferus Linnaeus——This summer resident
is common throughout the State, in open country frequently near wetlands.
A few individuals overwinter in Kansas, especially in the southern counties.
Breeding schedule-—The 29 records of breeding span the period March 21
to June 30; the modal date of laying is May 20. The distribution of com-
pleted clutches (Fig. 3) suggests that Killdeers are here double-brooded.
Number of eggs.—Clutch-size is 4 eggs.
Eggs are laid on the surface of the ground, frequently on gravel, field stubble,
plowed earth, and pasture.
Mountain Plover: Eupoda montana (Townsend).—This is an uncommon
and local summer resident in western short-grass prairie. Breeding records
come from Greeley and Decatur counties.
Breeding schedule-—Wolfe (1961) wrote that the species in Decatur County
laid eggs in the “last of May” in the early 1900s. The only other dated
breeding record is of downy young (KU 5512, 5513) taken on June 21.
Number of eggs.—Clutch-size is usually 3 eggs.
Eggs are laid in slight depressions in the ground, “lined with a few grass
stems,” according to Wolfe (1961).
American Woodcock: Philohela minor (Gmelin).—This is a rare summer
THE BREEDING Brirps oF KANSAS 613
resident in wet woodlands in eastern Kansas. Arrival in the northeast is from
mid-March through April, with departures southward occurring from Septem-
ber to December; the last date on which the species has been seen in any
year is December 5. There are nesting records only from Woodson County;
probably the species nests in Douglas County (Fitch, 1958:194).
Breeding schedule.—Eggs are laid in April.
Number of eggs.—Clutch-size is usually 4 eggs.
Nests are depressions in the dry ground within swampy places, usually
under heavy plant cover.
Long-billed Curlew: Numenius americanus americanus Bechstein.—This is
an uncommon summer resident in western Kansas, in prairie grassland. Breed-
ing records are from Stanton and Morton counties.
Breeding schedule.—Eggs are laid at least in May and June.
Number of eggs.—Clutch-size is 4 eggs.
Eggs are laid in slight depressions in the ground in grassy cover.
Upland Plover: Bartramia longicauda (Bechstein).—This is a locally com-
mon summer resident, most conspicuously in the Flint Hills, in grassland.
Breeding records are from Trego, Hamilton, Finney, Morton, Meade, Marion,
Chase, Kearny, Butler, Cowley, Douglas, Johnson, Wabaunsee, Franklin,
Anderson, and Coffey counties. Dates of first arrival in spring span the period
April 2 to May 5 (the median is April 19), and dates last seen in autumn are
from September 3 to October 6 (the median is September 13).
Breeding schedule —Sixteen records of breeding span the period April 21
to June 10; the modal date for egg-laying is May 5.
Number of eggs.—Usually 4 eggs are laid.
Eggs are placed on vegetation on the ground surface, in pasture, field
stubble, or gravel, frequently under heavy plant cover.
Spotted Sandpiper: Actitis macularia (Linnaeus).—This summer resident
is locally common on wet ground and along streams. Dates of arrival in spring
are from March 29 to April 30 (the median is April 24), and dates of last
observation in autumn span the period September 2 to October 10 (the median
is September 18).
Breeding schedule—Egg records are all from the northeastern sector, and
all are for May.
Number of eggs.—Usually 4 eggs are laid.
Nests are of plant fibers in depressions in dry ground on gravel banks,
pond or stream borders, or in pastureland.
American Avocet: Recurvirostra americana Gmelin.—This is a local summer
resident in marshes in central and western Kansas. There are breeding records
from Finney, Barton, and Stafford counties. Extreme dates within which
avocets have been recorded are April 2 to November 21.
Breeding schedule-—Forty-one records of breeding span the period May 11
to June 20 (26 records shown in Fig. 3); the modal date for laying is June 5.
Number of eggs.—Usually 4 eggs are laid.
Nests are placed on the surface of the ground, near water.
Wilson Phalarope: Steganopus tricolor Vieillot—This is a local summer
resident in marshes in central and western Kansas, but breeding records are
614 UNIVERSITY OF KANnsAs Pusts., Mus. Nat. Hist.
available only from Barton County. The earliest date of occurrence is April 7
and the latest is October 14.
Breeding schedule—Ten records indicate eggs are laid in May and June.
Number of eggs.—Three or 4 eggs are laid.
Nests are of plant stems in slight depressions in the ground.
Forster Tern: Sterna forsteri Nuttall—This is a local summer resident in
central Kansas, in marshes. There are breeding records only from Cheyenne
Bottoms, Barton County (Zuvanich, 1963:1). First dates of arrival in spring
span the period April 9 to 29 (the median is April 22), and apparent departure
south in autumn occurs from August 1 to November 1 (the median is Sep-
tember 3).
Breeding schedule—Twenty-three records of nesting are from late May to
mid-June; all records are for the year 1962.
Number of eggs.—Usually 4 eggs are laid.
Nests are frequently floating platforms of vegetation (algae, cattail, and the
like) in shallow water; old nests of Pied-billed Grebes are sometimes used as
bases, and occasionally the birds nest on the ground.
Least Tern: Sterna albifrons athalassos Burleigh and Lowery.—This tern is
a local summer resident in marshes and along streams in central and western
Kansas. There are breeding records from Hamilton, Meade, and Stafford
counties. First dates of arrival in spring are from May 14 to 80 (the median
is May 28), and last dates of occurrence in autumn are from August 9 to
September 7 (the median is August 25).
Breeding schedule—Twenty-one records of egg-laying are from May 21
to June 80 (Fig. 4); the modal date for laying is June 5.
Number of eggs—Two, 8 or 4 eggs are laid.
Eggs are laid on the bare ground, usually a sandy surface, near water.
Black Tern: Chlidonias niger surinamensis (Gmelin).—This is a local
summer resident in marshlands in central Kansas. There are breeding records
only from Barton County for 1961 and 1962; possibly the species breeds in
Douglas County. First dates of arrival in spring are from May 8 to 29 (the
median is May 14), and last dates of occurrence in autumn are from Sep-
tember 2 to 80 (the median is September 11).
Breeding schedule-—Twenty-four sets of eggs (Parmelee, 1961:25; M.
Schwilling ) were complete between June 11 and July 12.
Number of eggs—Clutch-size is 8 eggs.
Nests are of dead plant matter placed on floating parts of emergent green
plants in shallow water.
Rock Dove: Columba livia Gmelin.—This species was introduced into
North America by man from European stocks of semi-domesticated ancestry.
“Pigeons” now are feral around towns and farms, and cliffsides in the west,
and are locally common permanent residents throughout the State.
Breeding schedule—Eggs are laid in every month of the year. The main
season of breeding is spring, and this is depicted in Figure 4; the 26 records
of breeding by feral birds are from January 11 to June 10, and the modal
date of laying is probably April 5.
Number of eggs.—Pigeons usually lay 2 eggs.
THE BREEDING Birps OF KANSAS 615
Nests are of sticks and other plant matter placed on ledges and recesses
of buildings, bridges, and cliffs, 10 to 60 feet high.
Mourning Dove: Zenaidura macroura marginella (Woodhouse).—This is
a common summer resident throughout the State, in open country and wood-
land edge. The species is also present in winter in much reduced numbers,
and many are transient in periods of migration. The time of greatest abun-
dance is from March to November. Doves of extreme eastern Kansas have
by some workers been referred to the subspecies Z. m. carolinensis (Linnaeus);
specimens at the Museum of Natural History indicate that these doves are
Jan. Feb. Mch. Yan. Feb. Mch, Apr. May June July Aug. sep. May June July Aug. Sep.
Sterna albifrons
Columba livia
iki :
L) 8
va _ aida macroura
983
Zenaida macroura
43
Flees oo
Cocc a americanus
Bubo virginianus
Sit
30 @
eis
zou 2 Ss t /
aS sa Oo cunicularia
Ke )rs 2
oO
One
Jan. Feb. Mch. Apr. May June July Aug. Sep.
Fic. 4.—Histograms representing breeding schedules of the Least
Tern, two doves, the Yellow-billed Cuckoo, and two owls in
Kansas. See legend to Figure 1 for explanation of histograms.
616 UNIVERSITY OF KAnsas Pusts., Mus. Nat. Hist.
best regarded as members of populations of intermediate subspecific, or
morphologic, affinities, and that they are satisfactorily included within Z. m.
marginella.
Breeding schedule—-Numerous (983) records of egg-laying from north-
central Kansas are from April 1 to September 10; the modal date for laying
is May 15. Forty-three records of breeding from northeastern Kansas span
the period March 21 to August 10; the modal date of laying is May 15.
These samples are depicted in Figure 4.
Both sets of data are shown here to illustrate some of the differences
between large and small samples of heterogeneous data. The small sample
tends to be incomplete both early and late in the season, and the mode tends
to be conspicuous. Yet, the modes for the two samples coincide. Also, the
data from the north-central sector indicate that egg-laying in March would
be found less than once in 983 records, but the small sample from the north-
east includes one record for March. Such an instance doubtless reflects, at
least in part, the fact that the two geographic sectors have different environ-
mental conditions, but it is likely that the instance also partly reflects the
unpredictable nature of sampling.
Number of eggs—Doves lay two eggs. About one per cent of all nests
have 8 eggs, but it is not known for any of these whether one or two females
were responsible.
Nests are placed in a wide variety of plants, or on the ground. The
commonest plants are those used most frequently; in north-central Kansas
one-third of all nests are placed in osage orange trees, but in the northeast
elms are most frequently used. Nestsites are from zero to 15 feet high.
Yellow-billed Cuckoo: Coccyzus americanus americanus (Linnaeus ).—This
is a common summer resident in riparian and second-growth habitats throughout
the State. Twenty-three dates of first arrival in spring fall between April 29
and May 22 (the median is May 12), and nine dates of last observation in
autumn run from September 13 to October 12 (the median is September 23).
Breeding schedule.—Sixty-nine records of egg-laying span the period May
11 to September 10 (Fig. 4); the modal date of laying is June 5.
Number of eggs.—Clutch-size is 3 eggs (3.1, 2-5; 54).
Nests are placed about six feet high (from four to 20 feet) in sumac, rose,
pawpaw, mulberry, elm, cottonwood, willow, redbud, oak, osage orange, walnut,
boxelder, usually on horizontal surfaces, and in heavy cover.
Black-billed Cuckoo: Coccyzus erythropthalmus (Wilson ).—This is an un-
common summer resident, occurring in heavy riparian shrubbery and second-
growth. Breeding records are chiefly from eastern Kansas, but specimens
have been taken in the breeding season in all parts of the State. Eleven
dates of first arrival in spring are from May 7 to May 30 (the median is May
19), and four dates of last observed occurrence in autumn are between Sep-
tember 4 and October 7 (the average is September 18).
Breeding schedule.—Seventeen records of egg-laying are between May 21
and August 10; the mode is at June 5.
Number of eggs.—Clutch-size is 2 to 3 eggs (2.5, 2-8; 13).
Nests are placed about four feet high in heavy cover in plum, elm, locust,
and the like.
THE BREEDING Birps OF KANSAS 617
Roadrunner: Geococcyx californianus (Lesson ).—This is a local resident in
southern Kansas in xeric scrub or open edge habitats. Breeding records are
from Cowley and Sumner counties.
Breeding schedule—Eggs are laid at least from early April to mid-July.
Number of eggs.—Clutch-size is about 5 eggs (4.5, 3-6; 4).
Nests are placed on the ground under plant cover, or occasionally low in
bushes.
Barn Owl: Tyto alba pratincola Bonaparte.—This resident has a low density
throughout Kansas in open woodland and near agricultural enterprises of man.
Breeding schedule—The few records available indicate egg-laying occurs
at least from April to July; elsewhere the species is known to have a more
protracted breeding schedule.
Number of eggs.—Clutch-size is about 5 eggs (4.7, 2-6; 4).
Nests are informal aggregations of sticks and litter placed in recesses in
stumps, hollow trees, rocky and earthen banks, and dwellings and outbuildings
of man.
Screech Owl: Otus asio (Linnaeus ).—This is a common resident in wood-
land habitats throughout Kansas. O. a. aikeni (Brewster) occurs west of
Rawlins, Gove, and Comanche counties, and O. a. naevius (Gmelin) occurs
in the remainder of the State except for the eastern south-central sector, oc-
cupied by O. a. hasbroucki Ridgway.
Breeding schedule——Fifteen records of egg-laying span the period March
20 to May 10; there is a strong mode at April 5.
Number of eggs—Clutch-size is 4 eggs (4.0, 3-6; 12).
Nests are placed in holes and recesses in trees, three to 20 feet high.
Great Horned Owl: Bubo virginianus (Gmelin).—This is a common resi-
dent throughout Kansas, especially near woodlands and cliffsides. B. v. vir-
ginianus (Gmelin) occurs east of a line through Rawlins and Meade counties
and B. v. occidentalis Stone occurs to the west.
Breeding schedule—Fifty-seven records of egg-laying span the period Jan-
uary 11 to March 20 (Fig. 4); the modal date for laying is near February 10.
Number of eggs.—Clutch-size is 2 eggs (2.4, 2-3; 22).
Nests are placed about 30 feet high in cottonwood, elm, osage orange, hack-
berry, juniper, locust, cliffsides, and buildings of man. Old nests of hawks,
crows, and herons are frequently appropriated.
Burrowing Owl: Speotyto cunicularia hypugaea (Bonaparte).—This is an
uncommon summer resident in western Kansas in grassland and open scrub
habitats. Stations of breeding all come from west of a line running through
Cloud and Barber counties. Arrival in spring is between March 22 and April
17 (the median for 7 records is April 9), and dates last seen in autumn span
the period September 8 to November 14 (the median for 9 records is Sep-
tember 26).
Breeding schedule—-Twenty-one records of egg-laying run from April 11
to July 10 (Fig. 4); the mode of laying is May 15.
Number of eggs —Clutch-size is 7 or 8 eggs.
Nests are informal aggregations of plant and animal fibers in chambers of
earthen burrows usually made by badgers or prairie dogs.
618 UnIveErsITy OF Kansas Pusts., Mus. Nat. Hist.
Barred Owl: Strix varia varia Barton.—This is a loca] resident in eastern
Kansas, in heavy woodland. The species is said by implication (A. O. U.
Check-list, 1957) to occur in western Kansas, but no good breeding records
are available, all such records coming from and east of Morris County.
Specimens from southeastern Kansas show morphologic intergradation with
characters of S. v. georgica Latham.
Breeding schedule-—Three records of egg-laying are for the first half of
March.
Number of eggs——Clutch-size in our sample is 2 eggs.
Nests are situated in cavities in trees or in old hawk or crow nests.
Long-eared Owl: Asio otus wilsonianus (Lesson).—This ow] is a local resi-
dent or summer resident in woodland with heavy cover throughout the State.
Breeding records are available from Trego, Meade, Cloud, and Douglas
counties.
Breeding schedule ——Four records of egg-laying are for the period March 11
to April 10.
Number of eggs.—Clutch-size is 5 or 6 eggs.
Nests are placed in hollows of trees, stumps, cliffsides, on the ground sur-
face, or in old hawk, crow, or magpie nests (Davie, 1898).
Short-eared Owl: Asio flammeus flammeus (Pontoppidan).—This is a local
resident or summer resident in open, marshy, and edge habitats; records of
nesting come from Republic, Marshall, Woodson, and Bourbon counties.
Breeding schedule-—Eggs are laid at least in April.
Number of eggs.—Clutch-size is about 6 eggs (Davie, 1898).
Nests are simple structures of sticks and grasses, placed on the ground in
grasses, frequently near cover of downed timber or bushes.
Saw-whet Owl: Aegolius acadicus acadicus (Gmelin).—This is a rare and
local resident, in woodland. There is one breeding record (summer, 1951,
Wyandotte County; Tordoff, 1956:331).
Chuck-will’s-widow: Caprimulgus carolinensis Gmelin.—This is a locally
common summer resident in woodland habitats in eastern Kansas. Stations
of occurrence of actual breeding fall south of Wyandotte County and east of
Shawnee, Greenwood, Stafford, and Sedgwick counties.
Breeding schedule——Five records of breeding come between April 21 and
May 31, with a peak perhaps in the first third of May.
Number of eggs.—Clutch-size is 2 eggs.
Eggs are laid on heavy leaf-litter, usually under shrubby cover.
Whip-poor-will: Caprimulgus vociferus vociferus Wilson.—This is a local
summer resident in woodland in eastern Kansas. Breeding records are avail-
able only from Doniphan, Leavenworth, and Douglas counties; there are sight
records in summer from Shawnee County.
Breeding schedule-——Two records of breeding cover the period May 21 to
June 20.
Number of eggs.—Clutch-size is 2 eggs.
Eggs are laid on heavy leaf-litter in shrubby cover.
Poor-will: Phalaenoptilus nuttallii nuttallii (Audubon ).—This is a common
summer resident in western Kansas, in xeric, scrubby woodland. Breeding
Tue BREEDING Birps OF KANSAS 619
TABLE 12.—OccURRENCE IN TIME OF SUMMER RESIDENT CAPRIMULGIDS AND
APoDIDS IN KANSAS
Arrival Departure
SPECIES
Range Median Range Median
Chuck-will’s-
widow.........| Apr. 20-May 1 | Apr. 28 | Oct.—Dec. Oct.?
Whip-poor-will..... Apr. 6—Apr. 25 | Apr. 17 | Sept. 10-Oct. 11 | Sept. 21
Poor-willls 5. < Sipsu: Ayr, U20 glee ory chee Sept. 20°") Gisele
Common Night-
AWK tae eee: Apr. 29-May 23 | May 15 | Sept. 13-Oct. 18 | Sept. 23
Chimney Swift..... Apr. 2-Apr. 30 | Apr. 22 | Sept. 18-Oct. 30 | Oct. 4
Ruby-throated
Hummingbird. .| Apr. 2-May 19 | May 6 | Sept. 3-Oct. 15 | Sept. 10
records are chiefly from west of Riley County, but there is one from Franklin
County; specimens taken in the breeding season are available from Doniphan,
Douglas, Anderson, Woodson, and Greenwood counties.
Breeding schedule.—Six records of egg-laying are from the period May 1
to June 20.
Number of eggs.—Clutch-size is 2 eggs.
Eggs are laid on the ground, with or without plant cover.
Common Nighthawk: Chordeiles minor (Forster).—This is a common sum-
mer resident throughout Kansas. Temporal occurrence is indicated in Table
1l. Three subspecies reach their distributional limits in the State, C. m.
minor (Forster) in northeastern Kansas, C. m. chapmani Coues in southeastern
Kansas, and C. m. howelli Oberholser west of the Flint Hills.
Breeding schedule-—Twenty-two records of breeding span the period May
11 to June 30; the modal date for egg-laying is June 10 (Fig. 5).
Number of eggs.—Clutch-size is 2 eggs.
Eggs are laid on the ground in rocky or gravelly areas, on unpaved roads,
or on flat, gravelled tops of buildings of man.
Chimney Swift: Chaetura pelagica (Linnaeus ).—This is a common summer
resident in eastern Kansas, around towns. Temporal occurrence in the State
is indicated in Table 12.
Breeding schedule.—Thirty-six records of breeding span the period May 11
to June 30; the modal date for egg-laying is May 25 (Fig. 5).
Number of eggs.—Clutch-size is about 4 eggs.
Nests are secured by means of a salivary cement to vertical surfaces, usu-
ally near the inside tops of chimneys in dwellings of man, but occasionally in
abandoned buildings and hollow trees.
Ruby-throated Hummingbird: Archilochus colubris (Linnaeus ).—This is an
uncommon summer resident in eastern Kansas, and is rare in the west, in towns
and along riparian vegetation. Temporal occurrence in the State is listed in
Table 12.
Breeding schedule——Eight records of breeding fall within the period May
21 to July 10; there seems to be a peak to laying in the last third of June. —
620 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Number of eggs.—Clutch-size is 2 eggs.
Most nests are on outer branches of shrubs and trees, in forks or on pendant
branches, 10 to 20 feet high.
Belted Kingfisher: Megaceryle alcyon alcyon (Linnaeus).—This summer
resident is common throughout the State in streamside and lakeside habitats.
Timing of arrival and departure of the breeding birds is not well-documented
owing to the fact that the species is also transient and a winter resident in the
State.
Breeding schedule-—Eggs are laid at least from April 21 to May 20.
Number of eggs.—Clutch-size is near 6 eggs.
Eggs are laid on the floor of the chamber at the inner end of a horizontal
tunnel excavated in an earthen bank. The tunnel is two to six feet long and
many tunnels are strewn with bones and other dietary refuse.
Yellow-shafted Flicker: Colaptes auratus (Linnaeus).—This is a common
resident and summer resident in eastern Kansas, meeting, hybridizing with,
and partly replaced by Colaptes cafer westward, in open woodlands. C. a.
auratus (Linnaeus) occurs in southeastern Kansas, and C. a. luteus Bangs oc-
curs in the remainder, intergrading west of the Flint Hills with C. cafer.
Breeding season.—Forty-eight records of breeding span the period April
11 to June 10; the modal date for egg-laying is May 10 (Fig. 5). This sample
is drawn from central and eastern Kansas, but includes records of breeding by
some birds identified in the field as C. cafer.
Number of eggs.—Clutch-size is about 6 eggs.
Nests are piles of wood chips in cavities excavated in stumps and dead
limbs of trees such as willow, cottonwood, mulberry, and catalpa, ordinarily
about six feet above the ground.
Red-shafted Flicker: Colaptes cafer collaris Vigors—This woodpecker is
a common summer resident in western Kansas, meeting, hybridizing with, and
largely replaced by C. auratus in central and eastern sectors. The vast majority
of specimens taken in Kansas show evidence of intergradation with C. auratus.
Breeding schedule-—The few records of flickers identified in the field as
C. cafer have been combined with those of C. auratus (Fig. 5).
Number of eggs.—Clutch-size is perhaps 6 eggs.
Nests are like those of C. auratus.
Pileated Woodpecker: Dryocopus pileatus (Linnaeus).—This is a rare and
local resident in the east, in heavy timber. The species has been seen, chiefly
in winter, in all sectors of eastern Kansas in recent years, but actual records
of breeding come only from Linn and Cherokee counties. D. p. abieticola
(Bangs) occurs in the northeast, and D. p. pileatus (Linnaeus) in the south-
east.
Breeding schedule.—Eggs are laid at least in April.
Number of eggs.—Clutch-size is 3 or 4 eggs.
Nests are of wood chips in cavities excavated 45 to 60 feet high in main
trunks of cottonwood, sycamore, and pin oak.
Red-bellied Woodpecker: Centurus carolinus zebra (Boddaert ).—In wood-
land habitats this is a common resident in eastern Kansas, local in the west.
Breeding schedule——tThirty-seven records of breeding span the period
March 1 to June 30 (Fig. 5); the modal date of egg-laying is around April 25.
THE BREEDING Birps OF KANSAS 621
Mch. (Sa SAT May dune Neg Aug. Apr. May June July
roe ares al en Ar
eee Ss minor Tyrannus tyrannus
63
Chaetura pelagica Tyrannus verticalis
Muscivora forficata
28
a eae auratus
aa Centurus caro/linus
Contopus virens
‘- aac as erylhrocephalu. 9
is
Dendrocopos villosus
28
oer
=
Myrarchus crinitus
22
p
=
Empidonax traiflit
Ze
eh
f
40
ws ne ieee pubesens
30 ©
45
ted Sayornis phoebe
oO
Peels 136
0
[Swe re Colt cern ee eel
Mch. Apr. MayJune July Wen ee a junouuly Ade. Apr. May June July
Fic. 5.—Histograms representing breeding schedules of the Common Night-
hawk, Chimney Swift, woodpeckers, and flycatchers in Kansas. See legend to
Figure 1 for explanation of histograms.
622 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
Number of eggs.—Clutch-size is about 5 eggs.
Nests are of wood chips in cavities excavated in elm, cottonwood, box elder,
ash, hickory, or willow, about 25 feet high (nine to 60 feet).
Red-headed Woodpecker: Melanerpes erythrocephalus (Linnaeus ).—This
is a common summer resident and uncommon permanent resident in open
woodland; in winter it is noted especially around groves of oaks. M. e.
erythrocephalus (Linnaeus) occurs in eastern Kansas and M. e. caurinus Brod-
korb occurs in central and western Kansas.
Breeding schedule.—Fifty-eight records of breeding span the period May 1
to August 10 (Fig. 5); the modal date of egg-laying is June 5.
Number of eggs.—Clutch-size is 3 or 4 eggs.
Nests are of wood chips in cavities excavated about 25 feet high in willow,
cottonwood, and elm.
Hairy Woodpecker: Dendrocopos villosus villosus (Linnaeus ).—This resi-
dent is common in woodlands throughout the State.
Breeding schedule-——Twenty-eight records of breeding span the period
March 21 to May 80 (Fig. 5); the modal date of egg-laying is May 5.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are of wood chips in cavities excavated about 13 feet high in elm,
honey locust, and ash.
Downy Woodpecker: Dendrocopos pubescens (Linnaeus).—This resident is
common in woodland throughout the State. D. p. pubescens (Linnaeus)
occurs in southeastern Kansas, and D. p. medianus (Swainson) in the re-
mainder.
Breeding schedule.—Forty-one records of breeding span the period April
11 to June 10 (Fig. 5); the modal date of egg-laying is May 5.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are of wood chips in cavities excavated about 20 feet high in willow,
honey locust, ash, apple, and pear.
Eastern Kingbird: Tyrannus tyrannus (Linnaeus).—This summer resident
is common throughout the east; it is local in the west but there maintains
conspicuous numbers in favorable places, such as riparian woodland; preferred
habitat in eastern sectors is typically in woodland edge. Temporal occurrence
is indicated in Table 18.
Breeding season.—Sixty-three dates of egg-laying span the period May 11
to July 20 (Fig. 5); the modal date for completion of clutches is June 15.
