Jaw Musculature of the Mourning and White-Winged Doves is a public-domain classic of science by Robert L. Merz.
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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
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
29-7865
Jaw Musculature Of the Mourning and White-winged Doves
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 investigations 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 13 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. Specimens used were: White-winged Doves, KU 19141, 19142, 19143, 19144, 19145, 19146, 19147, 23138, 23139, 24337, 24339, 24341, 23592, 23593, 24340, 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 auriculata), 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 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: 377; 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 musculature 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. 3, 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 3.
~M. pterygoideus dorsalis:~ part of Mm. pterygoidei, Gadow, 1891:323-325, 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, Edgeworth, 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 superficialis, second part, Adams, 1919:100-101.
~M. pseudotemporalis profundus:~ M. quadrato-maxillaris, Gadow, 1891:322-323--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 3.
~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. 3 and 11--part of M. spheno-pterygo-quadratus, Edgeworth, 1935:57.
~M. depressor mandibulae:~ M. digastricus s. depressor mandibulae, Gadow, 1891:318-319--M. biventer maxillae, Shufeldt, 1890:18-19, figs. 3, 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, Edgeworth, 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 3.
~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. 3 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 equivalents 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 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 ventralis 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).
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