Nearly 70 per cent of all eggs are laid in June.
Number of eggs.—Clutch-size is 8 eggs (3.8, 2-8; 10). Clutches are
probably larger than the average in May and smaller in June and July.
Nests are placed in crotches, terminal forks, and some on tops of limbs,
about 16 feet high, in elm, sycamore, honey locust, willow, oak, apple, and
red cedar.
Western Kingbird: Tyrannus verticalis Say—This summer resident is com-
mon in the west, but is local and less abundant in the east. Preferred habitat
is in woodland edge, open country with scattered trees, and in towns. Tem-
poral occurrence is indicated in Table 13.
THE BREEDING Birps OF KANSAS 623
Breeding schedule.—The 124 dates of egg-laying span the period May 11 to
July 31 (Fig. 5); the modal date for egg-laying is June 15. More than 70
per cent of all clutches are laid in June.
Number of eggs.—Clutch-size is 4 eggs (3.6, 3-4; 8).
Nests are placed in crotches, lateral forks, or on horizontal limbs, about 26
feet high, in cottonwood, elm, osage orange, hackberry, honey locust, mulberry,
oak, and on power poles.
Scissor-tailed Flycatcher: Muscivora forficata (Gmelin).—This summer
resident is common in central and southern Kansas; it is rare to absent in the
northwestern sector, and is local in the northeast. Preferred habitat is in open
country with scattered trees. Temporal occurrence is indicated in Table 13.
Breeding schedule.—Twenty-eight records of breeding occur from May 21
to July 10 (Fig. 5); the modal date of egg-laying is June 25. The present
sample of records is small, and there is otherwise no evidence suggesting that
the breeding schedule of this species differs from those of the other two king-
birds in Kansas.
Number of eggs—Clutch-size is 8 eggs (3.2, 2-5; 17). Mean clutch-size
for the first peak of laying shown in Figure 5 is 4.0 eggs; that for the second
peak is 2.7 eggs.
Nests are placed in forks or on horizontal limbs of osage orange, red haw,
elm, and on crosspieces of power poles, about 15 feet high (ranging from five
to 85 feet).
Great Crested Flycatcher: Myiarchus crinitus boreus Bangs.—This summer
resident is common in eastern Kansas, but is less numerous in the west. Pre-
ferred habitat is in woodland and woodland edge. Temporal occurrence is
indicated in Table 13.
Breeding schedule-—The twenty-two records of egg-laying are in the period
May 11 to July 10 (Fig. 5); the modal date for egg-laying is June 5. The
shape of the histogram (Fig. 5) indicates that some breeding for which records
are lacking occurs earlier in May.
TABLE 13.—OccURRENCE IN TIME OF SUMMER RESIDENT FLYCATCHERS IN
KANSAS
Arrival Departure
SPECIES
Range Median Range Median
Eastern Kingbird. .| Apr. 22-Apr. 30 | Apr. 28 | Sept. 1-Sept. 24 | Sept. 13
Western Kingbird..| Apr. 23-Apr. 30 | Apr. 28 | Sept. 1-Sept. 26 | Sept. 8
Scissor-tailed
Flycatcher..... Apr. 15-Apr. 28 | Apr. 18 | Sept. 21-Oct. 22 | Oct. 12
Great Crested
Flycatcher..... Apr. 15-May 4 | Apr. 29 | Sept. 1-Sept. 21 | Sept. 9
Eastern Phoebe....| Mar. 3—Mar. 31 | Mar. 22 | Oct. 3-Oct. 27 | Oct. 9
Sayuehoebe: ©... ..... Apr. <4-Apr. 22) Apres 12 Wy un. cca eres eae eee
Eastern Wood
Acadian Flycatcher | Apr. 30-May 19 | May 9 | Sept. 3-Sept.17 | Sept. 4
BEWEGK oi fas it's Apr. 2-May 28 | May 19 | Aug. 30-Sept. 18 | Sept. 6
624 UnIvErsITY OF KAnsAs Pusts., Mus. Nat. Hist.
Number of eggs.—Clutch-size is 5 eggs (4.8, 4-6; 6).
Nests are placed in hollows and crevices in elm, maple, cottonwood, willow,
pear, apple, oak, drain spouts, and, occasionally, “bird houses” made by man,
about 17 feet high (four to 45 feet high).
Eastern Phoebe: Sayornis phoebe (Latham).—This summer resident is
common in eastern Kansas, but is local in the west. Preferred habitat is in
woodland edge and riparian groves, where most birds are found near bridges,
culverts, or isolated outbuildings of man. Temporal occurrence is indicated
in Table 13.
Breeding schedule-—The 186 records of breeding span the period March 21
to July 20 (Fig. 5); the modal date for egg-laying is April 25 (for first clutches)
and June 5 (for second clutches); this species seems to be the only double-
brooded flycatcher in Kansas.
Number of eggs.—Clutch-size is 4 to 5 eggs (4.2, 8-5; 58). The seasonal
progression in clutch-size can be summarized as follows:
March 21-April 10: 4.0 eggs (2 records)
April 11-May 10: 4.4 eggs (37 records)
May 11-June 10: 3.9 eggs (10 records)
June 11-July 20: 3.6 eggs (9 records)
Nests are placed on horizontal, vertical, or overhanging surfaces of culverts,
bridges, houses of man, earthen cliffs, rocky ledges, and entrances to caves, at
an average height of 7.8 feet.
Say Phoebe: Sayornis saya saya (Bonaparte).—This is a common summer
resident in western Kansas, breeding at least east to Cloud County, in open
country. Occurrence in time is listed in Table 13.
Breeding schedule—Ten records of breeding fall in the period May 1 to
July 20; the modal date for egg-laying is in late May.
Number of eggs——Clutch-size is about 5 eggs.
Nests are placed under bridges, in houses, or on cliffsides and earthen
banks.
Acadian Flycatcher: Empidonax virescens (Vieillot)—This is an uncom-
mon summer resident in eastern Kansas, in woodland and riparian habitats.
Temporal occurrence is indicated in Table 18.
Breeding schedule—tThe available records of breeding by this species in
Kansas are too few to indicate reliably the span of the breeding season. In-
formation on hand suggests that Acadian Flycatchers lay most eggs in late
May or early June, and this places their nesting peak some 10 to 20 days
earlier than peaks for Wood Pewees and Traill Flycatchers.
Number of eggs.—Five records show 3 eggs each.
Nests are placed about six feet high on terminal twigs of oak and alder.
Traill Flycatcher: Empidonax traillii trailii (Audubon).—This flycatcher
has only recently been found nesting within Kansas; the species is not in-
cluded in analyses above. Twenty-three nesting records are here reported, for
the species in Kansas City, Jackson and Platte counties, Missouri. Most of
these records are from within a few hundred yards of the political boundary
of Kansas. The Traill Flycatcher is a local summer resident in extreme north-
eastern Kansas (Doniphan County), in wet woodland and riparian groves.
THE BREEDING Birps OF KANSAS 625
Temporal occurrence is not well-documented; first dates run from May 19 to
25; the last dates of annual occurrence, possibly not all for transients, run
from August 14 to September 24.
Breeding schedule—Twenty-three records of breeding are from May 21
to July 10 (Fig. 5); the modal date for egg-laying is June 15.
Number of eggs.—Clutch-size is 3 eggs (3.4, 2-5; 22).
Nests are placed in forks, crotches, and occasionally near trunks, chiefly of
willow, from 4.5 to 12 feet high (averaging six feet).
Eastern Wood Pewee: Contopus virens (Linnaeus).—This summer resident
is common in the east, but is rare in the west. Preferred habitat is in edge of
forest and woodland. Temporal occurrence is indicated in Table 18.
Breeding schedule——Nineteen dates of egg-laying span the period June 1
to July 20 (Fig. 5); the modal date for completion of clutches is June 15, and
more than half of all clutches are laid in the period June 11 to 20.
Number of eggs.—Clutch-size is about 3 eggs.
Nests are placed on upper surfaces of horizontal limbs of oak, elm, and
sycamore, about 22 feet high.
Horned Lark: Eremophila alpestris (Linnaeus ).—Breeding populations are
resident in open country with short or cropped vegetation. E. a. praticola
(Henshaw) lives in the east, and E. a. enthymia (Oberholser) in the west.
Breeding schedule-—Twenty-one records of breeding span the period March
11 to June 10 (Fig. 6); the modal date for egg-laying is March 25. The
histogram (Fig. 6) is constructed on a clearly inadequate sample, and records
of breeding both earlier and later are to be expected. The peak of first nest-
ing activity is probably reasonably well-indicated by the available records.
Number of eggs.—Clutch-size is 8 eggs (3.6, 3-5; 16).
Nests are placed on the ground, usually amid short vegetation such as
cropped prairie grassland or cultivated fields (notably soybeans and wheat),
and occasionally on bare ground.
Tree Swallow: Iridoprocne bicolor (Vieillot).—This is a summer resident
in extreme northeastern Kansas; nesting birds have been found only along
the Missouri River in Doniphan County. Habitat is in open woodland, and in
Kansas is always associated with water. Temporal occurrence in the State
is indicated in Table 14.
TABLE 14.—OcCURRENCE IN TIME OF SUMMER RESIDENT SWALLOWS IN KANSAS
Arrival Departure
SPECIES
Range Median Range Median
Tree Swallow...... Apr. 5-Apr. 30 | Apr. 24 | Sept. 30-Oct. 21 |] Oct. 8
Bank Swallow..... Apr. 9-May 19 | May 7 | Sept. 3-Sept. 20 | Sept. 10
Rough-winged
Swallow....... Mar. 29-—May 30 | Apr. 22 | Sept. 23-Oct. 21 | Oct. 10
Cliff Swallow...... Apr. 14-May 27 | May 11 | Sept. 3-Oct. 25 | Sept. 11
Barn Swallow..... Mar. 31-Apr. 29 | Apr. 21 | Sept. 22-Oct. 25 | Oct. 7
Purple Martin..... Mar. 5-Apr. 9 | Mar. 26 | Aug. 28-Sept. 23 | Sept. 3
626 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Breeding schedule——Eight records of breeding span the period May 21 to
June 20; the modal date for egg-laying is May 25. The small sample may
not accurately reflect the peak of nesting activity.
Number of eggs.—Clutch-size is 5 or 6 eggs (5.5, 5-6; 4).
Nests are placed chiefly in abandoned woodpecker diggings in willows, four
to ten feet high, over water.
Bank Swallow: Riparia riparia riparia (Linnaeus).—This summer resident
is common wherever cut-banks suitable for nesting activities allow relatively
undisturbed behavior. The species is almost always found near water. Tem-
poral occurrence is indicated in Table 14.
Breeding schedule—Sixty records of breeding span the period May 11 to
June 20 (Fig. 6); the modal date for completion of clutches is June 5.
Nearly 75 per cent of all clutches are laid in the period May 21 to June 10.
Under unusual circumstances time of breeding can be greatly delayed; such
circumstances occurred in 1961 in many places along the Kansas River in
eastern Kansas, where the soft, sandy-clay banks were repeatedly washed
away in May and June by high water undercutting the cliffs, Bank Swallows
attempted to work on burrows in late May, but stabilization of the banks
occurred only by late June, and the peak of egg-laying for many colonies was
around July 12. Records for 1961 are omitted from the sample used here
(Fig. 6).
Number of eggs.—Clutch-size is 5 eggs (4.8, 3-7; 60). Yearly clutch-size
at one colony 3 miles east of Lawrence, Douglas County, is as follows:
1959: 5.2, 19 records 1961: 8.7, 11 records
1960: 5.0, 12 records 1962: 4.8, 18 records
The sample for 1961 is that taken in early July when breeding occurred
after a delay of more than a month, as described above.
Nesting chambers are excavated in sandy-clay banks, piles of sand, piles
of sawdust, or similar sites, at ends of tunnels one to more than three feet
in depth from the vertical face of the substrate.
Rough-winged Swallow: Stelgidopteryx ruficollis serripennis (Audubon).—
This summer resident is common in most places; it is not restricted to a single
habitat, but needs some sort of earthen or other substrate with ready-made
burrows for nesting. Temporal occurrence is indicated in Table 14.
Breeding schedule-—The 14 records of breeding are in the period May 11
to June 30; the modal date of egg-laying is June 5. Seventy per cent of all
eggs are laid in the period May 21 to June 10.
Number of eggs.—Clutch-size is 5 eggs (5.0, 4-6; 4).
Nesting chambers are in old burrows of Bank Swallows, Kingfishers, rodents,
or in crevices remaining subsequent to decomposition of roots of plants;
frequently this swallow uses a side chamber off the main tunnel, near the
mouth, of a burrow abandoned or still in use by the other species mentioned
above.
Cliff Swallow: Petrochelidon pyrrhonota pyrrhonota ( Vieillot).—This com-
mon summer resident occurs wherever suitable sites for nests are found. Tem-
poral occurrence is indicated in Table 14.
Breeding schedule.—The 610 records of breeding span the period May 21
to June 80 (Fig. 6); the modal date for egg-laying is June 5, and 85 per cent
THE BREEDING Binns OF KANSAS 627
of all clutches are laid from May 21 to June 10. Such synchronous breeding
activity is probably a function of strong coloniality with attendant “social
facilitation” of breeding behavior.
Number of eggs.—Clutch-size is 5 eggs (4.9, 3-7; 7).
Mch. Apr. May June July Aug.
Ae A era) Beet ere ee) reer
Eremophila alpestris
21
cae subis
.- ae riparia
ie ruficollis
E aes rustica
”
cots Petrochelidon pyrrhonota
oO
30 5 610
Io
20 S a
Oi, <3
>
TE ERE ERRn Aa Al
Mch. Apr. May June July niaar ny alana qu laden
Fic. 6.—Histograms representing breeding schedules
of the Horned Lark and swallows in Kansas. See
legend to Figure 1 for explanation of histograms.
628 UNIVERSITY OF KANSAS PuBts., Mus. Nat. Hist.
Nests are built in mud jugs plastered to vertical rock faces, bridges, culverts,
and buildings from a few feet to more than 100 feet above the ground.
Barn Swallow: Hirundo rustica erythrogaster Boddaert.—This summer resi-
dent is common in most habitats, occurring chiefly about cultivated fields and
pastures. Temporal occurrence is indicated in Table 14.
Breeding schedule.—Sixty-three records of breeding in northern Kansas
span the period May 1 to July 31 (Fig. 6); the modal date for completion
of first clutches is May 25, and that for the second is July 5. The schedule of
breeding in southern Kansas (chiefly Cowley County), to judge by 41 records,
conforms to the one for northern Kansas; the season spans the period May 1
to August 10, and the modal date for first clutches is May 15. The ten-day
lag in peak of first clutches of the northern over the southern sample is about
what would be expected on the basis of differential inception of the biological
growing season from south to north each spring.
Number of eggs.—Clutch-size does not vary geographically, to judge only
from the present samples, and all are included in the listing to follow. The
modal size of clutches is 5 eggs (4.7, 3-7; 43); clutches from the period May 1
to 80 show an average of 5.0 eggs, from June 1 to 20 an average of 4.9 eggs,
and from June 21 to August 10, 4.4 eggs.
Nests are usually placed on horizontal surfaces in barns, sheds, or other
such structures; more rarely they are put on bridges, and less frequently yet
on vertical walls of culverts or sheds.
Purple Martin: Progne subis subis (Linnaeus).—This summer resident is
common in the east but rare in the west. The only documented colony west
of the 99th meridian was in Oberlin, Decatur County (Wolfe, 1961), occupied
some 50 years ago. Temporal occurrence is indicated in Table 14.
Breeding schedule——The breeding season spans the period May 11 to June
20 (Fig. 6); the modal date of egg-laying is June 5, and 57 per cent of all
clutches are laid in the period June 1 to 10.
Number of eggs.—Clutch-size is 5 eggs (4.2, 8-6; 33). Mean clutch-size
is 4.3 eggs in May and 4.2 in June. Adults tend to lay clutches of 5 eggs and
first-year birds clutches of 4. Replacement clutches by birds of any age
tend to be of 3 eggs.
Nests are built of sticks and mud placed in cavities; in Kansas these are
almost always in colony houses erected by man. Use of holes and crevices
in old buildings is known to have occurred on the campus of The University
of Kansas in the nineteen thirties (W. S. Long, 1936, MS), in Oberlin, De-
catur County in 1908-1914 (Wolfe, loc. cit.), and presently in Ottawa, Frank-
lin County (Hardy, 1961).
Blue Jay: Cyanocitta cristata bromia Oberholser.—This resident is common
throughout Kansas in woodland habitats. Most first-year birds move south
in winter, but adults tend to be strictly permanent residents. Groups of ten
to more than 50 individuals can be seen moving south in October and north in
April. All individuals taken from such mobile groups are in first-year feather.
Breeding schedule.—Eighty-three records of breeding span the period April
10 to July 10 (Fig. 7); the modal date of egg-laying is May 15, and about
50 per cent of all clutches are laid in the period May 11-31.
THE BREEDING Brirps OF KANSAS 629
Number of eggs.—Clutch-size is 4 eggs (4.1, 3-6; 15).
Nests are placed from eight to 70 feet high (averaging 24 feet) in forks,
crotches, and on horizontal limbs of elm, maple, osage orange, cottonwood,
and ash.
Black-billed Magpie: Pica pica hudsonia (Sabine).—This resident is com-
mon in western Kansas, along riparian groves and woodland edge. Records of
nesting are from as far east as Clay County. Wolfe (1961) outlines the his-
tory of magpies in Decatur County as follows: the species was purported to
have appeared in rural districts near Oberlin in 1918, but Wolfe saw the birds
only by 1921, at which time he also found the first (used) nests. The first
reported occupied nest was one in Hamilton County in 1925 (Linsdale, 1926).
Earlier records, chiefly of occurrence in winter, can be found in Goss (1891).
Breeding schedule——Fourteen records of breeding span the period April 11
to June 20; the modal date for egg-laying is May 15.
Number of eggs.—There are no data on clutch-size in Kansas; elsewhere
Black-billed Magpies lay 3 to 9 eggs, and clutches of 7 are found most fre-
quently (Linsdale, 1937:104).
Nests are placed from 10 to 18 feet high (averaging 13 feet) in forks or
lateral masses of branches in cottonwood, box elder, ash, and willow.
White-necked Raven: Corvus cryptoleucus Couch.—This summer resident
is common in western Kansas, probably occupying locally favorable sites in
prairie grassland and woodland edge west of a line from Smith to Seward
counties. The species is known to nest in Cheyenne, Sherman, and Finney
counties.
Breeding schedule——There are few data from Kansas; Aldous (1942) states
that the birds begin activities leading to building sometime in April in Okla-
homa; the peak of egg-laying probably occurs in May, which coincides with
the records from Kansas.
Number of eggs.—Outside Kansas, this species lays 3 to 7 eggs; these fig-
ures seem applicable to Kansas, where brood sizes are known to run from 1 to
7 young.
Nests are placed about 20 feet high in cottonwood and other trees.
Common Crow: Corvus brachyrhynchos brachyrhynchos Brehm.—This resi-
dent is common in most of Kansas, but numbers are lower in the west. Distri-
bution in the breeding season is west at least to Cheyenne, Logan, and Meade
counties.
Breeding schedule.—Sixty-nine records of breeding span the period March
10 to May 31 (Fig. 7); the modal date for egg-laying is April 5, and 60 per
cent of all eggs are laid between March 21 and April 10.
Number of eggs.—Clutch-size is 4 eggs (4.2, 3-5; 19).
Nests are placed about 20 feet high in crotches near trunks or heavy
branches of such trees as red cedar, elm, oak, osage orange, cottonwood, honey
locust, box elder, and pine.
Black-capped Chickadee: Parus atricapillus Linnaeus.—This resident is
common north of the southernmost tier of counties, in forested and wooded
areas. P. a. atricapillus Linnaeus occurs chiefly east of the 98th meridian, and
P. a. septentrionalis Harris occurs west of this; a broad zone of intergradation
exists between these two subspecies.
630 UNIVERSITY OF KAnsAs Pusts., Mus. Nat. Hist.
Breeding schedule —Fifty-one records of breeding span the period March
21 to June 10 (Fig. 7); the modal date for laying is April 15, and 64 per cent
of all eggs are laid between April 11 and 30.
Number of eggs.—Clutch-size is 5 eggs (5.4, 4-7; 10).
Nests are placed in cavities about ten feet high (ranging from four to 20
feet) in willow, elm, cottonwood, honey locust, apricot, or nestboxes placed by
man.
Mch, Apr. May June July Aug. Mch. Apr. May JuneJuly Aug. Mch., Apr. ,May June July Aug.
Cyanocitta cristata Nene rufum
83
Corvus brachyrhynachos
65 ¢ Turdus migratorius
Parus atricapillus
5!
Porus bicolor
lage
=
wal” sialis
Lanius ludovicianus
| ee aedon
ey 16
Thryomanes bewicki/ bL’ cee
SCALE
OF CLUTCHES
- ™ ee polyglottos Vireo ae
TS Dumetella carolinensis
Mch. Apr. May June A Acco. Aug. Mch.Apr. May June July Aug.
ie mustelina
Fic. 7.—Histograms representing breeding schedules of crows, chickadees,
wrens, thrashers, thrushes, and their allies in Kansas. See legend to Figure 1
for explanation of histograms.
THE BREEDING Brirps OF KANSAS 631
Carolina Chickadee: Parus carolinensis atricapilloides Lunk.—This resident
is common in the southernmost tier of counties, from Comanche County east,
in forest and woodland edge. Actual records of breeding are from Barber and
Montgomery counties.
Breeding schedule—There are no data on breeding of this species in
Kansas.
Number of eggs.—Clutch-size is about 5 eggs.
Nests are placed in cavities of trees.
Tufted Titmouse: Parus bicolor Linnaeus——This resident is common in the
eastern half of Kansas, in woodlands. Specimens taken in the breeding sea-
son and nesting records come from east of a line running through Cloud,
Harvey, and Sumner counties, and the species probably breeds in Barber
County.
Breeding schedule—Twenty-two records of breeding span the period
March 21 to June 10 (Fig. 7); the modal date for laying is April 25, and 54
per cent of all clutches are laid in the period April 11 to 30.
Number of eggs.—Clutch-size is 4 to 5 eggs (4.5; 6).
Nests are placed in cavities about 12 feet high (ranging from three to 30
feet) in elm, oak, cottonwood, hackberry, redbud, osage orange, and nest-
boxes placed by man.
White-breasted Nuthatch: Sitta carolinensis Latham.—This resident in east-
ern Kansas, in well-developed woodland, is uncommon. S. c. cookei Ober-
holser occurs east of a line running through Douglas and Cherokee counties,
on the basis of specimens taken in the breeding season and actual nesting rec-
ords, and S. c. carolinensis Latham occurs in Montgomery and Labette coun-
ties. S. c. nelsoni Mearns has been recorded in Morton County but probably
does not breed there.
Breeding schedule——Eggs are laid in March and April; young have been
recorded being fed by parents throughout May.
Number of eggs——Clutch-size is between 5 and 10 eggs.
Nests are placed in cavities about 30 feet high in elm and sycamore.
House Wren: Troglodytes aedon parkmanii Audubon.—This summer resi-
dent is common in the east and uncommon in the west. Preferred habitat is
in woodland, brushland, and urban parkland. House Wrens arrive in eastern
Kansas in the period April 3 to 27 (the median is April 19), and are last seen
in autumn in the period September 19 to October 13 (the median is Sep-
tember 30).
Breeding schedule.—The 116 records of breeding span the period April 11
to July 31 (Fig. 7); the modal date of laying is May 20. About 45 per cent
of all clutches are laid in the period May 11 to 31.
Number of eggs.—Clutch-size is 7 eggs (5.8, 3-7; 20). Clutches laid in
May average 6.1 eggs (4-7; 14); those laid in June and July average 5.0 eggs
(38-7; 6).
Nests are placed in cavities about ten feet high (ranging from two to 50
feet) in cottonwood, elm, willow, and a wide variety of structures, mostly
nestboxes, built by man.
Bewick Wren: Thryomanes bewickii Audubon.—This wren is an uncommon
resident in Kansas, except for the northeastern quarter, in woodland under-
632 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
story and brushland. T. b. bewickii Audubon occurs north and east of stations
in Riley, Pottawatomie, Douglas, and Linn counties, and T. b. cryptus Ober-
holser is found south of stations in Greeley, Stafford, and Linn counties; a
zone of intergradation occurs between the two named populations. The species
occupies marginal habitat in most of Kansas and periodically is reduced in
numbers by severe winters.
Breeding schedule-——Twenty-two records of breeding span the period March
21 to July 10 (Fig. 7); the modal date for first clutches is April 15 and for
second clutches June 15.
Number of eggs.—Clutch-size is 5 eggs (5.5, 5-7; 12).
Nests are placed in crevices about five feet high (ranging from zero to nine
feet) in trees (oak, cherry, and pear), boulders, and a wide variety of struc-
tures, some of them nestboxes, built by man; appropriation and modification
of nests of Barn Swallows is known to occur.
Carolina Wren: Thryothorus ludovicianus ludovicianus Latham.—This com-
mon resident of southeastern Kansas in woodland understory and brushland
is uncommon in the northeastern and south-central sectors. Stations of breed-
ing all fall east of a line running through Doniphan, Riley, and western Reno
counties. North and west of southeastem Kansas the Carolina Wren is in
marginal habitat and periodically is reduced in numbers by severe winters.
Breeding schedule.—Fourteen records of breeding span the period April 11
to August 10; the modal date for laying is April 15, to judge only from the
present sample. The species probably breeds also in late March and early
April.
Number of eggs —Clutch-size is 4 eggs (4.2, 3-8; 9).
Nests are placed near the ground in stumps, and a wide variety of struc-
tures built by man, or in crevices in earthen banks.
Long-billed Marsh Wren: Telmatodytes palustris dissaéptus (Bangs ).—This
is an uncommon summer resident in eastern Kansas in and around marshes.
Presumably breeding individuals occur east of stations in Doniphan, Shawnee,
and Sedgwick counties, but actual records of breeding come only from
Doniphan County (Linsdale, 1928:505). First dates of arrival in spring run
from April 19 to 29 (the median is April 22), and dates of last autumnal occur-
rence are from September 26 to October 81 (the median is October 8).
Breeding schedule —Eggs are laid from May to August.
Number of eggs.—Clutch-size is 5 or 6 eggs; the range is from 8 to 10
(Welter, 1935).
Nests are woven of broad-bladed grasses, usually no farther than two feet
from water or mud, suspended in vertical plant stalks or branches in marshes.
Short-billed Marsh Wren: Cistothorus platensis stellaris (Nauman).—This
rare and irregular summer resident in northeastern Kansas occurs in wet
meadowland. Breeding records are available from Douglas and Coffey counties.
Temporal occurrence in the State is at least from April 29 to October 25;
early dates are most likely of transients.
Breeding schedule-—Eggs are laid in late July and August.
Number of eggs.—Clutch-size is 6 or 7 eggs.
Nests are woven of plant fibers and placed in vertically-running stalks and
stems of grasses and short, woody vegetation, within two feet of the ground.
THE BREEDING Birps OF KANSAS 633
Rock Wren: Salpinctes obsoletus obsoletus (Say).—This species is a com-
mon summer resident in western Kansas, in open, rocky country. Specimens
taken in the breeding season and actual nests found come from west of stations
in Decatur, Trego, and Comanche counties. Dates of occurrence are from
April 2 to October 25. Autumnal, postbreeding movement brings the species
east at least to Cloud County (October 7, 8, and 12) and Douglas County
(October 25).
Breeding schedule.—Sixteen records of breeding span the period May 11 to
July 20; the modal date for egg-laying is June 15.
Number of eggs.—Clutch-size is 5 eggs (4.6, 3-7; 5).
Nests are placed in holes in rocks, occasionally in rodent burrows, from
ground level to 80 feet high on faces of cliffs, but there averaging about
20 feet.
Northern Mockingbird: Mimus polyglottos (Linnaeus ).—This is a common
resident in parkland and brushy savannah throughout Kansas. M. p. polyglottos
(Linnaeus) occurs in the east, and M. p. leucopterus (Vigors) in the west;
a broad zone of intergradation exists between the two. Most specimens from
Kansas are of intermediate morphology.
Breeding schedule—Sixty-nine records of breeding span the period April
21 to July 31 (Fig. 7); the modal date for first clutches is June 5, but is
weakly indicated in the histogram (Fig. 7).
Number of eggs.—Clutch-size is 3 eggs (8.5, 3-5; 27). Size of clutch does
not vary seasonally or geographically in the present sample.
Nests are placed about four feet high (two to 10 feet) in osage orange,
red cedar, mulberry, scotch pine, catalpa, cottonwood, rose, and arbor vitae.
Catbird: Dumetella carolinensis (Linnaeus).—This is a common summer
resident in the eastern half of Kansas, but is local in the west, in and near
woodland edge and second-growth. First dates of arrival in spring are from
April 25 to May 14 (the median is May 6), and last dates of autumnal occur-
rence are between September 20 and November 16 (the median is Septem-
ber 26).
Breeding schedule—Seventy-seven records of breeding span the period
May 11 to July 81 (Fig. 7); the modal date for egg-laying is May 25, and
57 per cent of all clutches are laid from May 21 to June 10.
Number of eggs.—Clutch-size is 4 eggs (3.3, 2-5; 43). Clutches laid
between May 11 and June 10 tend to be of 4 eggs (8.5, 2-5; 27), and clutches
laid between June 11 and July 81 tend to be of 3 eggs (2.9, 2-4; 16).
Nests are placed about four feet high in shrubs (rose, lilac, plum, elder-
berry) and about seven feet high in trees (red cedar, honey locust, willow,
elm, apple, and in vines in such trees).
Brown Thrasher: Toxostoma rufum (Linnaeus).—This is a common sum-
mer resident in woodland understory, edge, and second-growth. T. r. rufum
(Linnaeus) occurs in eastern Kansas, to the western edge of the Flint Hills,
and T. r. longicauda Baird occurs west of stations in Decatur, Lane, and Meade
counties; the intervening populations are of intermediate morphologic char-
acter. Some individuals overwinter in Kansas, but most are regular migrants
and summer residents, arriving in spring from April 1 to April 25 (the
634 UNIVERSITY OF KANsAsS Pusts., Mus. Nat. Hist.
median is April 19), and departing in autumn between September 19 and
October 13 (the median is September 28).
Breeding schedule —The 237 records of breeding span the period May 1
to July 20 (Fig. 7); the modal date for egg-laying is May 15, and one-third
of all eggs are laid in the period May 11 to 20.
Number of eggs.—Clutch-size is 4 eggs, ranging from 2 to 5. Seasonal
variation and mean values are shown in Table 15.
TABLE 15.—SEASONAL VARIATION IN CLUTCH-SIZE OF THE BROWN THRASHER
TIME Mean clutch-size Number of records
May lle ten ac teeein oe 3.3 15
May A020, Aa et Bee aps ors 3.9 38
Mayol Slee u scale an: 4.1 13
Junewi=10; 4 eee ee SED 13
June 11=20 sass eee ae Synth) 12
June Zl=sOR peace ee 34: 9
July pela lO. con eee wane ee 3 1
Puly TI ZO eta wees anaes 3 1
Mea Ree Cees eave 2 3.63 102
Nests are placed about four feet high (ranging from 1% to 15 feet) in osage
orange, elm, ornamental evergreens, gooseberry, barberry, honey locust, cotton-
wood, red cedar, rose, plum, honeysuckle, spirea, arbor vitae, willow, oak,
apple, dogwood, and maple.
Robin: Turdus migratorius migratorius Linnaeus.—This summer resident is
common in the east, and is locally common in the west. Some individuals,
usually in small groups, can be seen throughout the winter in eastern Kansas,
and their presence makes it difficult to document dates of arrival and departure
of the strictly summer resident birds; these can be said to arrive in March and
to leave in October, but these indications are the barest approximations.
Breeding schedule-—The 334 records of breeding span the period April 1
to July 20 (Fig. 7); the modal date of laying of first clutches is April 25,
but subsequent peaks are indistinct. Nearly half of all eggs are laid in the
period April 11 to 30.
Number of eggs.—Clutch-size is 8 eggs (3.6,3-6; 57). Clutches laid prior
to May 10 average 3.6 eggs (3-6; 47), and those laid subsequent to May 10
average 3.5 eggs (3-4; 10).
Nests are placed about 18 feet from the ground (ranging from two to 80
feet) in elm, omamental conifers, fruit trees, cottonwood, mulberry, walnut,
hackberry, oak, ash, maple, osage orange, and coffeeberry. Robins rarely
nest in manmade structures, such as on rafters in sheds and barns, on bridge
stringers, and, exceptionally, on electrical utility pole installations.
Wood Thrush: Hylocichla mustelina (Gmelin).—This is an uncommon
summer resident in eastern Kansas, presently absent from the State west of
stations in Cloud and Barber counties. Preferred habitat is found in understory
of forest and woodland. Wood Thrushes appear to have nested in small
numbers as far west as Oberlin, Decatur County (Wolfe, 1961), some 50
years ago, but have since disappeared from such places, probably as a result
THE BREEDING Birps oF KANSAS 635
of progressive modification of watershed and riparian timber by man. First
dates of arrival in spring are from April 19 to May 20 (the median is May 9),
and departure southward is in the period September 8 to October 1 (the median
is September 15).
Breeding schedule.—Thirty-eight records of breeding fall in the period May
11 to August 10 (Fig. 7); the modal date of egg-laying is June 5 for first
clutches. Fifty-five per cent of all eggs are laid between May 21 and June 10.
Number of eggs.—Clutch-size is 3 eggs (8.4, 3-4; 9).
Nests are placed about 11 feet high in elm, dogwood, willow, linden, and
oak.
Eastern Bluebird: Sialia sialis sialis (Linnaeus ).—This locally common resi-
dent and summer resident in eastern Kansas, is only casual west of Comanche
County, in open parkland and woodland edge.
Breeding schedule.—Fifty-four records of breeding span the period April
1 to July 20 (Fig. 7); the modal date for first clutches is April 25 and for
second clutches is June 5.
Number of eggs.—Clutch-size is 5 eggs (4.9, 4-6; 15).
Nests are placed in cavities about eight feet high in trees (elm, box elder,
fruit trees, willow, and ash), and about four feet high in stumps, fence posts,
and nestboxes placed by man.
Blue-gray Gnatcatcher: Polioptila caerulea caerulea (Linnaeus).—This
summer resident is common in eastern Kansas in brushy woodland, edge, and
second growth. Specimens taken in the breeding season and nesting records
come from east of stations in Riley and Cowley counties, but there is a breeding
specimen from Oklahoma just south of Harper County, Kansas. The species
is present from March 80 to September 18.
Breeding schedule—Twelve records of breeding span the period April 20
to June 20; the modal date for egg-laying is May 10.
Number of eggs.—Clutch-size is about 5 eggs.
Nests are placed in forks or on limbs about 17 feet high in oak, elm, honey
locust, red haw, pecan, and walnut.
Cedar Waxwing: Bombycilla cedrorum Vieillot—This waxwing is a rare,
local, and highly irregular summer resident in northeastern Kansas, in wood-
land and forest edge habitats. The known nesting stations are in Wyandotte
and Shawnee counties; six nests have been found in the period 1949 to 1960.
The species has been recorded in all months.
Breeding schedule—Eggs are laid in June and early July.
Number of eggs.—Clutch-size is about 4 eggs (Davie, 1898).
Nests are placed four to 24 feet high in a variety of deciduous and coniferous
trees and shrubs.
Loggerhead Shrike: Lanius ludovicianus Linnaeus.—This common resident
and summer resident favors open country with scattered shrubs and thickets.
L. l. migrans Palmer occurs in eastern Kansas, west to about the 96th meridian,
and L. I. excubitorides Grinnell occurs in western Kansas, east to about the
100th meridian; populations of intermediate character occupy central Kansas.
These shrikes tend to be resident in southern counties, but are migratory in
the north. Dates of spring arrival in Cloud County are between March 9 and
636 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist.
31 (the median is March 21) and the birds leave southward between October
19 and December 19 (the median is November 1).
Breeding schedule—Fifty-seven records of breeding span the period April
1 to June 30 (Fig. 7); the modal date for egg-laying is April 15.
Number of eggs.—Clutch-size is 5 eggs (5.3, 4-7; 32). There is no sea-
sonal variation in the sample.
Nests are placed about six feet high (ranging from four to 10 feet) in
osage orange, small pines, honeysuckle vines, and elm.
Starling: Sturnus vulgaris Linnaeus.—This species is a common resident in
towns and around farms, foraging in open fields of various kinds. Starlings
(introduced into North America from European stocks of S. v. vulgaris) first
appeared in eastern Kansas in the early 1930s and were established as suc-
cessful residents by 1985 or 1936. Occupancy of Kansas to the west took
only a few years. There are no specimens taken in the breeding season or
actual nesting records from southwest of Ellis and Stafford counties; Starlings
seem to be resident in Cheyenne County, but no nesting record exists from
there.
Breeding schedule.—Sixty-seven records of breeding span the period March
1 to June 30 (Fig. 7); the modal date for first clutches is April 15, and for
second clutches is June 5.
Number of eggs.—Clutch-size is 5 eggs (5.2, 4-8; 19).
Nests are placed about 22 feet high (ranging from eight to 50 feet) in
crevices in elm, locust, hackberry, nestboxes placed by man, and in a variety
of other structures of man.
Black-capped Vireo: Vireo atricapilla Woodhouse—This was a summer
resident, apparently of limited distribution but in good numbers, in Comanche
County, in oak woodland and brushland edge. No specimens have been taken
in Kansas since 1885.
Breeding schedule—Eggs are probably laid in May and June. Goss (1891:
851) found a nest under construction on May 11, 1885, and this is the only
nesting record of the species in the State.
Number of eggs.—Clutch-size is about 4 eggs (Davie, 1898).
Nests are placed low, perhaps around four feet high, in deciduous trees
and shrubs ( Davie, op. cit.).
White-eyed Vireo: Vireo griseus noveboracensis (Gmelin).—This is a local
summer resident in eastern Kansas, in woodland and forest edge. Stations
of breeding occurrence are in Doniphan, Douglas, Johnson, Anderson, Labette,
and Montgomery counties. The species is present within the extreme dates
of April 23 to October 5 (Table 16).
Breeding schedule——Ten records of breeding span the period May 10 to
June 30; the modal date for egg-laying is June 10. The present sample is not
adequate to indicate extreme or modal dates with reasonable accuracy.
Number of eggs.—Clutch-size is 4 eggs (3.6, 3-4; 5).
Nests are placed relatively low in forks in trees and shrubs.
Bell Vireo: Vireo bellii bellii Audubon—This summer resident is common
in riparian thickets and second-growth scrub. Temporal occurrence is indi-
cated in Table 16.
Tue BREEDING Birps OF KANSAS 637
Breeding schedule.—Sixty-six records of breeding span the period May 1
to July 20 (Fig. 7); the modal date for egg-laying is May 25, and a little
under 40 per cent of all eggs are laid in the period May 21-31. Renesting
following disruption of first nests is regular, and the small peak in the histogram
in the period June 11-20 is representative of this.
Number of eggs—Clutch-size is 4 eggs (4.6, 3-6; 21). Clutches in May
have an average of 3.7 eggs, and those in June and July 3.6 eggs.
Nests are placed about two feet high (ranging from one to five feet) in
terminal or lateral forks of small branches in elm, hackberry, osage orange,
coralberry, dogwood, plum, honey locust, mulberry, willow, cottonwood, and
box elder.
Yellow-throated Vireo: Vireo flavifrons Vieillot—This is a rare and local
summer resident in deciduous forest and woodland in eastern Kansas. Stations
of breeding occurrence fall east of Shawnee and Woodson counties. Temporal
occurrence is indicated in Table 16.
Breeding schedule-—Eggs are laid at least in May.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are placed 16 to 30 feet high in forks of mature deciduous trees.
Red-eyed Vireo: Vireo olivaceus olivaceus (Linnaeus).—This summer resi-
dent is common in the east, but is local and less abundant in the west, in
woodland and deciduous forest. Temporal occurrence is indicated in Table 16.
TasBLE 16.—OccCURRENCE IN TIME OF SUMMER RESIDENT VIREOS IN KANSAS
| Arrival Departure
SPECIES |
| Range Median Range Median
White-eyed Vireo. .| Apr. 23-May 25 | May 8| Oct. 5 —_|........
8
Bell Wireo;..5:% 5; Apr. 14-May 20 | May Aug. 26-Sept. 27 | Sept. 6
Yellow-throated
Vireo sasetee =. Apr. 27-May 22 | May 7 | Aug. 23-Oct. 1 | Aug. 31
Red-eyed Vireo....| Apr. 21-May 10 | May 4 | Sept. 2-Oct. 7 | Sept. 10
Warbling Vireo....| Apr. 20-May 9 | Apr. 28 | Sept. 2-Oct. 6 | Sept. 9
Breeding schedule-—Eight records of breeding fall in the period May 21
to July 31; most records of egg-laying are in the first week of June.
Number of eggs.—Clutch-size is 4 eggs (4.0, 8-5; 5).
Nests are placed in forks of mature deciduous trees, usually fairly high—
perhaps 15 to 25 feet (Davie, 1898).
Warbling Vireo: Vireo gilvus gilvus (Vieillot)—This summer resident is
common in woodland and forest edge. Temporal occurrence is indicated in
Table 16.
Breeding schedule—Seventeen records of breeding span the period May 1
to June 20, but it is likely that breeding later in June and July will be recorded.
The modal date for egg-laying is June 5, and this seems to be a reliable
index to the major effort in egg-laying in spite of the small sample.
Number of eggs.—Clutch-size is 4 eggs (3.6, 3-4; 5).
638 UNIVERSITY OF KAnsAs Pusts., Mus. Nat. Hist.
Nests are placed three to 25 feet high in a variety of deciduous shrubs
and trees.
Black-and-white Warbler: Mniotilta varia (Linnaeus).—This loca] and un-
common summer resident lives in deciduous forest and woodland. Specimens
taken in the breeding season and actual records of nesting come from Doni-
phan, Douglas, Coffey, Greenwood, Sedgwick, Labette, and Cherokee counties.
Temporal occurrence in the State is indicated in Table 17.
Breeding schedule.—Eggs are laid in May and June.
Number of eggs.—Clutch-size is around 5 eggs ( Davie, 1898).
Nests are placed on the ground, in depressions or niches, under heavy
cover.
Prothonotary Warbler: Protonotaria citrea (Boddaert).—This is a local
summer resident in eastern Kansas, in understory of riparian timber and
swampy woodland. Specimens taken in the breeding season and actual records
of nesting come from Doniphan, Douglas, Linn, and Cowley counties, Tem-
poral occurrence is indicated in Table 17.
Breeding schedule-—Twenty-two records of breeding span the period May
11 to July 10 (Fig. 8); the modal date for egg-laying is June 5, and 75 per
cent of all clutches are laid in the period June 1 to 20.
Number of eggs.—Clutch-size is 5 eggs (4.5, 3-6; 15).
Nests are placed in holes and niches in willow, red haw, elm, and a variety
of stumps, about eight feet high (ranging from five to 20 feet), usually over
water. A pair nested once in a gourd under the eave of a house in Winfield,
Cowley County, and another pair in a tin cup on a shelf at a sawmill (Goss,
ex Long, 1936).
Parula Warbler: Parula americana (Linnaeus).—This summer resident in
eastern Kansas usually can be found in heavy woodland and flood-plain timber.
Specimens taken in the breeding season and actual records of breeding come
from Doniphan, Riley, Douglas, Montgomery, Labette, and Cherokee counties.
Temporal occurrence is indicated in Table 17.
Breeding schedule-—Eggs are laid at least from mid-May to mid-June.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are placed in debris in root tangles along stream banks, and, pre-
sumably, in pendant arboreal lichens.
Yellow Warbler: Dendroica petechia (Linnaeus).—This summer resident
is common in the east, in woodland and riparian growths. D. p. aestiva
(Gmelin) occupies eastern Kansas west at least to Barber County, but it is
not known how far west representatives of this population breed. D. p. mor-
comi Coale breeds in western Kansas. D. p. sonorana Brewster, a name ap-
plicable to Yellow Warblers of the southwestern United States and northern
Mexico, has been considered a “straggler” (Long, 1940) or probable summer
resident (Tordoff, 1956; Johnston, 1960) in southwestern Kansas, on the basis of
one specimen taken on June 24, 1911, at a point two miles south of Wallace,
Wallace County. This specimen, which is pale, was identified in 1935 as
D. p. sonorana by H. C. Oberholser. Specimens taken subsequently from
Cheyenne, Hamilton, and Morton counties in the breeding season can be
referred adequately to D. p. morcomi. Probably the specimen of 1911 is a
pale variant of D. p. morcomi within its normal distributional range.
THE BREEDING Birps OF KANSAS 639
Breeding schedule—Thirty-five records of breeding span the period May 11
to June 20 (Fig. 8); this probably is inadequate to show the extent of the
season, and some egg-laying into July is likely to be found in the future. The
modal date of egg-laying is May 25, and this is likely to be reliable.
Number of eggs.—Clutch-size is 4 eggs (4.2, 3-5; 29).
Nests are placed about nine feet high (ranging from five to 20 feet) in
crotches of trees and shrubs including willow, elderberry, cottonwood, crab-
apple, plum, and coralberry.
Prairie Warbler: Dendroica discolor discolor (Vieillot)—This rare, local
summer resident occurs in deciduous second-growth. The only breeding rec-
ords are from Wyandotte and Johnson counties.
Breeding schedule-—Eggs are laid at least in June.
Number of eggs.—Clutch-size is about 4 eggs (Davie, 1898).
Nests are placed low, perhaps about four feet high, in a wide variety of
small trees and shrubs.
Louisiana Waterthrush: Seiurus motacilla (Vieillot)—This uncommon to
rare summer resident in eastern Kansas lives in woodland understory near
streams. Nesting records come from Douglas, Miami, Linn, and Crawford
counties. Wolfe (1961) reports he found a nest with young near Oberlin,
Decatur County, on June 10, 1910, under an overhanging bank of Sappa Creek;
Decatur County is some 250 miles west of the present western limit of the
breeding range of the Louisiana Waterthrush, and western habitats are not
favorable for their occurrence. Temporal characteristics of their distribution
are indicated in Table 17.
Breeding schedule.—Eggs are laid in May and June.
Number of eggs.—Clutch-size is about 5 eggs (Davie, 1898).
Nests are placed in concealed places in banks or stumps always where it is
wet.
TABLE 17.—OcCCURRENCE IN TIME OF SUMMER RESIDENT Woop WARBLERS IN
KANSAS
Arrival Departure
SPECIES
Range Median Range Median
Black-and-white
Warbler. str. Apr. 2-May 12 | May 5 | Sept. 10-Oct. 14 | Sept. 22
Prothonotary
Warbler. 45.0: Apr. 24—-May 25 | May 8} Aug. 6-Sept.10 | Aug. 22
Parula Warbler....| Apr. 6—-May 5 | Apr. 23 | Sept. 12-Oct. 7 | Sept. 18
Yellow Warbler....| Apr. 21-May 7 | Apr. 30 | Aug. 28-Oct. 1 | Sept. 4
Louisiana
Waterthrush....| Apr. 2-May 2 | Apr. 16 | Aug. ?
Kentucky Warbler | Apr. 24-May 15 | May 3 | Sept. 13
Yellowthroat...... Apr. 21-May 10 | May 3 | Sept. 8-Oct. 3 | Sept. 17
Yellow-breasted
Chatrene eee Ge Apr. 29-May 19 | May 11 | Aug. 29-Oct. 1 | Sept. 8
American Redstart | Apr. 22-May 20 | May 12 | Sept. 1-Oct. 7 | Sept. 10
5—1476
640 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Kentucky Warbler: Oporornis formosus (Wilson).—This is an uncommon
summer resident in eastern Kansas, in deciduous forest and woodland. Speci-
mens taken in the breeding season and actual records of nesting come from
Riley, Doniphan, Douglas, Leavenworth, Linn, Montgomery, and Labette
counties. Temporal occurrence is indicated in Table 17.
Breeding schedule —Eggs are laid in May and June.
Number of eggs.—Clutch-size is 4 or 5 eggs.
Nests are placed near or on the ground, usually at the base of small shrubs
or clumps of grass.
Yellowthroat: Geothlypis trichas (Linnaeus).—This summer resident in
and near marshes is common in the east and is local and somewhat less com-
mon in the west. G. t. brachydactylus (Swainson) breeds east of stations in
Clay, Greenwood, and Montgomery counties, G. t. occidentalis Brewster breeds
west of stations in Decatur, Stafford, and Pratt counties, and the intervening
area is occupied by warblers of intermediate morphologic characters. Tem-
poral occurrence is indicated in Table 17.
Breeding schedule——Nine records of breeding span the period May 11 to
June 10; the modal date of egg-laying is June 1. The season is probably more
extended in time than is indicated by the available records.
Number of eggs.—Clutch-size is 5 eggs (4.8, 4-5; 6).
Nests are placed in cattails and sedges one to two and one-half feet high.
Yellow-breasted Chat: Icteria virens (Linnaeus).—This summer resident
is common in willow thickets and rank second-growth. I. v. virens (Linnaeus)
breeds in eastern Kansas, from Nemaha County south, I. v. auricollis (Deppe)
breeds in western Kansas, from Norton County south, and the intervening
sector is occupied by chats of intermediate morphologic character. Temporal
occurrence is indicated in Table 17.
Breeding schedule-—Twenty-six records of breeding span the period May 11
to July 20 (Fig. 8); the modal date for completion of clutches is June 5.
Forty-two per cent of all eggs are laid in the period June 1 to 10.
Number of eggs.—Clutch-size is 4 eggs (3.9, 3-5; 21). Clutches in May
are larger than those in June and July.
Nests are placed in forks and crotches about three feet high in dogwood,
willow, rose, coralberry, cottonwood, and thistles.
Hooded Warbler: Wilsonia citrina (Boddaert).—This warbler is a rare
summer resident in eastern Kansas, in wet, open woodland. Specimens (a
total of four) taken in the breeding season are from Leavenworth and Shawnee
counties, and the one nesting record is from Anderson County.
Breeding schedule —Eggs are laid at least in May.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are low (some as high as six feet) in woody vegetation.
American Redstart: Setophaga ruticilla ruticilla (Linnaeus ).—This summer
resident occurs locally in woodlands east from stations in Cloud and Sumner
Counties. Temporal occurrence is indicated in Table 17.
Breeding schedule—Eggs are laid in May and June.
Number of eggs.—Clutch-size is about 4 eggs (Davie, 1898), but there
are two records of 5 in Kansas.
THE BREEDING Birps OF KANSAS 641
Nests are placed six to 30 feet high, but usually about 12 feet, in forks or
saddled on a branch, in deciduous trees.
House Sparrow: Passer domesticus (Linnaeus).—This sparrow, introduced
from stocks in Ohio and New York (originally from England and Germany),
has been present since about 1876 in eastern Kansas; it is a common resident
in towns and at farmsteads throughout the state.
Nomenclaturally, House Sparrows in North America consistently have been
Apr. May JunejJuly pAug., .Mch, Apr. May JunaJuly; Aug,Sep.,
Frotonotaria citrea Icterus spurius
22
|
118
Dendroica petechia /cterus galbula
35 83
ee Quiscalus quiscula
/eteria virens 233
26
ae Molothrus ater
87
Passer domesticus mel
men ~
|
Sturnel/a magna
40
pis]
is K Sturnella neglecta
M23
Fichmondena cardinalis
7,
oe)
FHasserina cyanea
piza americana
30 a 4|
20wE Agelaius phoeniceus
10 3° 109
°S
Ons
Wipe lliviay dune uuly Aug. IMch! Apr. ‘May June July Aug.' Sep.
Fic. 8.—Histograms representing breeding schedules of wood warblers, the
House Sparrow, icterids, and cardinal grosbeaks in Kansas. See legend to
Figure 1 for explanation of histograms.
642 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
referred to the European ancestral stocks, P. d. domesticus, but none in North
America today duplicates morphologically the European birds. This is evi-
dence of meaningful adaptation of the North American populations to environ-
ments in which they now live, and continued use of P. d. domesticus in mis-
leading. Studies on local differentiation in North American House Sparrows
are in progress, and when the biology of sparrows in the midwest is better
understood, suitable nomenclatural proposals will be made.
Breeding schedule——Fifty-one records of breeding span the period March
20 to July 20 (Fig. 8); the modal date for laying of first clutches is April 5,
and for second clutches May 5.
Number of eggs ——Clutch-size is 4 eggs (3.9, 3-7; 13).
Nests are placed in niches of various sorts seven to 50 feet high in build-
ings, nestboxes, and trees, or freely situated in forks and crotches of large
trees.
Bobolink: Dolichonyx oryzivorus (Linnaeus).—This species is a rare and
local summer resident, in and about grassy meadows. There are but two sta-
tions of breeding in Kansas: Jamestown State Lake, Cloud County, and Big
Salt Marsh, Stafford County. Temporal occurrence is indicated in Table 18.
Breeding schedule.—Eggs are laid in June.
Number of eggs.—Clutch-size is about 5 eggs.
Nests are placed on the ground amidst grasses.
Eastern Meadowlark: Sturnella magna (Linnaeus ).—This summer resident
and resident is common in eastern Kansas, in moist grassland. S. m. argutula
Bangs occurs in Montgomery, Labette, and Cherokee counties and intergrades
to the north and west with S. m. magna (Linnaeus). Good numbers of birds
are found east of the Flint Hills, but to the west the species is of restricted
and local distribution. Extreme outliers of the species are found no farther
west than stations in Jewell, Stafford, and Barber counties.
Breeding schedule——Forty records of breeding span the period April 10
to July 20 (Fig. 8); the modal date for egg-laying is May 5. Fifty-seven
per cent of all eggs are laid in the period May 1 to 20.
Number of eggs.—Clutch-size is 5 eggs (5.2, 4-7; 26). Prior to May 11,
clutch-size is 5.8 eggs (18 records), and after that date it is 5.1 eggs (13
records).
Nests are placed on the ground, with cover of grasses or forbs.
Western Meadowlark: Sturnella neglecta neglecta (Audubon).—This is a
common resident and summer resident in western Kansas, and is restricted and
local in the east; preferred habitat is in grassy uplands.
Breeding schedule—Twenty-three records of breeding span the period
April 10 to July 30 (Fig. 8); the modal date for egg-laying is May 5 for first
nests and June 5 for second nests.
Number of eggs.—Clutch-size is 4 eggs (4.3, 3-6; 16).
Nests are placed on the ground with cover of grasses or forbs.
Yellow-headed Blackbird: Xanthocephalus xanthocephalus (Bonaparte ).—
This uncommon and local summer resident occurs chiefly in the west, in
marshes. Nesting records are from Wallace, Meade, Barton, Stafford, Doni-
phan, and Douglas counties. Temporal occurrence is indicated in Table 18.
Breeding schedule—Fifty-one records of breeding span the period May 20
Tue BREEDING Birps oF KANSAS 643
to June 30; the modal date of egg-laying is June 5. The sample is probably
not large enough to be wholly reliable.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are placed within a few feet of water in cattail, rush, sedge, and
willow.
Red-winged Blackbird: Agelaius phoeniceus (Linnaeus).—This is a com-
mon summer resident in marshes, wet pasture, and scrubby parkland through-
out the State. A. p. phoeniceus (Linnaeus) occurs in most of Kansas and
A. p. fortis (Ridgway) occurs in the west, east to about Decatur County. A
few birds can be found in eastern Kansas in winter; the full breeding popu-
lation is present between April and October.
Breeding schedule—The 109 records of breeding in Cloud County span
the period May 1 to July 30 (Fig. 8); the modal date for laying is May 25,
and 71 per cent of all eggs are laid in the period May 11 to June 10. Eighty-
eight records of breeding from northwestern Kansas make a histogram almost
exactly duplicating the one from Cloud County.
Number of eggs.—Clutch-size at Concordia, Cloud County, is 4 eggs (3.7,
3-5; 48); in northeastern Kansas mean clutch-size is 3.7 eggs (8-5; 46). For
the total sample, mean clutch-size in May is 4.0 eggs, in June, 3.7 eggs, and
in July, 3.8 eggs.
Nests are placed about four feet high (one to nine feet) in willow, cattail,
sedge, grass, elm, exotic conifer, elderberry, coralberry, buttonbrush, honey-
suckle, smartweed, ash, osage orange, and yellow clover.
In central Kansas red-wings are host to the Brown-headed Cowbird in a
frequency of one parasitized nest out of nine; in northeastern Kansas the
ratio is 1:25.
TABLE 18.—OcCURRENCE IN TIME OF SUMMER RESIDENT ICTERIDS IN KANSAS
Arrival Departure
SPECIES
Range Median Range Median
Bobolink. 32358)... May 4-May 21 | May 11 | Aug. 28-Oct. 1 | Sept. 12
Yellow-headed
Blackbird...... Mar. 31—Apr. 29 | Apr. 19 | Sept. 19-Oct. 18 | Sept. 24
Orchard Oriole..... Apr. 25-May 14 | May 4] Aug. 5-Sept.15 | Aug. 9
Baltimore Oriole...| Apr. 24-May 5 | Apr. 29 | Sept. 6-Sept. 29 | Sept. 10
Common Grackle..} Mar. 2—Mar. 27 | Mar. 17 | Oct. 15—Nov. 14 | Oct. 31
Orchard Oriole: Icterus spurius (Linnaeus).—This summer resident is
common in parkland, woodland, and old second-growth. Temporal occur-
rence is indicated in Table 18.
Breeding schedule—The 118 records of breeding span the period May 11
to August 10 (Fig. 8); the modal date for completion of clutches is June 5,
and 45 per cent of all eggs are laid in the first ten days of June.
Number of eggs.—Clutch-size is 4 eggs (4.1, 3-6; 41). Clutches laid at
the peak of the season average 4.38 eggs (3-6; 26), and replacement clutches
average 3.8 eggs (3-4; 9).
644 UNIVERSITY OF KANsAs Pusts., Mus. Nat. Hist.
Nests are hung about 15 feet high (ranging from six to 55 feet) in elm,
cottonwood, hackberry, locust, catalpa, willow, alder, osage orange, walnut,
pear, linden, and ash.
Baltimore Oriole: Icterus galbula (Linnaeus ).—This common summer resi-
dent is most numerous in the east, in woodland and riparian timber. The
species hybridizes freely with the Bullock Oriole in western Kansas, and indi-
viduals morphologically typical of Baltimore Orioles are rare west of the
100th meridian. Evidence of such hybridization can be found in specimens
taken in eastern Kansas, but the linear nature of distribution along water-
courses to the west restricts gene-flow, and evident hybrids are not yet con-
spicuous. Temporal occurrence is indicated in Table 18.
Breeding schedule.—Eighty-three records of breeding span the period May
11 to July 10 (Fig. 8); the modal date of egg-laying is June 5, and 66 per
cent of all eggs are laid between May 21 and June 10.
Number of eggs.—Clutch-size is 4 eggs.
Nests are hung about 24 feet high (ranging from nine to 70 feet) in elm,
cottonwood, sycamore, maple, and oak.
Bullock Oriole: Icterus bullockii (Swainson)—This summer resident is
common in western Kansas in woodland and riparian situations. The species
hybridizes freely with the Baltimore Oriole, and most Bullock Orioles in
Kansas show evidence of such interbreeding. Almost all records of breeding
come from west of the 100th meridian, but the species in recognizable form
probably breeds locally at least as far east as Stafford County.
Breeding schedule——Few nesting records are available, but these suggest
that the breeding schedule of the Bullock Oriole resembles those of the
preceding two species in Kansas.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are hung about 26 feet high (ranging from 10 to 50 feet) in cotton-
wood, elm, and other large trees.
Common Grackle: Quiscalus quiscula versicolor Vieillot—This summer resi-
dent is common in parkland, and around towns and farms. Most individuals
move out of Kansas in winter, and the temporal occurrence of these birds is
indicated in Table 18.
Breeding schedule.—The 233 records of breeding span the period April 11
to June 80 (Fig. 8); the modal date for egg-laying is May 5, and two-thirds
of all eggs are laid between May 1 and May 20.
Number of eggs.—Clutch-size is 5 eggs (4.5, 8-6; 33). Clutches laid at
the peak of the season average 4.7 eggs (3-6; 21), and those laid as replace-
ment clutches average 4.3 eggs (3-6; 12).
Nests are placed in forks and crotches about 22 feet high (ranging from
six to 50 feet) in elm, red cedar, cottonwood, oak, box elder, and pine.
Brown-headed Cowbird: Molothrus ater ater (Boddaert).—Many indi-
viduals of this common summer resident overwinter in the southern part of the
State and it is difficult to determine dates of arrival and departure in Kansas.
Conspicuous abundance in the north covers the period April to October.
Breeding schedule.—The 141 instances of egg-laying span the period April
21 to July 20 (Fig. 8); the modal date of laying is May 15, and 58 per cent
of all eggs are laid in the period May 11 to June 10. Inception of laying is
THE BREEDING Brirps OF KANSAS 645
here fairly reliably indicated, but in exceptionally early springs laying does
occur earlier; a few eggs were found on April 6, 1963, too late for incorpora-
tion into this report other than in this sentence.
Number of eggs.—Clutch-size in cowbirds is not readily determined. On
the basis of ovarian examination of five females taken in mid-season, the birds
here lay about five eggs at a time. There is no question that the birds are
“double-brooded” in Kansas, and the season is sufficiently long for as many as
five “clutches” to be laid by a given female.
Eggs are laid in nests of some forty species of birds in Kansas; 39 of these
are passerines. No preference for any one species is detectable; the most
frequently parasitized species are simply the common species, and these are
the kinds for which nesting records are easily gathered by man. In the
following list of host species, the names marked with an asterisk are the con-
spicuously parasitized species.
Mourning Dove, Eastern Kingbird, Eastern Phoebe,* Say Phoebe,* Acadian
Flycatcher, Barn Swallow, Horned Lark, Carolina Wren, Rock Wren, Brown
Thrasher,* Mockingbird, Catbird, Wood Thrush,* Eastern Bluebird, Yellow-
throated Vireo, Bell Vireo,* White-eyed Vireo,* Parula Warbler, Yellow
Warbler, Black-and-white Warbler, Kentucky Warbler, Louisiana Waterthrush,
Yellow-breasted Chat, Yellowthroat, Eastern Meadowlark, Western Meadow-
lark, Red-winged Blackbird,* Orchard Oriole,* Cardinal,* Black-headed Gros-
beak, Indigo Bunting,* Blue Grosbeak, Dickcissel,* Pine Siskin,* Rufous-sided
Towhee,* Grasshopper Sparrow, Lark Sparrow,* Chipping Sparrow, Field
Sparrow.*
Scarlet Tanager: Piranga olivacea (Gmelin).—This rare summer resident
in northeastern Kansas occurs in deciduous forest and bottomland timber.
Specimens taken in the breeding season and records of nesting come from
Clay, Doniphan, Douglas, Wyandotte, Johnson, and Linn counties, but the
species probably occupies the entire eastern third of the State. Dates of
arrival in spring are from April 29 to May 25 (the median is May 11), and
dates of departure in autumn are from August 4 to September 23 (the median
is August 10).
Breeding schedule—Six records of breeding fall in the period May 11 to
June 20.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are placed 20 to 35 feet high in elm, linden, hickory, and walnut.
Summer Tanager: Piranga rubra rubra (Linnaeus).—This uncommon sum-
mer resident in eastern Kansas occurs in woodland. Specimens taken in the
breeding season and records of nesting come from east of stations in Doniphan,
Shawnee, and Montgomery counties. Dates of arrival in spring run from
April 24 to May 18 (the median is April 29), and the species departs southward
in September and October.
Breeding schedule.—Eleven records of egg-laying cover the period May 21
to July 20; the modal date for laying is June 5.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are situated ten to 20 feet high on horizontal limbs of large trees.
Cardinal: Richmondena cardinalis cardinalis (Linnaeus).—This species is
a common resident in eastern Kansas, west to about the 99th meridian; west
646 UnIvERSITY OF Kansas Pusts., Mus. Nat. Hist.
of this line the species becomes local and uncommon to rare. Habitat in the
east is found in woodland, edge, second-growth and open riparian timber,
and in the west the species is restricted to riparian growths, chiefly along the
Republican, Solomon, Smoky Hill, Arkansas, and Cimarron rivers, and their
larger tributaries.
Breeding schedule ——The 117 records of breeding span the period April 1
to September 20 (Fig. 8); the modal date for laying of first clutches is May 1,
subsequent to which breeding activity is regular but asynchronous.
Number of eggs.—Clutch-size is 3 eggs (3.5, 3-6; 65). Seasonal variation
in clutch-size is as follows:
Date Mean clutch-size Number of records
v0) 9 Ui IE et ena A enn ee Oe el 3.0 6
April: 2i=sMay: WO Ao seek cee bake 3.8 25
Mayr tleMaysGlee nee ae ee cee 8.8 15
June —junesZ0 sek oee cic eee 3.6 ll
Junes2=Julya20 een ates cen se Got 3.3 a
Nests are placed about five feet high (ranging from 10 inches to 40 feet)
in osage orange, elm, grape, rose, red cedar, coralberry, willow, cottonwood,
gooseberry, oak, elderberry, box elder, arbor vitae, Lombardy poplar, Forsythia,
pines, honeysuckle, wisteria, lilac, red haw, hickory, dogwood, and sycamore.
Rose-breasted Grosbeak: Pheucticus ludovicianus (Linnaeus).—This is a
local and at times common summer resident in eastern Kansas, in woodland,
edge, and riparian timber. Specimens taken in the breeding season and actual
records of breeding come from Clay, Riley, Doniphan, Leavenworth, and
Douglas counties. This species meets and hybridizes with the Black-headed
Grosbeak west of the Flint Hills. Temporal occurrence in the State is indi-
cated in Table 19.
Breeding schedule——Eleven records of breeding span the period May 11
to July 10; the modal date for laying is probably June 5.
Number of eggs.—Clutch-size is 3 or 4 eggs.
Nests are placed in deciduous trees, in forks and crotches six to 30 feet high.
Black-headed Grosbeak: Pheucticus melanocephalus melanocephalus
(Swainson).—This summer resident is common in western Kansas, chiefly
along streams. Individuals referable to this species by sight records alone
breed in fair numbers as far east as Cloud and Sedgwick counties, but to the
east of these stations numbers are reduced, partly as a result of presumed
competition with the Rose-breasted Grosbeak. Hybrids between these two
grosbeaks are regularly produced. The easternmost record of breeding by this
species is at St. Mary’s, Pottawatomie County, where a male was seen as
probably mated with a female Rose-breasted Grosbeak. Temporal occurrence
is indicated in Table 19.
Breeding schedule.—Sixteen records of breeding span the period May 11 to
July 10; the modal date for egg-laying is June 5.
Number of eggs.—Clutch-size is about 4 eggs (3.7, 3-4; 4).
Nests are placed about 12 feet high in a variety of deciduous trees.
Blue Grosbeak: Guiraca caerulea (Linnaeus).—This is a common to un-
common summer resident in most of Kansas, in brushland and streamside
thickets. G. c. caerulea (Linnaeus) breeds in the east, east of stations in
Douglas, Greenwood, and Cowley counties, and G. c. interfusa Dwight and
THE BREEDING Birps OF KANSAS 647
Griscom breeds in the west, west of stations in Cloud, Stafford, and Clark
counties; a broad zone of intergradation exists between the two named popu-
lations. Temporal occurrence is indicated in Table 19.
TABLE 19.—OccCURRENCE IN TIME OF SUMMER RESIDENT CARDINAL GROSBEAKS
IN KANSAS
Arrival Departure
SPECIES
Range Median Range Median
Rose-breasted
Grosbeak...... Apr. 25-May 5] May 2 | Sept. 4-Oct. 1 | Sept. 13
Black-headed
Grosbeak...... Apr. 26-May 11 |} May 5 | Aug. 17-Sept.18 | Sept. 2
Blue Grosbeak... . . Apr. 25—-May 26 | May 13 | Aug. 15-Sept. 3 | Aug. 27
Indigo Bunting....| Apr. 20-May 15 | May 6 | Aug. 23-Oct. 31 | Oct. 1
Lazuli Bunting. ...:] May 5—-May 24 | May 10 |........0.0.2...-}- ones ==
Painted Bunting...) Apr. 30-May 25 | May 9 -].....:.622.5%. S6ulheseee
Dickcisseliiss .5'-. Apr. 21-May 10 | May 4 | Sept. 7-Oct. 11 | Sept. 18
Breeding schedule.—Seven records of breeding span the period May 21 to
June 80; the modal date of laying seems to be in late May or early June.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are placed from three to 30 feet high in a variety of deciduous plants.
Indigo Bunting: Passerina cyanea (Linnaeus).—This summer resident is
common in mixed-field and heavy brushland habitats. The species extends
westerly, in riparian situations, in reduced numbers, ultimately meeting and
hybridizing with the Lazuli Bunting. Specimens referrable to the Indigo
Bunting have been taken as far west as Finney County, but most specimens
from that far west show evidence of interbreeding with Lazuli Buntings.
Temporal occurrence is indicated in Table 19.
Breeding schedule Twenty-four records of breeding span the period May
11 to August 20 (Fig. 8); the modal date for egg-laying is June 15.
Number of eggs.—Clutch-size is 3 eggs (3.1, 2-4; 17).
Nests are placed about three feet high (ranging from one to nine feet) in
coralberry, sumac, thistle, sycamore sprouts, hickory sprouts, grape, elderberry,
cottonwood, dogwood, ragweed, and grasses.
Lazuli Bunting: Passerina amoena (Say).—This uncommon summer resi-
dent of western Kansas occurs in edge habitats and streamside thickets. The
one breeding record is from Morton County, and there is a breeding specimen
taken at Sharon Springs, Wallace County. The species hybridizes with the
Indigo Bunting in the western half of the State. Temporal occurrence in
spring is indicated in Table 19.
Breeding schedule.—Eggs are laid in June and July.
Number of eggs.—Clutch-size is about 4 eggs ( Davie, 1898).
Nests are placed a few feet from the ground, probably much as are nests of
the Indigo Bunting.
Painted Bunting: Passerina ciris pallidior Mearns.—This is an uncommon
648 UNIVERSITY OF KAnsAS Pusts., Mus. Nar. Hist.
summer resident in the southeastern third of Kansas, in edge habitats and
streamside brush. Specimens taken in the breeding season and actual nesting
records come from Douglas, Shawnee, Geary, Barber, and Crawford counties.
Temporal occurrence in spring is indicated in Table 19.
Breeding schedule-—Eggs are laid in June and July.
Number of eggs.——Clutch-size is about 4 eggs (Davie, 1898).
Nests are placed in deciduous shrubs and trees.
Dickcissel: Spiza americana (Gmelin).—This species is a common summer
resident in eastern Kansas and is local and irregular in the west, in grassland
habitats. Temporal occurrence is indicated in Table 19.
Breeding schedule——¥orty-one records of breeding span the period May
1 to July 10 (Fig. 8); the modal date for egg-laying seems to be May 5, but
the curiously abrupt inception of breeding described by this sample suggests
that more records are needed to document fully the breeding schedule of this
species. Breeding in April almost certainly will be found.
Number of eggs.—Clutch-size is about 4 eggs (4.1, 3-5; 14).
Nests are placed about two feet high (ranging from ground level to 12
feet) in grasses, osage orange, sedge, box elder, honey locust, clover, thistle,
and blackberry.
Pine Siskin: Spinus pinus pinus (Wilson).—This irregular summer resident
occurs locally north of the 38th parallel, chiefly around planted conifers.
Known stations of breeding are in Hays, Ellis County, Concordia, Cloud
County, and Onaga and St. Marys, Pottawatomie County.
Breeding schedule—Twelve records of breeding span the period March 11
to May 20 (Fig. 9); most nests have been established in late April or by early
May.
Number of eggs.—Clutch-size is about 4 eggs. Of ten nests examined for
eggs, five had at least one egg of the Brown-headed Cowbird; if it is assumed
that each cowbird egg replaced one of the siskins, mean clutch-size is 3.7 eggs.
Nests are placed about seven feet high (ranging from 3.5 to 18 feet) in
red cedar, exotic conifers, and Lombardy poplar.
American Goldfinch: Spinus tristis tristis (Linnaeus).—This resident is
common in woodland edge, scrubby second-growth, old fields, and riparian
thickets. Occurrence tends to be local and at low density in the southwestern
sector.
Breeding schedule—Twelve records of breeding span the period June 20
to September 10 (Fig. 9); the modal date for laying is August 5.
Number of eggs.—Clutch-size is 4 eggs (4.4, 3-6; 8).
Nests are placed from two to eight feet high in woody or herbaceous vege-
tation.
Red Crossbill: Loxia curvirostra Linnaeus.—This is an uncommon and irreg-
ular winter visitant to Kansas, but it nested once in Shawnee County. L. c.
minor (Brehm), on geographic grounds, probably nested here, but five other
subspecies have been recorded in the State and any one of these might have
undertaken the aberrant breeding.
Breeding record.—Three eggs, set completed March 24, 1917, Shawnee
County; successfully fledged (Hyde, 1917:166).
Tue BREEDING Birps OF KANSAS 649
The species usually lays 4 eggs and places its nests in conifers.
Rufous-sided Towhee: Pipilo erythrophthalmus erythrophthalmus (Lin-
naeus ).—This is an uncommon summer resident in eastern Kansas, in under-
story of woodland and streamside timber. Specimens taken in the breeding
season and actual records of nesting come from east of stations in Cloud,
Marion, and Cherokee counties. Temporal occurrence is indicated in Table
20; records of P. e. arcticus (Swainson) have been eliminated from the sample
as far as has been possible.
Breeding schedule—Nineteen records of breeding span the period April
21 to August 10 (Fig. 9); the modal date for egg-laying is May 5.
Number of eggs.—Clutch-size is 4 eggs (4.0, 3-7; 14).
Nests are placed on the ground, in heavy cover.
Lark Bunting: Calamospiza melanocorys Stejneger.—This species is ordi-
narily a common summer resident in western Kansas, in grassland and open
Mch. Apr. \May June July Aug. Sep.
Spinus pinus
l2
Spinus tristis
I
Pipilo erythophthalmus
a 9
Chondestes grammacus
39
40 |
30 = Spizella pusilla
i
20 az = 29
loos
o x
Mch! Apr. ie ener July ‘Aug. Sep.
Fic. 9.—Histograms representing breeding sched-
ules of cardueline and emberizine finches in Kan-
sas. See legend to Figure 1 for explanation of
histograms.
650 UNIVERSITY OF KANnsAs PuBts., Mus. Nat. Hist.
scrub. Specimens taken in the breeding season and all breeding records ex-
cept one for western Franklin County come from west of stations in Decatur,
Ellis, and Comanche counties. Irregular fluctuations in breeding density have
been recorded from Decatur County (Wolfe, 1961). Temporal occurrence is
indicated in Table 20.
Breeding schedule-—Fourteen records of breeding span the period May 21
to June 20; the modal date of egg-laying cannot be determined from the
present sample.
Number of eggs.—Clutch-size is 4 eggs (4.1, 3-5; 7).
Nests are placed on the ground, at bases of clumps of grasses.
Grasshopper Sparrow: Ammodramus savannarum perpallidus (Coues).—
This species is a local and at times common summer resident throughout
Kansas, in grassland. Temporal occurrence is indicated in Table 20.
Breeding schedule——Seven records of breeding fall in the period May 1 to
June 30; the modal date of laying seems to be about May 21.
Number of eggs.—Clutch-size is 5 eggs (4.8, 4-5; 5).
Nests are placed on the ground or in low vegetation, with cover of grasses
or forbs.
Henslow Sparrow: Passerherbulus henslowii henslowii (Audubon).—This
is an uncommon and local summer resident in eastern Kansas, in grassland.
Breeding records are from Cloud, Shawnee, Douglas, Morris, and Anderson
counties. ‘Temporal occurrence is indicated in Table 20.
Breeding schedule-——Eggs are laid in May and June.
Number of eggs.—Clutch-size is about 5 eggs.
Nests are placed on the ground, usually in bluestem pasture, but in any
case grasses.
Lark Sparrow: Chondestes grammacus (Say).—This is a common summer
resident in grassland edge habitats. C. g. grammacus (Say) breeds east of
the Flint Hills, east of stations in Pottawatomie, Anderson, and Montgomery
counties, and C. g. strigatus Swainson breeds west of stations in Clay, Dickin-
son, Harvey, and Sedgwick counties; specimens from the intervening area are
of intermediate subspecific character. Temporal occurrence is indicated in
Table 20.
Breeding schedule.—Thirty-nine records of breeding span the period May
1 to July 20 (Fig. 9); the modal date for egg-laying is probably May 25, but
the sample may not be reliable in this respect.
Number of eggs.—Clutch-size is 4 eggs (4.1, 3-5; 28).
Nests are usually placed on the ground, in cover of pasture grasses, clover,
thistle, milo maize, and soybean; there is one record of a nest one and one-
half feet high in a small pine.
Cassin Sparrow: Aimophila cassinii (Woodhouse ).—This is a common sum-
mer resident in open scrub and grassland edge, to the south and west of Wal-
lace and Comanche counties. Specimens taken in the breeding season and
actual nesting records are from Wallace, Hamilton, Kearny, Finney, Morton,
and Comanche counties; the A.O.U. Check-list (1957) cites Hays, Ellis
County, as a breeding locality, but it is doubtful that the species now occurs
there.
TuHeE BREEDING BirDs OF KANSAS 651
Breeding schedule —Eggs are laid in May and June.
Number of eggs.—Clutch-size is about 4 eggs.
Nests are placed on the ground, at bases of small bushes.
Chipping Sparrow: Spizella passerina passerina (Bechstein).—This is an
uncommon summer resident in open woodland, second-growth, and edge.
TABLE 20.—OCCURRENCE IN TIME OF SUMMER RESIDENT AMERICAN BUNTINGS
IN KANSAS
Arrival Departure
SPECIES
Range Median Range Median
Rufous-sided
Towhee........ Apr. 2-Apr. 19 | Apr. 9 | Sept. 20-Oct. 8 | Sept. 29
Lark Bunting...... May,, 5—-May 14. |)May 10 |. ...2..4e. 202 5. cesleeeeeine
Grasshopper
Sparrow....... Apr. 12-May 11 | Apr. 29 | Aug. 20-Oct. 6 | Aug. 31
Henslow Sparrow. .| Apr. 14-Apr. 30 | Apr. 22 | Oct. 15 —_........ - -
Lark Sparrow..... Mar. 29-Apr. 21 | Apr. 18 | Sept. 13-Oct. 16 | Oct. 12
Chipping Sparrow..| Mar. 6-Apr. 29 | Apr. 23 | Oct. 3-Nov. 15 | Oct. 20
Field Sparrow. .... Mar. 4-Apr. 28 | Apr. 7 | Oct. 5-Nov. 12 | Oct. 30
S. p. passerina is found east of stations in Barber and Shawnee counties;
Chipping Sparrows are not known to breed farther to the west, but records
for north-central Kansas are likely to be found. The subspecific affinities of
our Chipping Sparrows are entirely with the nominate subspecies, and there
is no basis for earlier reports (Long, 1940; Tordoff, 1956; Johnston, 1960)
that S. p. arizonae Coues (=S. p. boreophila Oberholser) occurs in Kansas.
Breeding schedule.—Nine records of breeding fall in the period May 1 to
May 10, in no way indicating the whole span of the breeding season; the
species probably lays eggs in May and July, as well as in June.
Number of eggs.—Clutch-size is 4 eggs.
Nests are placed four to 40 feet high in evergreens of a variety of kinds.
Field Sparrow: Spizella pusilla (Wilson).—This species is a common sum-
mer resident in grassland and edge habitats. S. p. pusilla (Wilson) breeds in
eastern Kansas chiefly east of the Flint Hills; S$. p. arenacea Chadbourne breeds
in central and western Kansas, intergrading easterly with S. p. pusilla.
Breeding schedule.—Twenty-nine records of breeding span the period April
21 to September 10 (Fig. 9); the modal date for first clutches is May 5.
Number of eggs—Clutch-size is 4 eggs (4.1, 3-5; 21).
Nests are placed about 10 inches high (ranging from ground level to three
feet) in or among coralberry, osage orange, elm, oak, rose, and, once, peony.
Chestnut-collared Longspur: Calcarius ornatus (Townsend).—This was
formerly a summer resident in western Kansas, in short-grass habitat. The
only known nesting area was in the vicinity of Ft. Hays, Ellis County. The
species is to be looked for in prairie with short grass type of vegetation.
652 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
ACKNOWLEDGMENTS
Many persons have contributed field observations such as dates of arrival
and departure for migrants, and the various activities of the breeding cycle
for most of the species here discussed. An alphabetic listing of their names
follows.
Galen Abbot, Ruth Abbot, Ted Anderson, Ted F. Andrews, Jon Barlow,
Amelia Betts, Grace Thompson Bigelow, L. C. Binford, Bessie Boso, William J.
Brecheisen, J. Walker Butin, L. B. Carson, Mrs. Eunice Dingus, Charles S.
Edwards, A. S. Gaunt, Sue Griffith, Mrs. Mary F. Hall, J. W. Hardy, Stanley
Hunter, Katherine Kelley, E. E. Klaas, W. C. Kerfoot, John A. Knouse, Eugene
Lewis, Eulalia Lewis, John Lenz, Nathan H. McDonald, Marno McKaughan,
Merrill McHenry, Robert M. Mengel, Robert Merz, Jim Myers, Mary Louise
Myers, Mrs. Kathryn Nelson, T. W. Nelson, Steven Norris, Dan Michener, P. W.
Ogilvie, Gary C. Packard, Mrs. Marion J. Mengel, Dwight Platt, William
Reynolds, Frank Robl, S. D. Roth, Jr., Nancy Saunders, Richard H. Schmidt,
Marvin D. Schwilling, T. M. Sperry, Steve Stephens, Max Thompson, Fr. Mat-
thew Turk, Emil Urban, J. W. Wallace, H. E. Warfel, A. W. Wiens, Mrs.
Joyce Wildenthal, George Young, and Richard Zenger.
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1956. The molt and testis cycle of the Anna hummingbird. Condor,
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1961. The breeding birds of Decatur County, Kansas: 1908-1915. Bull.
Kansas Ornith. Soc., 12:27-30.
ZUVANICH, J. R.
1963. Forster terns breeding in Kansas. Bull. Kansas Orith. Soc., 14:1-8.
Transmitted November 21, 1963.
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\22.
Vol, 10.
Vols 125 94.,
23.
(Continued from inside of front cover )
Review of the insectivores of Korea. By J. Kno Pies i d Davi k
Johnson. Pp. 549-578. | February 23, ise et ee ee
peciation and evolution o e pygmy mice, genus Baiomys. By Robert L.
Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960. te
Index. Pp. 671-690.
1.
2.
3.
4,
5.
7G:
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.
Comparative breeding behavior of Ammospiza caudacuta and A. maritima.
By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.
ar ee aera cf the Peery ae Kansas Natural History Reservation.
y Henry S. Fitch an ona - McGregor. Pp. 77-
in text, 4 tables. December 31, 1956. i ee oe ae
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-
dae yee asta
irds found on the ctic slope \of- northern Alaska. B ames W. Bee.
Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1938.
The wood rats of Colorado: . distribution and ecology. By Robert B. Finley,
Jr. Pp. 213-552, 34 plates, 8 figures in text, 85 tables. November 7, 1958.
Home ranges and movements of the eastern cottontail in Kansas.. By Donald
W. Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, 1959.
8. ‘Natural history of the salamander, Aneides hardyi. By Richard F. Johnston
10.
and Gerhard A. Schad. Pp. 578-585. October 8, 1959.
A new subspecies of lizard, Cnemidophorus sacki, from Michoacan, México.
By William’ E. Duellman. Pp. 587-598, 2 figures in text. May 2, 1960.
A-taxonomic study of the middle American snake, Pituophis deppei. By
William E, Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960.
Index. Pp. 611-626.
Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960.
ND
Vol. 18.
36
*4,
Functional morphology of three bats: Eumops, Myotis, Macrotus. By. Terry
A. Vaughan. Pp. 1-153; 4 plates, 24 figures in text. July 8, 1959.
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.
The baculum in microtine rodents. By Sydney’ Anderson. Pp. 181-216, 49
figures in text. February 19, 1960.
A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By
Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in
text. May 2, 1960.
5. Natural history of the bell vireo. By Jon C. Barlow. Pp. 241-296, 6 figures
10.
1.
12.
18.
14,
in text. March 7, 1962.
Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C.
Fox. Pp. 297-307, 6 figures in text. May 21, 1962.
Vertebrates from the barrier island of Tamaulipas, México. By Robert K.
Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 309-
345, pls. 5-8. June 18, 1962.
Teeth of Edestid sharks. By Theodore H, Eaton, Jr. Pp. 347-362, 10 fig-
ures in text. October 1, 1962.
Variation in the muscles and nerves of the leg in two genera of. grouse
(Tympanuchus and Pedioecetes). By E. Bruce Holmes. Pp. 363-474, 20
figures. October 25, 1962. F
A new genus of Pennsylvanian Fish (Crossopterygii, Coelacanthiformes )” from
Kansas. By Joan Echols. Pp. 475-501, 7 figures. October 25, 1963.
Observations on the Mississippi Kite in southwestern Kansas. By Henry S.
Fitch. Pp. 508-519. October 25, 1963. ,
Jaw musculature of the Mourning and White-winged doves. By Robert L.
Merz. Pp. 521-551, 22 figures. October 25, 1963. __ :
Thoracic and coracoid arteries in two families of birds, Columbidae and
Hirundinidae. By Marion Anne Jenkinson. Pp. 553-573, 7 figures. March
2 64
as
~The breeding birds of Kansas. By Richard F. Johnston. Pp. 575-655, 10
figures. May 18, 1964. :
Index to come.
1,
2.
3.
Five natural hybrid combinations in minnows (Cyprinidae), By Frank B.
Cross and W.-L. Minckley. Pp. 1-18. June 1, 1960.
A. distributional study of the amphibians of the Isthmus of Tehuantepec,
México. By William E. Duellman. Pp. 19-72, pls. 1-8, 3 figures in text,
A t 16, 1960. ’ ;
A nee subspecies of the slider turtle (Pseudemys scripta) from Coahuila,
México. By John M. Legler. Pp. 73-84, pls. 9-12, 8 figures in text. August
16, 1960. : ty
Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, pls. 13-20,
26 figures ‘in text. November 30, 1960. Re Sie 3
Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and
"Rocky Mountains. By Henry S. Fitch and T. Paul Maslin. Pp. 289-808,
4fi in text. February 10, 1961. j
Fishes of thie Wakartse river in Kansas. By James E. Deacon and Artie L.
Metcalf. Pp. 309-322, 1 figure in text. February 10, 1961.
(Continued on outside of back cover)
7.
8.
9.
10.
(Continued from inside of back cover)
Geographic variation in the North American cyprinid fish, Hybopsis gracilis.
By. Leonard J. Olund and Frank B. Cross. Pp. 328-348, pls. 21-54, a Riratee
a text. Rebrnery 10, 1961. ae
escriptions of two species o ogs, genus Ptychohyla; studies of Ameri-
can hylid frogs, V. By William E. Duellman. Pp. 349-857, pl. 25, 2 figures
in text, April 27, 1961. ¥ 2 si
Fish populations, following ‘a drought, in the Neosho and Marais des Cygnes
rivers of Kansas. By James Everett Deacon, Pp. 359-427, pls. 26-30, 3
figures. August 1], 1961.
Recent soft-shelled turtles of North America (family Trionychidae). By
Rope: Webb. Pp. 429-611, pls, 31-54, 24 figures in text. February
Index. Pp. 613-624.
Vol. 14. 1.
2.
10.
Ly,
12.
13.
14.
15.
16.
17.
Vol. 15. 1.
2.
3.
4,
Neotropical bats from western México. By Sydney Anderson. . 1-8.
October 24, 1960. fae é Hey /pre
Geographic variation in the harvest mouse. Reithrodontomys megalotis, on
the central Great Plains. and in adjacent regions. By J. Knox Jones, Jr.,
and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961.
Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson.
Pp. 29-67, pls..1 and 2, 3 figures in text. July 24, 1961.
A new subspecies of the black myotis (bat) from eastern Mexico. By E.
Beyer Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December
North American yellow bats,! “‘Dasypterus,” anda list of the named kinds
of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr.
Pp. 73-98, 4 figures in text. December 29, 1961.
Natural history of the brush mouse (Peromyscus boylii) in Kansas with
description of a new subspecies. By Charles A. Long. Pp. 99-111, 1 figure
in text. December 29, 1961.
Taxonomic status of some’ mice of the Peromyscus boylii group in, eastern
Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 118-
120, 1 figure in text. December 29, 1961.
A new subspecies of ground squirrel (Spermophilus spilosoma) from Ta-
maulipas, Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962.
Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J.
rox Jones, Jr., and Ticul Alvarez. Pp. 125-133, 1 figure in-text. March ts
A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene,
of Texas. By E. Raymond Hall and Walter W. Dalquest. Pp. 185+138,
2 figures in text.. April 30, 1962.
A new subspecies of wood rat (Neotoma) from northeastern Mexico. By
Ticul Alvarez. Pp. 139-143. April 30, 1962.
Noteworthy mammals from Sinaloa, ‘Mexico, By J. Knox Jones, Jr., Ticul
Alvarez, and M. Raymond Lee. Pp. 145-159, 1 figure in text. May 18, 1962.
A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164,
1 figure in text. May 21, 1962.
The mammals of Veracruz. By E. Raymond Hall and Walter W. Dalquest.
Pp, 165-362, 2 figures. May 20, 1963.
The recent mammals of Tamaulipas, México. By Ticul Alvarez. Pp. 363-
478, 5 figures in text. May 20, 1963. Mt
A new subspecies of the fruit-eating bat, Sturnira ludovici, from western
eas) By J. Knox Jones, Jr. and Gary L. Phillips. Pp. A75-481, March 2,
1 E o
Records of the fossil mammal Sinclairella, Family Apatemyidae, from the
Chadronian and Orellan. By William C. Clemens. Pp. 483-491. March 2,
1964. /
More numbers will appear in volume 14.
The amphibians and reptiles. of Michoacin, México. By ‘William E. Duell-
man. Pp. 1-148, pls. 1-6, 11 figures in text. December 20, 1961.
Some reptiles and amphibians from Korea, By Robert G. Webb, J. Knox
Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962.
‘A new species of frog (Genus Tomodactylus) from western México. By
Robert G: Webb. | Pp. 175-181,.1 figure in text. March 7, 1962. :
Type specimens of amphibians and reptiles in the Museum of Natural His-
tory, the University of Kansas. By William E. Duellman and Barbara Berg.
Pp. 1838-204. October 26,1962.
Amphibians and Reptiles of the Rainforests of Southern El Petén, Guatemala.
aN erway E. Duellman. Pp: 205-249, pls. 7-10, 6 figures in text. October
XK revision of snakes of the genus Conophis (Family Colubridae, from Middle
ScneaG By John Wellman. Pp. 251-295, 9 figures in text. October 4,
968.
A review of the Middle American tree frogs of the genus Ptychohyla. By
Apert pee Duellman. Pp. 297-349, pls. 11-18, 7 figures in text. October
18, 1963.
Natural history of the racer Coluber constrictor. By Henry S. Fitch. Pp.
351-468, pls. 19-22, 20 figures in text. December 30, 1968.
A review of the frogs of the Hyla bistincta group. By William E. Duellman.
Pp. 469-491, 4 figures in text. March 2, 1964.
More numbers will appear in volume 15.
MUS. COMP. ZOOL
LIBRARY
JUL 21 1904
HARVARD
UNIVERSITY OF KANSAS PUBLICATIONSERSITY
MvuSEUM OF NATURAL HISTORY
Volume 12, No. 15, pp. 657-680, 11 figs.
May 18, 1964
The Adductor Muscles of the Jaw
In Some Primitive Reptiles
BY
RICHARD C. FOX
UNIVERSITY OF KANSAS
LAWRENCE
1964
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HisToRY
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 12, No. 15, pp. 657-680, 11 figs.
Published May 18, 1964
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY
HARRY (BUD) TIMBERLAKE, STATE PRINTER
TOPEKA, KANSAS
1964
80-1522
MUS. COMP. ZOOL
LIBRARY,
JUL 21 1964
The Adductor Muscles of the, ay the oc
In Some Primitive Reptiles
BY
RICHARD C. FOX
Information about osteological changes in the groups of reptiles
that gave rise to mammals is preserved in the fossil record, but the
musculature of these reptiles has been lost forever. Nevertheless,
a reasonably accurate picture of the morphology and the spatial
relationships of the muscles of many of these extinct vertebrates
can be inferred by studying the scars or other marks delimiting the
origins and insertions of muscles on the skeletons of the fossils and
by studying the anatomy of Recent genera. A reconstruction built
by these methods is largely speculative, especially when the fossil
groups are far removed in time, kinship and morphology from
Recent kinds, and when distortion, crushing, fragmentation and
overzealous preparation have damaged the surfaces associated with
the attachment of muscles. The frequent inadequacy of such direct
evidence can be partially offset by considering the mechanical de-
mands that groups of muscles must meet to perform a particular
movement of a skeletal member.
Both direct anatomical evidence and inferred functional rela-
tions were used to satisfy the purposes of the study here reported
on. The following account reports the results of my efforts to: 1,
reconstruct the adductor muscles of the mandible in Captorhinus
and Dimetrodon; 2, reconstruct the external adductors of the mandi-
ble in the cynodont Thrinaxodon; and 8, learn the causes of the
appearance and continued expansion of the temporal fenestrae
among the reptilian ancestors of mammals.
The osteology of these three genera is comparatively well-known.
Although each of the genera is somewhat specialized, none seems
to have departed radically from its relatives that comprised the
line leading to mammals.
I thank Prof. Theodore H. Eaton, Jr., for suggesting the study
here reported on, for his perceptive criticisms regarding it, and for
his continued patience throughout my investigation. Financial as-
sistance was furnished by his National Science Foundation Grant
(NSF-G8624) for which I am also appreciative. I thank Dr. Rainer
Zangerl, Chief Curator of Geology, Chicago Museum of Natural
History, for permission to examine the specimens of Captorhinus
(659)
660 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
and Dimetrodon in that institution. I am grateful to Mr. Robert
F. Clarke, Assistant Professor of Biology, The Kansas State Teachers
College, Emporia, Kansas, for the opportunity to study his speci-
mens of Captorhinus from Richard’s Spur, Oklahoma. Special
acknowledgment is due Mr. Merton C. Bowman for his able prepa-
ration of the illustrations.
Captorhinus
The outlines of the skulls of Captorhinus differ considerably from
those of the skulls of the primitive captorhinomorph Protorothyris.
Watson (1954:335, Fig. 9) has shown that in the morphological
sequence, Protorothyris—Romeria—Captorhinus, there has been
flattening and rounding of the skull-roof and loss of the primitive
“square-cut” appearance in transverse section. The quadrates in
Captorhinus are farther from the midline than in Protorothyris, and
the adductor chambers in Captorhinus are considerably wider than
they were primitively. Additionally, the postorbital region of Cap-
torhinus is relatively longer than that of Protorothyris, a specializa-
tion that has increased the length of the chambers within.
In contrast with these dimensional changes there has been little
shift in the pattern of the dermal bones that roof the adductor
chambers. The most conspicuous modification in Captorhinus is
the absence of the tabular. This element in Protorothyris was lim-
ited to the occiput and rested without sutural attachment upon the
squamosal (Watson, 1954:338); later loss of the tabular could have
had no effect upon the origins of muscles from inside the skull roof.
Changes in pattern that may have modified the origin of the ad-
ductors in Captorhinus were correlated with the increase in length
of the parietals and the reduction of the supratemporals. Other
changes that were related to the departure from the primitive
romeriid condition of the adductors included the development of
a coronoid process, the flattening of the quadrate-articular joint,
and the development of the peculiar dentition of Captorhinus.
The adductor chambers of Captorhinus are large. They are cov-
ered dorsally and laterally by the parietal, squamosal, postfrontal,
postorbital, quadratojugal and jugal bones. The chamber extends
medially to the braincase, but is not limited anteriorly by a bony
wall. The occiput provides the posterior limit. The greater part
of the adductor chambers lies mediad of the mandibles and thus
of the Meckelian fossae; consequently the muscles that arise from
the dermal roof pass downward and outward to their insertion on
the mandibular rami.
Appuctor MUSCLES OF JAW, PRimITIvVE REPTILES 661
Mandible
The mandibular rami of Captorhinus are strongly constructed.
Each ramus is slightly convex in lateral outline. Approximately the
anterior half of each ramus lies beneath the tooth-row. This half
is roughly wedge-shaped in its lateral aspect, reaching its greatest
height beneath the short posterior teeth.
The posterior half of each ramus is not directly involved in sup-
porting the teeth, but is associated with the adductor musculature
and the articulation of the ramus with the quadrate. The ventral
margin of this part of the ramus curves dorsally in a gentle arc
that terminates posteriorly at the base of the retroarticular process.
The dorsal margin in contrast sweeps sharply upward behind the
teeth and continues posteriorly in a long, low, truncated coronoid
process.
A prominent coronoid process is not found among the more
primitive members of the suborder, such as Limnoscelis, although
the mandible commonly curves upward behind the tooth-row in
that genus. This area in Limnoscelis is overlapped by the cheek
when the jaw is fully adducted (Romer, 1956:494, Fig. 213), thereby
foreshadowing the more extreme condition in Captorhinus.
The coronoid process in Captorhinus is not oriented vertically,
but slopes inward toward the midline at approximately 45 degrees,
effectively roofing the Meckelian fossa and limiting its opening to
the median surface of each ramus. When the jaw was adducted,
the coronoid process moved upward and inside the cheek. A space
persisted between the process and the cheek because the process
sloped obliquely away from the cheek and toward the midline of
the skull. The external surface of the process presented an area
of attachment for muscles arising from the apposing internal surface
of the cheek.
Palate
The palate of Captorhinus is of the generalized rhynchocephalian
type (Romer, 1956:71). In Captorhinus the pterygoids and pala-
tines are markedly arched and the relatively large pterygoid flange
lies almost entirely below the lower border of the cheek. The
lateral edge of the flange passes obliquely across the anterior lip
of the Meckelian fossa and abuts against the bottom lip of the fossa
when the jaw is closed.
The palatines articulate laterally with the maxillary bones by
means of a groove that fits over a maxillary ridge. This presumably
allowed the halves of the palate to move up and down rather freely.
The greatest amplitude of movement was at the midline. Antero-
662 UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist.
posterior sliding of the palate seems impossible in view of the firm
palatoquadrate and quadrate-quadratojugal articulations.
The subtemporal fossa is essentially triangular, and its broad
end is bounded anteriorly by the pterygoid flange. The fossa is
lateral to much of the adductor chamber; consequently muscles
arising from the parietals passed ventrolaterally, parallel to the
oblique quadrate ramus of the pterygoid, to their attachment on
the mandible.
Musculature
These osteological features indicate that the adductor muscles
of the jaw in Captorhinus consisted of two primary masses (Figs. 1,
2, 3). The first of these, the capitimandibularis, arose from the
internal surface of the cheek and roof of the skull and inserted on
the bones of the lower jaw that form the Meckelian canal and the
coronoid process.
The muscle was probably divided into a major medial mass, the
temporal, and a lesser, sheetlike lateral mass, the masseter. The
Fic. 1. Captorhinus. Internal aspect of skull, showing
masseter, medial adductor, and temporal muscles. Unnum-
bered specimen, coll. of Robert F. Clarke. Richard’s Spur,
Oklahoma. x 2.
Fic. 2. Captorhinus. Internal aspect of skull, showing an-
terior and posterior pterygoid muscles. Same specimen
shown in Fig. 1. x 2
AppUCTOR MUSCLES OF JAw, PrimitivE REPTILES 663
temporal was the largest of the adductors and arose from the lateral
parts of the parietal, the dorsal parts of the postorbital, the most
posterior extent of the postfrontal, and the upper parts of the
squamosal. The muscle may have been further subdivided, but
evidence for subordinate slips is lacking. The fibers of this mass
were nearly vertically oriented in lateral aspect since the parts of
the ramus that are available for their insertion lie within the antero-
posterior extent of the adductor chamber. In anterior aspect the
fibers were obliquely oriented, since the jaw and subtemporal fossa
are lateral to much of the skull-roof from which the fibers arose.
The masseter probably arose from the quadratojugal, the jugal,
and ventral parts of the squamosal, although scars on the quad-
ratojugal and jugal are lacking. The squamosal bears an indistinct,
gently curved ridge, passing upward and forward from the postero-
ventral corner of the bone and paralleling the articulation of the
squamosal with the parietal. This ridge presumably marks the
upper limits of the origin of the masseter from the squamosal.
Fic. 8. Captorhinus.
Cross-section of right
half of skull immedi-
ately behind the ptery-
goid flange, showing
masseter, temporal, and
anterior pterygoid mus-
cles. Same specimen
shown in Fig. l. x 2.
Fic. 4. Captorhinus. Internal aspect of left mandibular
fragment, showing insertion of posterior pterygoid muscle.
KU 8968, Richard’s Spur, Oklahoma. x 2.8.
664 UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist.
The masseter inserted on the external surface of the coronoid
process, within two shallow concavities separated by an oblique
ridge. The concavities and ridge may indicate that the muscle
was divided into two sheets. If so, the anterior component was
wedge-shaped in cross-section, and its thin posterior edge over-
lapped the larger mass that inserted on the posterior half of the
coronoid process.
From a functional standpoint it is doubtful that a major com-
ponent of the adductors arose from the quadrate wing of the
pterygoid, for when the jaw is closed the Meckelian fossa is directly
lateral to that bone. If the jaw were at almost any angle but maxi-
mum depression, the greatest component of force would be mediad,
pulling the rami together and not upward. The mediad component
would increase as the jaw approached full adduction. Neither is
there anatomical evidence for an adductor arising from the quad-
rate wing of the pterygoid. The bone is smooth, hard, and without
any marks that might be interpreted as muscle scars.
The internal adductor or pterygoid musculature in Captorhinus
consisted of anterior and posterior components. The anterior ptery-
goid arose from the lateral edge and the dorsal surface of the
pterygoid flange. The burred dorsal recurvature of the edge re-
sembles that of the flange of crocodiles, which serves as part of the
origin of the anterior pterygoid in those animals. In Captorhinus
the attachment of the anterior pterygoid to the edge of the flange
was probably tendinous, judging from the extent of the develop-
ment of the edge of the flange. From the edge the origin extended
medially across the dorsal surface of the flange; the ridging of this
surface is indistinct, leading to the supposition that here the origin
was more likely to have been fleshy than tendinous.
The anterior pterygoid extended obliquely backward and down-
ward from its origin, passed medial to the temporal muscle and
inserted on the ventral and medial surfaces of the splenial and
angular bones beneath the Meckelian fossa. The spatial relation-
ship between the palate and quadrate-articular joint indicate that
the muscle was probably a minor adductor in Captorhinus.
When the jaw was adducted, the insertion of the anterior ptery-
goid was in a plane nearly level with the origin. Contraction of
the anterior pterygoid when the jaw was in this position pulled the
mandible forward and did not adduct it. Maximum depression of
the mandible produced maximum disparity vertically between the
levels of the origin and insertion. The force exerted by the anterior
ApDDUCTOR MUSCLES OF JAW, PRIMITIVE REPTILES 665
pterygoid upon the mandible when fully lowered most nearly ap-
proached the perpendicular to the long axes of the mandibular
rami, and the resultant force acting on the mandible was adductive.
The adductive component of force therefore decreased as the
jaw swung upward, with the result that the anterior pterygoid could
only have been active in initiating adduction and not in sustaining it.
The evidence regarding the position and extent of the posterior
pterygoid is more veiled. On the medial surface of the mandible,
the prearticular and articular bones meet in a ridge that ventrally
rims the glenoid cavity (Fig. 4). The ridge extends anteriorly and
curves slightly in a dorsal direction and meets the Meckelian fossa.
The curved part of the ridge is made of the prearticular bone alone.
A small hollow above the ridge, anterior to the glenoid cavity, faces
the medial plane of the skull and is bordered by the articular bone
behind and above, and by the Meckelian fossa in front.
The surfaces of the hollow and the prearticular-articular ridge
bear tiny grooves and ridges that seem to be muscle scars. The
entire area of the hollow and its bordering features was probably
the area of insertion of the posterior pterygoid.
However, the area of insertion lies mostly ventral to the articulat-
ing surface of the articular bone and extends but slightly in front
of it. Seemingly little lever effect could be exercised by an adductor
attaching in this position, namely, at the level of the fulcrum of the
mandibular ramus.
The posterior pterygoid muscle probably arose from the anterior
portion of the pterygoid wing of the quadrate, from a ridge on the
ventromedial surface. From the relationship of the muscle to the
articulation of the jaw with the skull, it may be deduced that the
muscle was limited in function to the stabilization of the quadrate-
articular joint by keeping the articular surfaces in close contact
with each other and by preventing lateral slipping.
Finally there is evidence for an adductor between the temporal
and masseter masses. The anterior dorsal lip of the Meckelian
fossa supports a small knob to which this muscle attached, much as
in Sphenodon (Romer, 1956:18, Fig. 12). Presumably the muscle
was sheetlike and attached to the skull roof, medial to the attach-
ment of the masseter.
A pseudotemporal may have been present, but evidence to indi-
cate its extent and position is lacking. The muscle usually arises
from the epipterygoid and nearby areas of the braincase and skull
roof and inserts in the anterior parts of the fossa of the jaw. In
Captorhinus the lateral wing of the pterygoid cuts across the fossa,
666 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
effectively blocking it from the upper and medial parts of the skull,
the areas of origin for the pseudotemporal.
Dimetrodon
The morphology of the skull of Dimetrodon closely resembles
that of the primitive Haptodus (Haptodontinae, Sphenacodonti-
dae), and “hence may be rather confidently described as that of
the family as a whole” (Romer and Price, 1940:285). The major
differences between the two genera are in the increased specializa-
tion of the dentition, the shortening of the lacrimal, and the develop-
ment of long vertebral spines in Dimetrodon. The absence of gross
differences in the areas of the skull associated with the groups of
muscles with which this study is concerned, implies a similarity
in the patterns of musculature between the two groups. Romer
and Price suggest that Haptodus, although too late in time to be
an actual ancestor, shows “all the common features of the Dimetro-
don group on the one hand and the therapsids on the other.” The
adductors of the jaw of Dimetrodon were probably little changed
from those of the Haptodontinae and represent a primitive condi-
tion within the suborder.
Dimetrodon and Captorhinus differ in the bones associated with
the adductor mechanism; the area behind the orbit in Dimetrodon
is relatively shorter, reducing the comparative longitudinal extent
of the adductor chamber. Furthermore, the dermal roof above the
adductor chamber slopes gently downward from behind the orbit
to its contact with the occipital plate in Dimetrodon. Temporal
fenestrae are, of course, present in Dimetrodon.
Musculature
The adductor musculature of the lower jaw in Dimetrodon was
divided into lateral and medial groups (Figs. 5, 6). The lateral
division consisted of temporal and masseter masses. The temporal
arose from the upper rim of the temporal opening, from the lateral
wall of the skull behind the postorbital strut, and from the dorsal
roof of the skull. The bones of origin included jugal, postorbital,
postfrontal, parietal and squamosal. This division may also have
arisen from the fascia covering the temporal opening (Romer and
Price, 1940:53). The muscle passed into the Meckelian fossa of the
mandible and inserted on the angular, surangular, prearticular,
coronoid and dentary bones. Insertion on the lips of the fossa also
probably occurred.
The lateral division arose from the lower rim of the temporal
opening and from the bones beneath. Insertion was in the
ADDUCTOR MUSCLEs OF JAW, PRIMITIVE REPTILES 667
Fic. 5. Dimetrodon. Internal aspect of skull, showing masseter and
temporal muscles. Skull modified from Romer and Price (1940).
Approx. X 4.
Meckelian fossa and on the dorsal surface of the adjoining coronoid
process.
The reconstruction of the progressively widening masseter as it
traveled to the mandible follows from the progressively widening
depression on the internal wall of the cheek against which the
muscle must have been appressed. The depressed surface included
the posterior wing of the jugal, the whole of the squamosal, and
probably the anteriormost parts of the quadratojugal. Expansion
of the muscle rostrally was prevented by the postorbital strut that
protected the orbit (Romer and Price, 1940:53).
The sphenacodonts possess the primitive rhynchocephalian kind
of palate. In Sphenodon the anterior pterygoid muscle arises from
the dorsal surface of the pterygoid bone and from the adjacent
bones. A similar origin suggests itself for the corresponding muscle,
the second major adductor mass, in Dimetrodon.
From the origin the muscle passed posterodorsad and laterad of
the pterygoid flange. Insertion was in the notch formed by the
reflected lamina of the angular, as suggested by Watson (1948).
In Dimetrodon the relationship of the dorsal surface of the palate
and the ventromedial surface of the mandible in front of the articu-
lation with the quadrate is unlike that in Captorhinus. When the
mandible of Dimetrodon is at rest (adducted), a line drawn be-
668 UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist.
tween these two areas is oblique, between 30 and 40 degrees from
the horizontal. Depression of the mandible increases this angle.
The insertion of the anterior pterygoid is thus always considerably
below the origin, permitting the muscle to be active throughout
the movement of the mandible, from maximum depression to com-
plete adduction. This was a major factor in adding substantially
to the speed and power of the bite.
The presence and extent of a posterior pterygoid is more difficult
to assess, because of the closeness of the glenoid cavity and the
raised ridge of the prearticular, and the occupancy of at least part
of this region by the anterior pterygoid. In some specimens of
Dimetrodon the internal process of the articular is double (see
Romer and Price, 1940:87, Fig. 16) indicating that there was a
double insertion here. Whether the double insertion implies the
insertion of two separate muscles is, of course, the problem. Divi-
sion of the pterygoid into anterior and posterior portions is the
reptilian pattern (Adams, 1919), and such is adhered to here, with
the posterior pterygoid arising as a thin sheet from the quadrate
wing of the pterygoid and the quadrate, and inserting by means
of a tendon on the internal process of the articular, next to the
insertion of the anterior pterygoid.
Watson (1948) has reconstructed the musculature of the jaw in
Dimetrodon with results that are at variance with those of the
present study. Watson recognized two divisions, an inner temporal
Fic. 6. Dimetrodon. Internal aspect of
right cheek, showing anterior and posterior
pterygoid muscles. Skull modified from
Romer and Price (1940). Approx. X H.
ADDUCTOR MUSCLES OF JAW, PRIMITIVE REPTILES 669
and an outer masseteric, of the capitimandibularis, but has pictured
them (830: Fig. 4; 831: Fig. 5C) as both arising from the inner
surface of the skull roof above the temporal opening. But in
Captorhinus the masseter arose from the lower part of the cheek
close to the outer surface of the coronoid process. Watson has
shown (1948:860, Fig. 17B) the same relationship of muscle to
zygoma in Kannemeyeria sp. It is this arrangement that is also
characteristic of mammals and presumably of Thrinaxodon. In
view of the consistency of this pattern, I have reconstructed the
masseter as arising from the lower wall of the cheek beneath the
temporal opening.
Watson's reconstruction shows both the temporal and masseter
muscles as being limited anteroposteriorly to an extent only slightly
greater than the anteroposterior diameter of the temporal opening.
The whole of the posterior half of the adductor chamber is un-
occupied. More probably this area was filled by muscles. The
impress on the inner surface of the cheek is evident, and the extent
of both the coronoid process and Meckelian opening beneath the
rear part of the chamber indicate that muscles passed through this
area.
Watson remarked (1948:829-830) that the Meckelian opening in
Dimetrodon “is very narrow and the jaw cavity is very small. None
the less, it may have been occupied by the muscle or a ligament
connected to it. Such an insertion leaves unexplained the great
dorsal production of the dentary, surangular and coronoid. This
may merely be a device to provide great dorsal-ventral stiffness to
the long jaw, but it is possible and probable that some part of the
temporal muscle was inserted on the inner surface of the coronoid.
Indeed a very well-preserved jaw of D. limbatus? (R. 105: PI. I,
Fig. 2) bears a special depressed area on the outer surface of the
extreme hinder end of the dentary which differs in surface modelling
from the rest of the surface of the jaw, has a definite limit anteriorly,
and may represent a muscle insertion. The nature of these inser-
tions suggests that the muscle was already divided into two parts,
an outer masseter and an inner temporalis.” But, unaccountably,
Watson’s illustration (1948:830, Fig. 4) of his reconstruction limits
the insertion of the temporal to the anterior limit of the Meckelian
opening and a part of the coronoid process above it. No muscle
is shown entering the Meckelian canal. It seems more likely that
the temporal entered and inserted in the canal and on its dorsal
lips. The masseter inserted lateral to it, over the peak of the
coronoid process, and overlapping onto the dorsalmost portions of
670 UNIVERSITY OF KAnsAs Pusts., Mus. Nat. Hist.
its external face, as Watson has illustrated (Plate I, middle fig.).
I am in agreement with Watson’s reconstruction of the origins
for both the anterior and posterior pterygoid muscles. On a func-
tional basis, however, I would modify slightly Watson’s placement
of the insertions of these muscles. Watson believed that the jaw
of Dimetrodon was capable of anteroposterior sliding. The articular
surfaces of the jaws of Dimetrodon that I have examined indicate
that this capability, if present at all, was surely of a very limited
degree, and in no way comparable to that of Captorhinus. The
dentition of Dimetrodon further substantiates the movement of the
jaw in a simple up and down direction. The teeth of Dimetrodon
are clearly stabbing devices; they are not modified at all for grinding
and the correlative freedom of movement of the jaw that that func-
tion requires in an animal such as Edaphosaurus. Nor are they
modified to parallel the teeth of Captorhinus. The latter’s diet is
less certain, but presumably it was insectivorous (Romer, 1928).
With the requisite difference in levels of origin and insertion of
the anterior pterygoid in Dimetrodon insuring the application of
force throughout the adduction of the jaws, it would seem that the
whole of the insertion should be shifted downward and outward
in the notch. If this change were made in the reconstruction, the
anterior pterygoid would have to be thought of as having arisen by
a tendon from the ridge that Watson has pictured (1948:828, Fig. 3)
as separating his origins for anterior and posterior pterygoids. The
posterior pterygoid, in turn, arose by tendons from the adjoining
lateral ridge and from the pterygoid process of Romer and Price.
Tendinous origins are indicated by the limitations of space in this
area, by the strength of the ridges pictured and reported by Watson,
and by the massiveness of the pterygoid process of Romer and Price.
Discussion
A comparison of the general pattern of the adductor musculature
of Captorhinus and Dimetrodon reveals an expected similarity. The
evidence indicates that the lateral and medial temporal masses were
present in both genera. The anterior pterygoid aided in initiating
adduction in Captorhinus, whereas in Dimetrodon this muscle was
adductive throughout the swing of the jaw. Evidence for the
presence and extent of a pseudotemporal muscle in both Capto-
rhinus and Dimetrodon is lacking. The posterior division of the
pterygoid is small in Captorhinus. In Dimetrodon this muscle has
been reconstructed by Watson as a major adductor, an arrangement
that is adhered to here with but slight modification.
Appuctor MuscLes oF JAw, PRIMITIVE REPTILES 671
The dentition of Captorhinus suggests that the jaw movement
in feeding was more complex than the simple depression and ad-
duction that was probably characteristic of Dimetrodon and sup-
ports the osteological evidence for a relatively complex adductor
mechanism.
In Captorhinus the presence of an overlapping premaxillary beak
bearing teeth that are slanted posteriorly requires that the mandible
be drawn back in order to be depressed. Conversely, during
closure, the jaw must be pulled forward to complete full adduction.
The quadrate-articular joint is flat enough to permit such antero-
posterior sliding movements. The relationship of the origin and
insertion of the anterior pterygoid indicates that this muscle, in-
effective in maintaining adduction, may well have acted to pull
the mandible forward, in back of the premaxillary beak, in the last
stages of adduction. Abrasion of the sides of the inner maxillary
and outer dentary teeth indicates that tooth-to-tooth contact did
occur. Whether such abrasion was due to contact in simple vertical
adduction or in anteroposterior sliding is impossible to determine,
but the evidence considered above indicates the latter probability.
Similarities of Protorothyris to sphenacodont pelycosaurs in the
shape of the skull and palate already commented upon by Watson
(1954) and Hotton (1961) suggest that the condition of the ad-
ductors in Dimetrodon is a retention of the primitive reptilian
pattern, with modifications mainly limited to an increase in size
of the temporalis. Captorhinus, however, seems to have departed
rather radically from the primitive pattern, developing specializa-
tions of the adductors that are correlated with the flattening of the
skull, the peculiar marginal and anterior dentition, the modifications
of the quadrate-articular joint, and the development of the coronoid
process.
Thrinaxodon
The evidence for the position and extent of the external adductors
of the lower jaw in Thrinaxodon was secured in part from dissec-
tions of Didelphis marsupialis, the Virginia opossum. Moreover,
comparison of the two genera reveals striking similarities in the
shape and spatial relationships of the external adductors. These
are compared below in some detail.
The sagittal crest in Thrinaxodon is present but low. It arises
immediately in front of the pineal foramen from the confluence of
bilateral ridges that extend posteriorly and medially from the base
of the postorbital bars. The crest diverges around the foramen,
672 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
reunites immediately behind it, and continues posteriorly to its
junction with the supraoccipital crest (Estes, 1961).
In Didelphis the sagittal crest is high and dorsally convex in
lateral aspect, arising posterior to and medial to the orbits, reaching
its greatest height near the midpoint, and sloping down to its termi-
nation at the supraoccipital crest. Two low ridges extend poste-
riorly from the postorbital process to the anterior end of the sagittal
crest and correspond to ridges in similar position in Thrinaxodon.
The supraoccipital crest flares upward to a considerable extent
in Thrinaxodon and slopes posteriorly from the skull-roof proper.
The crest extends on either side downward to its confluence with
the zygomatic bar. The area of the crest that is associated with
the temporal musculature is similarly shaped in Didelphis.
The zygomatic bar in each genus is stout, laterally compressed,
and dorsally convex on both upper and lower margins. At the back
of the orbit of Thrinaxodon, the postorbital process of the jugal
extends posterodorsally. At this position in Didelphis, there is but
a minor upward curvature of the margin of the bar.
In Thrinaxodon the dorsal and ventral postorbital processes, aris-
ing from the postorbital and jugal bones respectively, nearly meet
but remain separate. The orbit is not completely walled off from
the adductor chamber. The corresponding processes in Didelphis
are rudimentary so that the confluence of the orbit and the adductor
chamber is complete.
The adductor chamber dorsally occupies slightly less than half
of the total length of the skull of Thrinaxodon; in Didelphis the
dorsal length of the chamber is approximately half of the total
length of the skull.
The coronoid process in Thrinaxodon sweeps upward postero-
dorsally at an angle oblique to the long axis of the ramus. Angular,
surangular and articular bones extend backward beneath and
Fic. 7. Thrinaxodon. Showing masseter and temporal muscles.
Skull after Romer (1956). Approx. x Yo.
ADDUCTOR MUSCLEs OF JAW, PRIMITIVE REPTILES 673
medial to the process. The process extends above the most dorsal
point of the zygomatic bar, as in Didelphis. The mandibular ramus
is ventrally convex in both genera.
The relationships described above suggest that Thrinaxodon and
the therapsids having similar morphology in the posterior region
of the skull possessed a temporal adductor mass that was split into
major medial and lateral components (Fig. 7). The more lateral of
these, the masseter, arose from the inner surface and lower margin
of the zygomatic bar and inserted on the lateral surface of the coro-
noid process.
The medial division or temporal arose from the sagittal crest and
supraoccipital crest and the intervening dermal roof. The muscle
inserted on the inner and outer surfaces of the coronoid process
and possibly on the bones beneath.
Thrinaxodon represents an advance beyond Dimetrodon in sev-
eral respects. The zygomatic bar in Thrinaxodon extends relatively
far forward, is bowed outward and dorsally arched. Consequently,
the masseter was able to extend from an anterodorsal origin to a
posterior and ventral insertion. The curvature of the jaw trans-
forms the anterodorsal pull of the muscle into a dorsally directed
adductive movement regardless of the initial angle of the jaw. This
is the generalized mammalian condition.
With the development of the secondary palate the area previously
available for the origin of large anterior pterygoid muscles was
reduced. The development of the masseter extending postero-
ventrally from an anterior origin presumably paralleled the reduc-
tion of the anterior pterygoids. The therapsid masseter, as an
external muscle unhindered by the crowding of surrounding organs,
was readily available for the many modifications that have been
achieved among the mammals.
In the course of synapsid evolution leading to mammals, the
temporal presumably became the main muscle mass acting in ad-
duction of the lower jaw. Its primacy is reflected in the phyletic
expansion of the temporal openings to permit greater freedom of
the muscles during contraction. In the synapsids that lead to mam-
mals, there is no similar change in the region of the palate that can
be ascribed to the effect of the pterygoid musculature, even though
these adductors, like the temporal, primitively were subjected to
severe limitations of space.
Didelphis
Dissections reveal the following relationships of the external ad-
ductors of the jaw in Didelphis marsupialis (Fig. 8).
674 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
1. MASSETER
Origin: ventral surface of zygomatic arch.
Insertion: posteroventral and lateroveniral surface of mandible.
2. EXTERNAL TEMPORALIS
Origin: sagittal crest; anteriorly with internal temporalis from frontal
bone; posteriorly with internal temporalis from interparietal bone.
Insertion: lateral surface of coronoid process of mandible.
8. INTERNAL TEMPORALIS
Origin: sagittal crest and skull roof, including posterior two-thirds of
frontal bone, whole of parietal, and dorsalmost portions of squamosal
and alisphenoid.
Insertion: medial surface of coronoid process; dorsal edge of coronoid
process.
Fic. 8. Didelphis marsupialis. Showing masseter and
temporal muscles. Skull KU 3780, 1 mi. N Lawrence,
Douglas Co., Kansas. x %.
Temporal Openings
In discussions of the morphology and functions of the adductor
mechanism of the lower jaw, the problem of accounting for the
appearance of temporal openings in the skull is often encountered.
Two patterns of explanation have evolved. The first has been the
attempt to ascribe to the constant action of the same selective force
the openings from their inception in primitive members of a
phyletic line to their fullest expression in terminal members. Ac-
cording to this theory, for example, the synapsid opening appeared
originally to allow freer expansion of the adductor muscles of the
jaw during contraction, and continued selection for that character
caused the openings to expand until the ultimately derived therapsid
or mammalian condition was achieved.
The second course has been the attempt to explain the appear-
ance of temporal openings in whatever line in which they occurred
by the action of the same constant selective force. According to
the reasoning of this theory, temporal fenestration in all groups was
ADDUCTOR MUSCLES OF JAW, PRIMITIVE REPTILES 675
due to the need to decrease the total weight of the skull, and
selection in all those groups where temporal fenestration occurs
was to further that end.
Both of these routes of inquiry are inadequate. If modern views
of selection are applied to the problem of explaining the appearance
of temporal fenestrae, the possibility cannot be ignored that:
1. Selective pressures causing the inception of temporal fenestrae differed
from those causing the continued expansion of the fenestrae.
2. The selective pressures both for the inception and continued expansion
of the fenestrae differed from group to group.
8. Selection perhaps involved multiple pressures operating concurrently.
4. Because of different genotypes the potential of the temporal region to
respond to selective demands varied from group to group.
Fic. 9. Captorhinus. Diagram, showing
some hypothetical lines of stress. Approx.
<a.
Fic. 10. Captorhinus. Diagram, Fic. 11. Captorhinus. Diagram,
showing areas of internal thickening. showing orientation of sculpture.
Approx. X 1. Approx. X l.
Secondly, the vectors of mechanical force associated with the
temporal region are complex (Fig. 9). Presumably it was toward
a more efficient mechanism to withstand these that selection on the
cheek region was operating. The simpler and more readily analyzed
of these forces are:
1. The force exerted by the weight of the skull anterior to the cheek and
the distribution of that weight depending upon, for example, the length of the
snout in relation to its width, and the density of the bone.
2. The weight of the jaw pulling down on the suspensorium when the jaw
is at rest and the compression against the suspensorium when the jaw is ad-
ducted; the distribution of these stresses depending upon the length and
breadth of the snout, the rigidity of the anterior symphysis, and the extent of
the quadrate-articular joint.
676 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
3. The magnitude and extent of the vectors of force transmitted through the
occiput from the articulation with the vertebral column and from the pull of
the axial musculature.
4. The downward pull on the skull-roof by the adductor muscles of the
mandible.
5. The lateral push exerted against the cheek by the expansion of the
mandibular adductors during contraction.
6. The necessity to compensate for the weakness in the skull caused by the
orbits, particularly in those kinds of primitive tetrapods in which the orbits
are large.
The distribution of these stresses is further complicated and
modified by such factors as:
1. The completeness or incompleteness of the occiput and the location and
extent of its attachment to the dermal roof.
2. The size and rigidity of the braincase and palate, and the extent and
rigidity of their contact with the skull.
The stresses applied to the cheek fall into two groups. The first
includes all of those stresses that ran through and parallel to the
plane of the cheek initially. The weight of the jaw and snout, the
pull of the axial musculature, and the necessity to provide firm
anchorage for the teeth created stresses that acted in this manner.
The second group comprises those stresses that were applied
initially at an oblique angle to the cheek and not parallel to its
plane. Within this group are the stresses created by the adductors
of the jaw, pulling down and medially from the roof, and sometimes,
during contraction, pushing out against the cheek.
It is reasonable to assume that the vectors of these stresses were
concentrated at the loci of their origin. For example, the effect of
the forces created by the articulation of the jaw upon the skull
was concentrated at the joint between the quadrate, quadratojugal,
and squamosal bones. From this relatively restricted area, the
stresses radiated out over the temporal region. Similarly, the
stresses transmitted by the occiput radiated over the cheek from
the points of articulation of the dermal roof with the occipital
plate. In both of these examples, the vectors paralleled the plane
of the cheek bones. Similar radiation from a restricted area, but
of a secondary nature, resulted from stresses applied obliquely to
the plane of the cheek. The initial stresses caused by the adductors
of the jaw resulted from muscles pulling away from the skull-roof;
secondary stresses, created at the origins of these muscles, radiated
out over the cheek, parallel io its plane.
The result of the summation of all of those vectors was a complex
grid of intersecting lines of force passing in many directions both
ADDUCTOR MUSCLES OF JAW, PRIMITIVE REPTILES 677
parallel to the plane of the cheek and at the perpendicular or at an
angle oblique to the perpendicular to the plane of the cheek.
Complexities are infused into this analysis with the division of
relatively undifferentiated muscles into subordinate groups. The
differentiation of the muscles was related to changing food habits,
increased mobility of the head, and increase in the freedom of move-
ment of the shoulder girdle and forelimbs (Olson, 1961:214). As
Olson has pointed out, this further localized the stresses to which
the bone was subjected. Additional localization of stresses was
created with the origin and development of tetrapods (reptiles)
that were independent of an aquatic environment and were sub-
jected to greater effects of gravity and loss of bouyancy in the
migration from the aqueous environment to the environment of air.
The localization of these stresses was in the border area of the
cheek, away from its center.
What evidence is available to support this analysis of hypo-
thetical forces transmitted through the fully-roofed skull of such
an animal as Captorhinus?
It is axiomatic that bones or parts of bones that are subject to
increased stress become thicker, at least in part. This occurs
ontogenetically, and it occurs phylogenetically through selection.
Weak bones will not be selected for. Figure 10 illustrates the pat-
tern of the areas of the skull-roof in the temporal region that are
marked on the internal surface by broad, low thickened ridges.
The position of these ridges correlates well with the position of the
oriented stresses that were presumably applied to the skull of
Captorhinus during life. It can be seen from Figure 10 that the
central area of the cheek is thinner than parts of the cheek that
border the central area. The thickened border areas were the
regions of the cheek that were subjected to greater stress than the
thin central areas.
External evidence of stress may also be present. The pattern of
sculpturing of Captorhinus is presented in Figure 11. The longer
ridges are arranged in a definite pattern. Their position and di-
rection correlates well with the thickened border of the cheek, the
region in which the stresses are distinctly oriented. For example,
a ridge is present on the internal surface of the squamosal along its
dorsal border. Externally, the sculptured ridges are long and
roughly parallel, both to each other and to the internal ridge.
The central area of the cheek is characterized by a reticulate pat-
tern of short ridges, without apparent orientation. The thinness
of the bone in this area indicates that stresses were less severe here.
678 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist.
The random pattern of the sculpture also indicates that the stresses
passed in many directions, parallel to the plane of the cheek and
obliquely to that plane.
Possible Explanation for the Appearance of Temporal Openings
Bone has three primary functions: support, protection and par-
ticipation in calcium metabolism. Let us assume that the require-
ments of calcium metabolism affect the mass of bone that is se-
lected for, but do not grossly affect the morphology of the bones
of that mass. Then selection operates to meet the needs for
support within the limits that are set by the necessity to provide
the protection for vital organs. After the needs for protection are
satisfied, the remaining variable and the one most effective in
determining the morphology of bones is selection for increased
efficiency in meeting stress.
Let us also assume that bone increases in size and/or compactness
in response to selection for meeting demands of increased stress,
but is selected against when requirements for support are reduced
or absent. Selection against bone could only be effective within
the limits prescribed by the requirements for protection and calcium
metabolism.
We may therefore assume that there is conservation in selection
against characters having multiple functions. Since bone is an
organ system that plays a multiple role in the vertebrate organism,
a change in the selective pressures that affect one of the roles of
bone can only be effective within the limits set by the other roles.
For example, selection against bone that is no longer essential for
support can occur only so long as the metabolic and protective
needs of the organism provided by that character are not compro-
mised. If a character no longer has a positive survival value and is
not linked with a character that does have a positive survival value,
then the metabolic demands for the development and maintenance
of that character no longer have a positive survival value. A useless
burden of metabolic demands is placed upon the organism because
the character no longer aids the survival of the organism. If selec-
tion caused, for example, muscles to migrate away from the center
of the cheek, the bone that had previously provided support for
these muscles would have lost one of its functions. If in a popula-
tion of such individuals, variation in the thickness of the bone of
the cheek occurred, those with thinner bone in the cheek would be
selected for, because less metabolic activity was diverted to building
and maintaining what is now a character of reduced functional
AppucTor MUSCLES OF JAW, PRIMITIVE REPTILES 679
significance. A continuation of the process would eliminate the
bone or part of the bone in question while increasing the metabolic
efficiency of the organism. The bone is no longer essential for
support, the contribution of the mass of bone to calcium metabolism
and the contribution of this part of the skeleton to protection have
not been compromised, and the available energy can be diverted
to other needs.
The study of Captorhinus has indicated that the central area of
the cheek was subjected to less stress than the border areas. A
similar condition in basal reptiles may well have been present. A
continued trend in reducing the thickness of the bone of the cheek
in the manner described above may well have resulted in the ap-
pearance of the first reptiles with temporal fenestrae arising from
the basal stock.
Such an explanation adequately accounts for an increased selec-
tive advantage in the step-by-step thinning of the cheek-wall prior
to the time of actual breakthrough. It is difficult to see the ad-
vantage during such stages if explanations of weight reduction or
bulging musculature are accepted.
After the appearance of temporal fenestrae, selection for the
classical factors is quite acceptable to explain the further develop-
ment of fenestration. The continued enlargement of the temporal
fenestrae in the pelycosaur-therapsid lineage undoubtedly was
correlated with the advantages accrued from securing greater space
to allow increased lateral expansion of contracting mandibular ad-
ductors. Similarly, weight in absolute terms can reasonably be
suggested to explain the dramatic fenestration in the skeletons of
many large dinosaurs.
Literature Cited
ApAMs, L. A.
1919. Memoir on the phylogeny of the jaw muscles in recent and fossil
vertebrates. Annals N. Y. Acad. Sci., 28:51-166, 8 pls.
EsTEs, R.
1961. Cranial anatomy of the cynodont reptile Thrinaxodon liorhinus.
Bull. Mus. Comp. Zool., 125(6):165-180, 4 figs., 2 pls.
Horton, N.
1960, The chorda tympani and middle ear as guides to origin and develop-
ment of reptiles. Evolution, 14(2):194-211, 4 figs.
Otson, E. C. ;
1961. Jaw mechanisms: rhipidistians, amphibians, reptiles. Am. Zoolo-
gist, 1(2):205-215, 7 figs.
Romer, A. S. ,
1928. Vertebrate faunal horizons in the Texas Permo-Carboniferous red-
beds. Univ. Texas Bull., 2801:67-108, 7 figs.
1956. Osteology of the reptiles. Univ. Chicago Press, xxii-+ 772 pp.,
248 figs.
680 UNIVERSITY OF Kansas PuBLs., Mus. Nat. Hist.
Romer, A. S. and Price, L. I.
1940. Review of the Pelycosauria. Geol. Soc. Amer. Special Papers, No.
28,x + 538 pp., 71 figs., 46 pls.
Watson, D. M. S.
1948. Dicynodon and its allies. Proc. Zool. Soc. London, 118:823-877,
20 figs., 1 pl.
1954. On Bolosaurus and the origin and classification of reptiles. Bull.
Mus. Comp. Zool., 111(9):200-449, 87 figs.
Transmitted December 5, 1963.
XO
30-1522
INDEX TO VOLUME 12
mus. COMP. ZOOL
New systematic names are in boldface type
abieticola, Dryocopus pileatus, 620
abbreviatus, Microtus (Stenocranius),
187
Acacia Greggii, 27
acadicus, Aegolius, 618
Acanthostega, 220
Accipiter
cooperi, 607
striatus, 607
velox, 607
Acer negundo, 248
Actitis macularia, 613
acuflavidus, Thalasseus sandvicensis,
330
acuta, Anas, 604
Adenostoma fasciculatum, 10
aedon, Troglodytes, 631
Aegolius acadicus, 618
aerodynamic considerations in bat
flight, 38
aestiva, Dendroica petechia, 638
Agassizodus, 349, 351, 352
Agelaius
fortis, 643
megapotamus, 334
phoeniceus, 643
agrestis, Microtus, 185, 189, 192, 203,
206, 207
aikeni, Otus asio, 617
Aimophila cassinii, 650
Aistopoda, 159
Aix sponsa, 605
Ajaia ajaja, 319
alascensis, Lemmus trimucronatus,
194
alba,
Crocethia, 325
Tyto, 617
albifrons, Sterna, 328, 614
albus, Casmerodius, 603
Alca torda, 367
alcomi, Microtus pennsylvanicus, 206
aleyon, Megaceryle, 620
alexandrinus, Charadrius, 612
alle, Mergulus, 367
alleni, Neofiber, 191, 209, 211
alpestris, Eremophila, 332, 625
altiloquus, Vireo, 266
Ambrosia psilostachya, 313
Ambystoma,
middle ear, 163, 164
rib-bearer, 169
Ambystomidae, 163, 175
LIBRARY
DEC 31 ls09
HARVARD
UNIVERSITY
americana,
Aythya, 605
Fulica, 612
Parula, 638
Recurvirostra, 326, 613
Spiza, 648
Ulmus, 248
americanus,
Coccyzus, 616
Numenius, 322, 613
Ammodramus
perpallidus, 650
savannarum, 650
amoena, Passerina, 647
amosus, Microtus montanus, 189, 204
Amphibamus, 157-159, 172, 177
Amphibia,
modem, ancestry of, 157
primitive, 219, 236
Anas
acuta, 604
clypeata, 605
discors, 605
platyrhynchos, 604
ancestory of modern Amphibia, 157
Anderson, Sydney, The baculum in
microtine rodents, 183
Andropogon scoparius, 248
annectens, Spermophilus spilosoma,
335
antillarum, Sterna albifrons, 328
Antrozous pallidus, 25, 34, 120, 124
Anura,
development of vertebrae, 166
resemblances to Urodela, 177
apiana, Salvia, 10
Archilochus colubris, 619
archticus,
Mormon, 367
Pipilo erythrophthalmus, 649
arcuatus,
Coelacanthus, 499
Synaptotylus, 480
Ardea
herodias, 318, 602
occidentalis, 319
treganzai, 31
wardi, 319
arenacea, Spizella pusilla, 651
Arenaria
interpres, 325
morinella, 325
—Univ. Kansas Pusts. Mus. Nat. Hist., Vou. 12, 1959-1964.
(681)
682
argentatus, Larus, 40, 326
argutula, Sternella magna, 642
arizonae, Spizella passerina, 651
Artemisia, 505
californica, 10
Artibeus hirsutus, 7
arvalis, Microtus, 185, 189, 192, 202
Arvicola richardsoni, 192, 199
Ascaphus, 159
foot, 173
larva, 176
pectoral girdle, 171, 172
ribs, vertebrae, 166
aserriensis, Chordeiles minor, 330, 331
asiatica, Zenaida, 523, 556
Asio
flammeus, 618
otus, 618
wilsonianus, 618
asio, Otus, 617
ater, Molothrus, 289, 644
athabascae, Clethrionomys gapperi,
195
athalassos, Sterna albifrons, 329, 614
atratus, Coragyps, 607
atricapilla, Vireo, 249, 256, 636
atricapilloides, Parus carolinensis, 631
atricapillus, Parus, 629
atricilla, Larus, 327
attwateri, Tympanuchus cupido, 367
Aulacomys, 199
aura, Cathartes, 320, 605
auratus,
Colaptes, 620
Mesocricetus, 186
auriculata, Zenaidura, 523, 536
aurita, Zenaida, 523, 536
auritus,
Phalacrocorax, 602
Plecotus, 120
Axelia, 482
Aythya
americana, 605
valisneria, 605
baculum in microtine rodents, 183
Barlow, Jon C.
Natural history of the Bell Vireo,
Vireo bellii Audubon, 243
Barrier Island of Tamaulipas, México,
birds, 316
mammals, 334
plant associations, 312
reptiles, 314
vertebrates, 311
Bartramia longicauda, 613
bat,
big-eared, 34
leaf-nosed, 8
pallid, 25, 34
western mastiff, 7
Batis maritima, 313
UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist,
bats,
adaptations for flight, 114
mechanics of flight, 119
Myology, 67
osteology, 44
Bell Vireo, natural history of, 243-293
courtship, 267
egg laying, incubation, 278
general behavior, 252
habitat, 248
nesting, 272
nestling period, 283
parasitism by cowbird, 289, 291,
292
seasonal movement, 250
singing, 255
territoriality, 259
berlandieri, Gopherus, 314
bewickii, Thryomanes, 332, 631, 632
bicincta, Treron, 563
bicolor,
Iridoprocne, 556, 625
Parus, 631
birds of Kansas, breeding, 577
distributional limits reached in
Kansas, 584
grassland species, 580
limnic species, 580
woodland species, 579
xeric scrub species, 581
zoolgeographical categories, 593
Bombinator, 171, 172
Bombycilla cedrorum, 635
bondae, Molossus, 7
Borborocoetes larva, 176
borealis, Buteo jamaicensis, 607
boreophila, Spizella passerina, 651
boreus, Myiarchus crinitus, 623
Borrichia fructescens, 313
Botaurus lentiginosus, 604
Bouteloua gracilis, 506
brachydactylus, Geothlypis trichas,
640
brachyrhynchos, Corvus, 629
Bradyodonti, 349
brasiliensis, Tadarida, 7, 20
brevis, Myotis velifer, 7
breweri, Microtus, 187
bromia, Cyanocitta cristata, 628
Bubo virginianus, 617
Buchloé dactyloides, 506
bulleri, Macrotus mexicanus, 7
bullockii, Icterus, 644
Buteo
borealis, 607
jamaicensis, 607
lineatus, 608
platypterus, 608
regalis, 609
swainsoni, 608
Butorides virescens, 603
INDEX TO VoLUME 12
cachinnans, Gallinula chloropus, 612
caerulea,
Florida, 319, 603
Guiraca, 646
Polioptila, 635
cafer, Colaptes, 620
Calamospiza melanocorys, 649
Calcarius omatus, 651
calendula, Regulus, song, 257
californianus, Geococcyx, 330, 617
californica, Artemisia, 10
californicus,
Clethrionomys occidentalis, 196
Eumops perotis, 7
Larus, 342
Lepus, 835
Macrotus, 5, 6, 7
Microtus, 205
Myotis, 26
Callipepla squamata pallida, 610
Campodus, 349-355
agassizianus, 350, 351
canicendts; Richmondena cardinalis,
83
Canis latrans, 338
capitata, Lespedeza, 248
Caprimulgus
carolinensis, 618
vociferus, 618
Captorhinus, 659
skull, 660
jaw muscles, 662
Carabidae in stomachs of bats, 35
cardinalis, Richmondena, 334, 645
carolina, Porzana, 320, 611
carolinensis,
Caprimulgus, 618
Dumetella, 633
Parus, 631
Sitta, 631
Zenaidura macroura, 615
carolinus, Centurus, 620
carpus in Amphibia, 173
Caseidae, 301
Casmerodius
albatus, 603
egretta, 603
caspia, Hydroprogne, 328
Cassia fasciculata, 313
Cassidix
mexicanus, 3338, 334
prosopidicola, 333, 334
cassinii, Aimophila, 650
Casuarius, 867
Cathartes
aura, 320
teter, 605
Catoptrophorus semipalmatus, 323
eras, Melanerpes erythrocephalus,
cave Myotis, 8
flight of, 26
cedrorum, Bombycilla, 635
683
celatus, Phenacomys intermedius, 188
Centurus
carolinus, 620
zebra, 620
Cercidium floridum, 28
Cereus giganteus, 533.
Chaerophon luzonus, 7
Chaetura pelagica, 41, 619
Chalcophaps indica, 563
chapmani, Chordeiles minor, 619
Charadrius
alexandrinus, 612
hiaticula, 321
semipalmatus, 321
tenuirostris, 612
vociferus, 612
wilsonia, 321
chihi, Plegadis, 604
Chilonycteris personata, 7
Chilotus oregoni, 191, 199, 200
chinensis, Streptopelia, 563, 564
Chionomys
d, 202
ongicaudus, 191, 202
nivalis, 191, 202
roberti, 202
Chlidonias
niger, 327, 614
surinamensis, 327, 614
chloropus, Gallinula, 612
Choeronycteris mexicana, 7
Chondestes
grammacus, 650
strigatus, 650
Chondrichthyes, 349
Chordeiles
aserriensis, 330, 331
chapmani, 619
howelli, 619
minor, 330, 331, 619
neotropicalis, 331
chrotorrhinus, Microtus, 192, 211
chrysolopha, Eudyptes, 367
cinerascens, Myiarchus, 331
cinereus, Lasiurus, 7
Circus
cyaneus, 609
hudsonius, 609
ciris, Passerina, 647
Cistothorus
platensis, 632
stellaris, 632
citrea, Protonotaria, 638
citrina, Wilsonia, 640
Clepsydrops, 497
Clethrionomys, 194, 211
athabascae, 195
gapperi, 188, 192, 195
californicus, 196
galei, 195
glareolus, 185, 188, 192
occidentalis, 185, 188, 192, 195
684 UNIVERSITY OF KANSAS PuBts., Mus. Nat. Hisrv,
Clethrionomys—Concluded
rufocanus, 185, 188, 192, 196, 197
rutilus, 188, 192, 194, 195
clypeata, Anas, 605
Cnemidophorus gularis, 316
coccinea, Richmondena cardinalis, 334
Coccothraustes coccothraustes, 524
Coccyzus
americanus, 616
erythrophthalmus, 616
Coelacanthi, 478
Coelacanthoidei, 478
Coelacanthus, 492, 494-496, 499
arcuatus, 499
newelli, 499
coibensis, Molossus, 7
Colaptes
auratus, 620
cafer, 620
collaris, 620
luteus, 620
colchicus, Phasianus, 610
Colinus
taylori, 610
texanus, 320
virginianus, 610
collaris, Colaptes cafer, 620
Colobomycter pholeter, 301
colubris, Archilochus, 619
Columba
guinea, 538
livia, 538, 556, 563, 614
palumbus, 525
Columbidae,
jaw muscles, 524-531
relationship with Hirundinidae,
565
serology, 537, 538
thoracic and coracoid arteries,
555-572
Commicarpus scandens, 313
compactus, Dipodomys ordii, 338
Conocarpus erectus, 313
Contopus virens, 625
cooperi,
Accipiter, 607
Synaptomys, 183, 188, 192, 212
Copodontidae, 349
Coragyps atratus, 607
Corus drummondi, 248
coronarius, Microtus, 187, 192
corrugatus,
Agassizodus, 352
Orodus, 350, 351
Corvus
brachyrhynchos, 629
cryptoleucus, 332, 629
imparatus, 332
monedula, 367
Corynorhinus townsendii, 34
Crataegus mollis, 248
crenulata, Fadenia, 353, 354
crenulatus, Edestus, 356
crinitus, Myiarchus, 623
cristata, Cyanocitta, 628
Crocethia alba, 325
Crossopterygii, 236, 477, 478
Croton
capitatus, 312
punctatus, 312
Cryptobranchus, 165
cryptoleucus, Corvus, 332, 629
cryptus, Thryomanes bewickii, 332,
632
cunicularia, Speotyto, 617
cupido, Tympanuchus, 367, 609
curtatus, Lagurus, 183, 210, 211
curti, Lepus californicus, 335, 340
curvirostra, Loxia, 648
cyanea, Passerina, 647
cyaneus, Circus, 609
Cyanocitta, 253
bromia, 628
cristata, 628
Dasypus
novemcinctus, 334
mexicanus, 334
davyi, Pteronotus, 7
dawsoni, Clethrionomys rutilus, 195
Delorhynchus priscus, 299-302
deltoides, Populus, 248
Dendrocopos
medianus, 622
pubescens, 622
villosus, 622
Dendroica
aestiva, 638
discolor, 639
morcomi, 638
parasitism by cowbird, 291
petechia, 638
sonorana, 638
diaphora, Eremophila alpestris, 332
Dichromanassa rufescens, 319
Dichrostonyx
groenlandicus, 188, 192, 193, 212
richardsoni, 193
rubricatus, 193
torquatus, 193
Didelphis
jaw muscles, 673, 674
marsupialis, 671, 672
Dimetrodon, 659
jaw muscles, 666
skull, 666
Diplocercidae, 478, 496, 497, 499
Diplocercides kayseri, 482
Diplocercidoidei, 478, 496
Diplocercinae, 478, 497, 499
Diplurus, 496
Dipodomys
compactus, 338, 341
ordii, 338, 341
INDEX TO VOLUME 12
Dipodomys—Concluded
parvabullatus, 337, 341
sennetti, 341
discolor,
Dendroica, 639
parasitism by cowbirds, 291
discors, Anas, 605
dissacptns, Telmatodytes palustris,
6
divaricata, Larrea, 28
Dolichonyx oryzivorus, 642
Dolomys, 213
domesticus, Passer, 641
dominica, Pluvialis, 41
Doves,
jaw muscles, 524-531
thoracic and coracoid arteries, 555-
572
Dromaeus, 367
Dromiceius, 367
drummondi, Cornus, 248
Dryocopus
abieticola, 620
pileatus, 620
Dumetella carolinensis, 633
duodecimcostatus, Pitymys, 209
Dvinosaurus, 162
ear in Amphibia, 161
Eaton, Theodore H., Jr.
The ancestry of modern Amphibia:
a review of the evidence, 157
A new order of fishlike Amphibia
from the Pennsylvanian of Kan-
sas, 219
Teeth of Edestid sharks, 349
Echols, Joan
A new genus of Pennsylvanian fish
(Crossopterygii, Coelacanthi-
formes ) from Kansas, 477
ecordi, Edaphosaurus, 497
Edaphosaurus ecordi, 497
Edestidae, teeth of, 349-361
Edestus, 361
crenulatus, 356
mirus, 355, 3856
newtoni, 355, 357
egretta, Casmerodius albus, 603
elegans,
Rallus, 611
Rhabdoderma, 477, 480, 483-486,
488, 495, 498
Ellobius, 213
Empidonax
trailli, 624
virescens, 624
endocranium, Coelacanth, 480-483
Enstoma exaltatum, 312
enthymia, Ermophila alpestries, 625
Eoanura, 159
Eothyrididae, 301, 302
Eptesicus fuscus, 7
685
Eremophila
alpestris, 332, 625
diaphora, 332
enthymia, 625
giraudi, 332
praticola, 625
Ereunetes pusillus, 314
Eriogonum fasciculatum, 10
erythrocephalus, Melanerpes, 622
erythrogaster, Hirundo rustica, 628
erythrophthalmus,
Coccyzus, 616
Pipilo, 649
erythrorhynchus, Pelecanus, 317
Eucnide urens, 35
Eudyptes chrysolopha, 367
Eugyrinus, 159, 177
Eumops
californicus, 7
foraging habits and flight, 14
perotis, 5, 6, 7, 38
roosting habits and terrestrial loco-
motion, 9
Eupoda montana, 612
Euporosteus, 478
Eusthenopteron, 495
evolutionary considerations, bats, 127
excelsus, Procyon lotor, 338
exilis, Ixobrychus, 604
Fadenia, 350, 352-354
crenulata, 353, 354
gigas, 353, 354
Falco
anatum, 609
peregrinus, 609
sparverius, 19, 609
fasciculatum,
Adenostoma, 10
Eriogonum, 10
fatioi, Microtus (Pitymys), 190, 192,
209
fedoa, Limosa, 322, 341
Fimbristylis castanea, 312
finitus, Microtus pennsylvanicus, 206
fissure deposits, Fort Sill, 304-306
Fitch, Henry S.
Observations on the Mississippi
Kite in southwestern Kansas,
505
flammeus, Asio, 618
flava, Triodia, 248
Flaveria oppositifolia, 312
flavifrons, Vireo, 271
flight,
adaptations for, in bats, 114
mechanics of, in bats, 119
Florida caerulea, 319, 603
floridum, Cercidium, 28
forficata, Muscivora, 331, 623
formosus, Oporornis, 640
686
forsteri, Sterna, 328
Fort Sill fissure deposits, 304-306
fortis,
Agelaius phoeniceus, 643
Microtus, 185, 189
fowl mite, 287
Fox, Richard C.
Two new Pelycosaurs from the
Lower Permian of Oklahoma,
299
The adductor muscles of the jaw in
some primitive reptiles, 659
Fregata
magnificens, 318
rothschildi, 318
Fulica americana, 612
fulviventer, Microtus, 192
fulvus, Pteronotus davyi, 7
fuscipes, Procyon lotor, 338
fuscus, Eptesicus, 7
fusus, Microtus montanus, 189, 204
Gaillardia pulchella, 312
galbula, Icterus, 674
galei, Clethrionomys gapperi, 195
Gallinula
cachinnans, 612
chloropus, 612
gallopavo, Meleagris, 610
gapperi, Clethrionomys, 188, 192, 195
gamettense, Hesperoherpeton, 219-
239, 497
Gavia immer, 368
Geococcyx californianus, 330, 617
Geomys
personatus, 335-337
tropicalis, 311, 336
georgica, Strix varia, 618
Geothlypis
brachydactylus, 640
occidentalis, 640
trichas, 640
gigas, Fadenia, 353, 354
gilmorei, Microtus oeconomus, 205
gilvus, Vireo, 637
giraudi, Eremophila alpestris, 332
glareolus, Clethrionomys, 185, 188,
192
Gleditsia triacanthos, 247
Glossophaga
leachii, 7
soricina, 7
Gopherus berlandieri, 314
gossii, Synaptomys cooperi, 194
grammacus, Chondestes, 650
gregalis, Microtus (Stenocranius),
190, 192
Greggii, Acacia, 27
groenlandicus, Dichrostonyx, 188,
192.5212,
grouse, muscles, 396
muscles and nerves of leg, 365-472
UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist,
grouse—Concluded
nerves, 376
skeleton, 369, 375
variation of muscles and nerves of
leg, 447-456
guatemalensis, Microtus
tomys), 191, 192, 198
gud, Microtus, 202
guentheri, Microtus, 185, 189, 192,
203
guinea, Columba, 538
Guiraca
caerulea, 646
interfusa, 646
gularis, Cnemidophorus, 316
(Herpe-
Haematopus ostralegus, 320
hasbroucki, Otus asio, 617
haydeni, Microtus ochrogaster, 207
Helicoprion, 349, 354, 355, 357-361
ferrieri, 358
henslowii, Passerherbulus, 650
herodias, Ardea, 318, 602
Hesperoherpeton
for limb, 233
garnettense, 219-239, 497
pectoral girdle, 231
ribs, 231
skull, 220
vertebrae, 229
Hesperoherpetonidae, 239
hesperus, Pipistrellus, 26, 36
hiaticula, Charadrius, 321
himantopus, Micropalama, 326
hipposideros, Rhinolophus, 120
hirsutus, Artibeus, 7
Hirundinidae,
myology and angiology, 557-559
relationship with Columbidae, 565
thoracic and coracoid arteries, 555
Hirundo
erythrogaster, 628
rustica, 556
hirundo, Sterna, 328
hispidus, Sigmodon, 339
hoactli, Nycticorax nycticorax, 603
Holbrookia propinqua, 314-316
Holmes, E. Bruce
Variation in the muscles and nerves
of the leg in two genera of
Grouse (Tympanuchus and Ped-
ioecetes), 365
Holocephali, 349
hoopesi, Sturnella magna, 334
howelli, Chordeiles minor, 619
hudsonia, Pica pica, 629
hudsonius, Circus cyaneus, 609
huttoni, Vireo, 284
Hydranassa tricolor, 319
Hydroprogne caspia, 328
Hylaeobatrachus, 164
Hylocichla mustelina, 634
INDEX TO VOLUME 12
Hynobiidae, 163, 175
Hypogeophis, 164
hypugaea, Speotyto cunicularia, 617
Icteria virens, 640
Icterus
bullockii, 644
galbula, 644
spurius, 643
Ictinia
breeding cycle, 511
feeding, 507
habitat, 506
mississippiensis, 505, 607
mortality and defense, 515
ratio immatures to adults, 516
immer, Gavia, 368
imparatus, Corvus, 332
inca, Scardafella, 556, 564, 567, 571,
572
indica, Chalcophaps, 563
inornatus, Catoptrophorus semipalma-
tus, 323
interfusa, Guiraca caerulea, 646
intermedius, Phenacomys, 185, 188,
192, 197, 198, 211
interpres, Arenaria, 325
Ipomoea pescaprae, 312
Iridoprocne bicolor, 556, 625
Iva, 312, 3138
Ixobrychus exilis, 604
jamaicensis,
Buteo, 607
Laterallus, 611
Oxyura, 605
gen Pedioecetes phasianellus, 367-
472
jaw muscles
in doves, 523
in primitive reptiles, 659
Jenkinson, Marion Anne
Thoracic and coracoid arteries in
two families of birds, Columbi-
dae and Hirundinidae, 555
Johnston, Richard F.
Vertebrates from the Barrier Island
of Tamaulipas, México, 311
The breeding birds of Kansas, 577
Kannemeyeria, 669
kansensis, Petrolacosaurus, 219, 497
kaysui, Diplocercides, 482
Kite, Mississippi, in southwestern
Kansas, 505-515
breeding cycle, 511
feeding, 507
habitat, 506
mortality and defense, 515
ratio immatures to adults, 516
Labyrinthodontia, 159, 238, 497
Lagurus, 187
687
Lagurus—Concluded
curtatus, 183, 192, 210, 211
levidensis, 210
Lamna, tooth series, 355, 358
Lanius
excubitorides, 635
ludovicianus, 635
migrans, 635
Larrea divaricata, 28
Larus
argentatus, 40, 326
atricilla, 327
californicus, 342
larva,
Borborocoetes, 176
Leiopelma, 175
modem Amphibia, 174
Lasiurus cinereus, 7
Laterallus jamaicensis, 611
latimeri, Vireo, 256
latrans, Canis, 338
leachii, Glossophaga soricina, 7
leaf-nosed bat, 8
Leiopelma, 166, 171, 172
archeyi, 176
hochstetteri, larva, 175, 177
Lemmi, 212
Lemmus, 212, 213
alascensis, 194
helvolus, 193
lemmus, 194
subarcticus, 194
trimucronatus, 188, 192, 193, 211
lentiginosus, Botaurus, 604
Lepospondyli, 159
Leptonycteris nivalis, 7
Lepus
californicus, 335, 340, 341
curti, 335, 340
merriami, 335, 341
Lespedeza capitata, 248
Leucaena, 3138
Leucophoyx thula, 319, 603
leucopterus, Mimus polyglottos, 332,
633
levidensis, Lagurus curtatus, 210
limicola, Rallus, 611
Limnodromus, 325
Limnoscelis, 661
Limomium carolinianum, 313
Limosa fedoa, 322, 341
lineatus, Buteo, 608
littoralis,
Microtus longicaudus, 202
Neotoma micropus, 338
Taxidea taxus, 338
livia, Columba, 538, 556, 563, 614
longicauda,
Bartramia, 613
Toxostoma rufum, 633
longicaudus,
Chionomys, 191, 202
688
longicaudus—Concluded
Microtus, 187, 191, 202, 211
Phenacomys, 197
Lophodus variabilis, 351
Loxia curvirostra, 648
ludovicianus,
Lanius, 635
Microtus, 187, 192
Pheucticus, 646
Thryothorus, 632
luteus, Colaptes auratus, 620
luzonus, Chaerophon, 7
Lycium carolinianum, 312
Lysorophus tricarinatus, 160
macfarlani, Microtus oeconomus, 205
Maclura pomifera, 247
Macrotus
californicus, 5-7, 38
foraging habits and flight, 32
mexicanus bulleri, 7
roosting habits and terrestrial loco-
motion, 30
marginella, Zenaidura macroura, 615
macroura, Zenaidura, 330, 523, 536,
556, 615
macularia, Actitis, 612
magna, Sturnella, 334, 642
magnificens, Fregata, 318
major, Parus, combat, 262
maximus, Thalasseus, 329
Mazonerpeton, 157
medianus, Dendrocopos pubescens,
622
Megaceryle alcyon, 620
Megalichthys, 236
Megalocephalus brevicomis, occipital
region, 162
megapotamus,
Agelaius phoeniceus, 334
Geomys personatus, 336
mehelyi, Microtus oeconomus, 205
Melanerpes
caurinus, 622
erythrocephalus, 622
melanocephalus, Pheucticus, 646
melanocorys, Calamospiza, 649
melanoleuca, Tringa, 322
Meleagris gallopavo, 610
mellifera, Salvia, 10
Mergulus alle, 367
merriami,
Lepus californicus, 335, 341
Perognathus, 337
Merz, Robert L.
Jaw musculature of the Mourning
and White-winged Doves, 523
Mesocricetus auratus, 186
mexicana, Choeronycteris, 7
mMmexicanus,
Cassidix, 333, 334
UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
mexicanus—C oncluded
Dasypus novemcinctus, 334
Macrotus, 7
Microtus, 187, 192
Micropalama himantopus, 326
micropus, Neotoma, 338
Microsauria, 159
Microti, 212, 213
microtine rodents, baculum, 183, 212
Microtus
abbreviatus, 187
agrestis, 185, 189, 192, 203
alcorni, 206
amosus, 189, 204
arvalis, 185, 189, 192, 202
bacula of, 189-191
breweri, 187
californicus, 205, 212
chrotorrhinus, 192, 211
coronarius, 187, 192
fatioi, 190, 192, 209
finitus, 206
fortis, 185, 189
fulviventer, 192
fusus, 186, 189, 204
gilmorei, 205
gregalis, 190, 192, 200
guatemalensis, 191, 192, 198, 211
gud, 202
guentheri, 185, 189, 192, 203
haydeni, 207
littoralis, 202
longicaudus, 187, 191, 202, 211
ludovicianus, 187, 192
macfarlani, 205
mexicanus, 192, 205, 212
mehelyi, 205
miurus, 190, 192, 212
modestus, 206
mogollonensis, 187, 205
mohavensis, 206
montanus, 192, 186, 204, 211
mordax, 202
nanus, 189, 204
nesophilus, 187
nivalis, 185, 191, 192, 202
ochrogaster, 184, 186, 187, 192,
207
oeconomus, 191, 192, 204, 211
orcadensis, 185, 189, 192, 202
oregoni, 185, 191, 192, 199, 200
parvulus, 185, 189, 208
pennsylvanicus, 187, 189, 192, 206,
211
phaeus, 205
provectus, 187, 192
pullatus, 206
quasiator, 190, 192, 208
richardsoni, 192, 199, 212
roberti, 202
sierrae, 202
socialis, 183
INDEX TO VOLUME 12
Microtus—Concluded
subsimus, 205
subterraneus, 185
taylori, 183, 207
terrestris, 199
townsendii, 185, 191, 204, 211
umbrosus, 187
uligocola, 206
xanthognathus, 192
migrans, Lanius ludovicianus, 635
migratorius, Turdus, 634
Mimomys, 199
Mimus
leucopterus, 332, 633
polyglottos, 633
minor
Chordeiles, 330, 331, 619
Philohela, 612
Miobatrachus, 157, 158
vertebra, 170
mirus, Edestus, 355, 356
mississippiensis, Ictinia, 505, 607
mite, fowl, 287
miurus, Microtus (Stenocranius), 190,
192, 200, 201
Mniotilta varia, 638
modestus, Microtus
206
mohavensis, Microtus californicus, 206
Molge
taeniatus, rib, 168
vulgaris, development of vertebrae,
167
mollis, Crataegus, 248
molossa, Tadarida, 7
Molossidae, 7, 25, 127
Molossus
bondae, 7
coibensis, 7
Obscurus, 7
Molothrus ater, 289, 644
Monanthochloé littoralis, 313
monedula, Corvus, 367
montana, Eupoda, 612
montanus, Microtus, 186, 192
morcomi, Dendroica petechia, 638
mordax, Microtus longicaudus, 202
morinella, Arenaria interpres, 325
morio, Psilorhinus, 342
Mormon arcticus, 367
motacilla, Seiurus, 639
muriei, Microtus miurus, 201
murinus, Thryomanes bewickii, 332
Muscivora forficata, 331, 623
muscles,
jaw, in doves, 523
jaw, in primitive reptiles, 659
leg, in grouse, 396
pectoral girdle and limb, bats, 70
unique to bats, 67
mustelina, Hylocichla, 634
pennsylvanicus,
689
Myiarchus
cinerascens, 331
crinitus boreus, 623
myology of bats, 67
Myopus, 212, 213
Myotis,
brevis, 7
californicus, 26
foraging habits and flight, 26
myotis, 120
roosting habits and terrestrial loco-
motion, 23
velifer, 5, 36, 38
yumanensis, 34
naevius, Otus asio, 617
nanus, Microtus montanus, 189, 204
National Science Foundation, 185,
220, 299, 477, 659
Nectridia, 159
Necturus maculosus, development of
vertebrae, 167
ear, 163, 164, 167
occipital region, 162
rib, 168
neglecta, Sturnella, 642
negundo, Acer, 248
nelsoni, Sitta carolinensis, 631
Neofiber alleni, 191, 192, 209, 211
Neotoma
littoralis, 338
micropus, 338
nerves, grouse, 376
Nesides, 480
newelli,
Coelacanthus, 499
Synaptotylus, 480, 482, 486, 487,
488, 491, 493, 496, 499
newtoni, Edestus, 355, 357
niger, Chlidonias, 327, 614
nigra, Rhynchops, 330
Nitosauridae, 299, 301, 302
nivalis,
Chionomys, 191, 192, 201
Leptonycteris, 7
Microtus, 185, 191
nobilis, Otidiphaps, 564
norvegicus, Rattus, 186
Notobatrachus,
foot, 173
pectoral girdle, 171, 172
noveboracensis, Vireo griseus, 636
novemcinctus, Dasypus, 334
Numenius
americanus, 613
parvus, 322
nuttallii, Phalaenoptilus, 618
Nyctanassa violacea, 603
Nycticorax
hoactli, 603
nycticorax, 319
690
obscurus, Molossus, 7
obsoletus, Salpinctes, 633
occidentalis,
Ardea herodias, 319
Clethrionomys, 185, 188, 192, 195
Geothlypis trichas, 640
Pelecanus, 318
ochrogaster,
Microtus, 183, 184, 190, 192
Pedomys, 190
Odocoileus virginianus, 339
oeconomus, Microtus, 191, 192, 204
Oenothera, 313
olivacea, Piranga, 645
Ondatra zibethicus, 192, 198, 211
Oporornis formosus, 640
Opuntia, 506
lindheimeri, 312
orbiculatus, Symphoricarpos, 248
orcadensis, Microtus, 185, 189, 192
ordii, Dipodomys, 337, 341
oregoni,
Chilotus, 191, 192, 199, 200
Microtus, 185, 191, 192, 199, 200
ornatus, Calcarius, 651
Omithonyseus sylviarum, 287
Orodontidae, 349
Orodus
corrugatus, 350, 351
ramosus, 350
Orthosaurus, occipital region, 162
Orthriomys umbrosus, 187
oryzivorus, Dolichonyx, 642
osteology, cranial, in doves, 531
ostralegus, Haematopus, 320
Otidiphaps, nobilis, 564
Otus
aikeni, 617
asio, 617
hasbroucki, 617
naevius, 617
otus, Asio, 618
Oxyura jamaicensis rubida, 605
Palaeochiropteryx, 128
Palaeoproteus, 164
pallida, Callipepla squamata, 610
pallidicinctus, Tympanuchus, 367, 610
pallidior, Passerina ciris, 647
pallidus, Antrozous, 25, 34
paludis, Synaptomys cooperi, 183,
188, 194
palumbus, Columba, 525
palustris, Telmatodytes, 632
Panicum virgatum, 248
papuana, Paradisea, 367
Paradisea papuana, 367
parasitism of Bell Vireo by cowbird,
289, 291, 292
parasphenoid, 480-483
parkmanii, Troglodytes aedon, 631
UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist.
Parula americana, 638
Parus
atricapillus, 629, 631
bicolor, 631
carolinensis, 631
major, combat of, 262
septentrionalis, 629
parvabullatus, Dipodomys ordii, 337,
341
parvulus, Microtus (Pitymys), 185,
191, 208
parvus, Numenius americanus, 322
Passer domesticus, 641
Passerherbulus henslowii, 650
Passerina
amoena, 647
ciris, 647
cyanea, 647
pallidior, 647
passerina, Spizella, 651
pectoral girdle,
Amphibia, 171
bats, 51
Pedioecetes, 365-472
jamesi, 367-472
phasianellus, 367-472
Pedomys, 212
ochrogaster, 190, 207
pelagica, Chaetura, 41, 619
Pelecanus
erythrorhynchus, 317
occidentalis, 318
pelvic
girdle, bats, 61
limb, bats, 61
Pelycosaurs, two new, from the Lower
Permian of Oklahoma, 299
pennsylvanicus, Microtus, 187, 189,
192-206, 207, 211
peregrinus, Falco, 609
Perognathus merriami, 337
Peromyscus, 291
perotis, Eumops, 5, 6, 7
perpallidus, Ammodramus
rum, 650
personata, Chilonycteris, 7
personatus, Geomys, 811, 355-337
Petalodontidae, 349
petechia, Dendroica, 638
Petrochelidon pyrrhonota, 626
Petrolacosaurus kansensis, 219, 497
phaeus, Microtus mexicanus, 205
Phalacrocorax, 318
auritus, 602
Phalaenoptilus nuttallii, 618
phasianellus, Pedioecetes, 367
Phasianus colchicus, 610
Phenacomys
celatus, 188, 197
intermedius, 185, 188, 192, 197,
198, 211
longicaudus, 197
savanna-
INDEX TO VOLUME 12 691.
Pheucticus
ludovicianus, 646
melanocephalus, 646
philadelphicus, Vireo, 256
Philohela minor, 612
phoebe, Sayornis, 623
phoeniceus, Agelaius, 334, 643
phoenicoptera, Treron, 563, 564
pholeter, Colobomycter, 301
Phyllostomidae, 8, 127
Pica
hudsonica, 629
pica, 629
pileatus, Dryocopus, 620
pinetorum, Microtus (Pitymys), 190,
192, 208
pines, Tympanuchus cupido, 367,
pinus, Spinus, 648
Pipilo
arcticus, 649
erythrophthalmus, 649
Pipistrellus hesperus, 26, 36
Piranga
olivacea, 645
rubra, 645
Pitymys, 212
duodecimcostatus, 209
fatioi, 190, 209
parvulus, 185, 191, 208
pinetorum, 190, 208
quasiator, 190, 208
platensis, Cistothorus, 632
platypterus, Buteo, 608
platyrhynchos, Anas, 604
plecotus auritus, 120
Plegadis chihi, 604
Plesiopoda, 238
Plethodontidae, 163, 175
Pluchea sericea, 28
Pluvialis dominica, 41
Poa pratensis, 248
podiceps, Podilymbus, 602
Podilymbus podiceps, 602
Polioptila caerulea, 635
polyglottos, Mimus, 332, 633
pomifera, Maclura, 247
Populus deltoides, 248, 506
Porzana carolina, 320, 611
pratensis, Poa, 248
praticola, Eremophila alpestris, 625
pratincola, Tyto alba, 617
priscus, Delorhynchus, 299-302
Procyon
excelsus, 338
fuscipes, 338
lotor, 338
Progne subis, 556, 567, 569, 570, 628
Prometheomys, 213
propinqua, Holbrookia, 314-316
prosopidicola, Cassidix mexicanus,
>
Prosopis
juliflora, 312
pubescens, 27
Proteus, ear, 164
Protobatrachus, 157, 158, 159, 164,
165,177
pectoral girdle, 171, 172
Protonotaria citrea, 638
Protorothyris, 660
provectus, Microtus, 187, 192
Psammodontidae, 349
Psilorhinus morio, 342
Pteronotus
davyi, 7
fulvus, 7
pubescens,
Dendrocopos, 622
Prosopsis, 27
pullatus, Microtus pennsylvanicus, 206
pusilla, Spizella, 651
pusillus, Ereunetes, 314
pyrrhonota, Petrochelidon, 626
quasiator, Microtus (Pitymys), 190,
192, 208
Quiscalus
quiscula, 644
versicolor, 644
quiscula, Quiscalus, 644
Rallus
elegans, 611
limicola, 611
na,
middle ear, 163, 164
pectoral girdle, 171
Rattus norvegicus, 186
Raun, Gerald G.
Vertebrates from the Barrier Island
of Tamaulipas, México, 311
Recurvirostra americana, 326, 613
regalis, Buteo, 609
Regulus
calendula, 257
song, 257
relictus, Synaptomys cooperi, 194
reptiles, adductor muscles of jaw, 659
Rhabdoderma, 478, 480, 481, 486,
490, 492, 496, 498, 499
elegans, 477, 480, 483-486, 488,
494, 495, 498
Rhabdodermatinae, 478, 497, 499
Rhea, 367
Rhinolophus hipposideros, 120
Rhynchops nigra, 330
ribs,
Amphibia, 165
bats, 51
richardsoni,
Dichrostonyx rubricatus, 193
Microtus (Arvicola), 192, 199
Ra
692
Richmondena
canicaudus, 334
cardinalis, 645
coccinea, 334
Riparia riparia, 556, 626
roberti, Microtus, 202
Romeria, 660
rothschildi, Fregata magnificens, 318
rubida, Oxyura jamaicensis, 605
rubra, Piranga, 645
rubricatus, Dichrostonyx, 193
rufescens, Dichromanassa, 319
ruficollis, Stelgidopteryx, 556, 626
rufocanus, Clethrionomys, 185, 188,
192
rufum, Toxostoma, 633
rustica, Hirundo, 556, 628
ruticilla, Setophaga, 640
rutilus, Clethrionomys, 188, 192, 194,
195
Salamandra, carpus, 174
Salamandridae, 163, 175
Salicornia, 313
Salix, 35
Salpinctes obsoletus, 633
Salvia
apiana, 10
mellifera, 10
sandvicensis, Thalasseus, 330
saturatus, Synaptomys cooperi, 188,
194
Saurerpeton, 159
savannarum, Ammodramus, 650
saya, Sayomis, 624
Sayornis
phoebe, 624
saya, 624
Scarabaeidae in stomachs of bats, 35
Scardafella inca, 556, 564, 567, 571,
572:
scoparius, Andropogon, 248
Seiurus motacilla, 639
Selander, Robert K.
Vertebrates from the Barrier Island
of Tamaulipas, México, 311
semipalmatus,
Catoptrophorus, 323
Charadrius hiaticula, 321
senegalensis, Streptopelia, 563
sennetti, Dipodomys ordii, 341
sericea, Pluchea, 28
serratidens, Thrausmosaurus, 302-304
serripennis, Stelgidopteryx ruficollis,
626
Setophaga nuticilla, 640
Seymouriamorpha, 160
shark, 497
sharks, Edestid, teeth of, 349-361
Sialia sialis, 635
sialis, Sialia, 635
sierrae, Microtus longicaudus, 202
UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist,
Sigmodon
hispidus, 339, 341
solus, 339, 341
Sitta
carolinensis, 631
nelsoni, 631
skeleton, grouse, 369, 375
socialis, Microtus, 183
solitarius, Vireo, 270, 271
sonorana, Dendroica petechia, 638
soricina, Glossophaga, 7
Spartina, 313
sparverius, Falco, 19, 609
Speotyto
cunicularia, 617
hypugaea, 617
Spermatodus, 489
Spermophilus
annectens, 335
spilosoma, 335
Sphenacodontidae, 302
Sphenodon, 665
Sphingidae, fed on by bats, 35
spilosoma, Spermophilus, 335
Spinus
pinus, 648
tristis, 648
Spiza americana, 648
Spizella
arenacea, 651
arizonae, 651
boreophila, 651
passerina, 651
pusilla, 651
sponsa, Aix, 605
Sporobolus virginicus, 313
spurius, Icterus, 648
spuamata, Callipepla, 610
Squatarola squatarola, 320
Steganopus tricolor, 613
Stelgidopteryx
ruficollis, 556, 626
serripennis, 626
stellaris, Cistothorus platensis, 632
Stenocranius
abbreviatus, 187
gregalis, 190
miurus, 190
Sterna
albifrons, 328
antillarum, 328
athalassos, 329, 614
forsteri, 328, 614
hirundo, 328
Stewart, Peggy Lou
A new order of fishlike Amphibia
from the Pennsylvanian of Kansas,
219
Streptopelia
chinensis, 563, 564
senegalensis, 563
tranquebarica, 563
INDEX TO VOLUME 12
striatus, Accipiter, 607
strigatus, Chondestes grammacus, 650
Strix
georgica, 618
varia, 618
Struthio, 367
Sturnella
argutula, 642
hoopesi, 334
magna, 642
neglecta, 642
Sturnus vulgaris, 636
subarcticus, Lemmus trimucronatus,
94
subis, Progne, 556, 567, 569, 570, 578
subsimus, Microtus mexicanus, 205
subterraneus, Microtus, 185
surinamensis, Chlidonias niger, 327,
614
swainsoni, Buteo, 608
sylviarum, Ornithonyseus, 287
Symphoricarpos orbiculatus, 248
Synaptomys
cooperi, 192, 194, 212
gossii, 194
paludis, 183, 188, 194
relictus, 194
saturatus, 188, 194
Synaptotylus, 482, 490, 492, 495, 498,
499
arcuatus, 499
newelli, 480, 482, 486-488, 491,
493, 496, 499
Tadarida
brasiliensis, 7, 20
molossa, 7
Tamarix, 27
tarsus, in Amphibia, 173
Taxidea
littoralis, 338
taxus, 338
taxus, Taxidea, 338
taylori, Colinus virginianus, 610
teeth
of Edestid sharks, 349-361
of Lamna, 355
symphysial, evolution of, 354
Telmatodytes
dissaéptus, 632
palustris, 632
Temnospondyli, 159, 161, 162
temporal openings, origin of, 674-679
denn ostals, Charadrius alexandrinus,
terrestris, Microtus, 199
teter, Cathartes arva, 605
texanus, Colinus virginianus, 320
Thalasseus
acuflavidus, 330
maximus, 329
sandvicensis, 329
thapsus, Verbascum, 248
693
Thrausmosaurus serratidens, 302-304
Thrinaxodon, 659, 669
jaw muscles, 672
skull, 671
Thryomanes
bewickii, 631, 632
cryptus, 332, 632
murinus, 332
Thryothorus ludovicianus, 632
thula, Leucophoyx, 319, 603
torda, Alca, 367
townsendii,
Corynorhinus, 34
Microtus, 185, 191, 204, 211
Toxostoma
longicauda, 633
rufum, 633
trailli, Empidonax, 624
tranauebarica, Streptopelia, ee
treganzai, Ardea herodias, 31
Treron
bicincta, 563
phoenicoptera, 563, 564
triacanthos, Gleditsia, 247
trichas, Geothlypis, 640
tricolor,
Hydranassa, 319
Steganopus, 613
Trimerorhachoidea, 159
trimucronatus, Lemmus,
193, 21]
Tringa melanoleuca, 322
Triodia flava, 248
tristis, Spinus, 648
Triton, development of vertebrae,
167
Troglodytes,
aedon, 631
parkmanii, 631
tropicalis, Geomys personatus, 311,
336
Turdus migratorius, 634
Tympanuchus, 365-472
attwateri, 367
cupido, 367
pallidicinctus, 367, 610
pinnatus, 367, 609
188, 192,
Typha, 28
Tyto,
alba, 617
pratincola, 617
uligocola, Microtus pennsylvanicus,
Ulmus americana, 248
umbrosus, Microtus (Orthiomys), 187
Uniola paniculata, 313
urens, Eucnide, 35
Urodela,
ear, 163
resemblances to Anura, 177
vertebrae, ribs, 165, 167
694
valisneria, Aythya, 605
varia,
Mniotilta, 638
Strix, 618
variabilis,
Agassizodus, 351
Campodus, 351, 352
Lophodus, 351
Vaughan, Terry A.
Functional morphology of three
bats, Eumops, Myotis, Macro-
tus, 3
velifer, Myotis, 5, 6, 7
velox, Accipiter striatus, 607
Verbascum thapsus, 248
versicolor, Quiscalus quiscula, 644
vertebrae,
Amphibia, 165
bats, 45
Vespertilionidae, 8, 25, 127
villosus, Dendrocopos, 622
violacea, Nyctanassa, 603
virens,
Contopus, 625
Icteria, 640
Vireo
altiloquus, 266
atricapilla, 249, 256, 636
bellii, 243-293, 636
flavifrons, 271, 637
gilvus, 637
griseus, 249, 636
huttoni, 284
latimeri, 256
noveboracensis, 636
olivaceus, 245, 637
philadelphicus, 256
solitarius, 270, 271
virescens,
Butorides, 603
Empidonax, 624
virgatum, Panicum, 248
UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist,
Odocoileus, 339
vocalization, Bell Vireo and other
vireos, 255
vociferus,
Caprimulgus, 618
Charadrius, 612
vulgaris, Sturnus, 636
wardi, Ardea herodias, 319
Wilks, B. J.
Vertebrates from the Barrier Island
of Tamaulipas, México, 311
Wilsonia citrina, 640
wilsonia, Charadrius, 321
wilsonianus, Asio otus, 618
Wimania, 482
wing, bats, 51
Xanthocephalus xanthocephalus, 642
xanthognathus, Microtus, 192
yumanensis, Myotis, 34
zebra, Centurus carolinus, 620
Zenaida
asiatica, 528, 556
aurita, 523, 536
cranial osteology, 531-533
generic relationship, 535-538
myology, 524-531
Zenaidura
auriculata, 523, 536
carolinensis, 615
cranial osteology, 531-533
generic relationship, 535-538
macroura, 830, 528, 536, 556
marginella, 615
myology, 524-531
yucatanensis, 536
zibethicus, Ondatra, 192, 198, 211
30-3964
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