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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

Jaw Musculature Of the Mourning and White-winged Doves

ROBERT L. MERZ

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 : White-winged Doves , 40323, 40324, 40328, 40392, 40393; Zenaida Doves , 40399, 40400; Mourning Doves , 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 , 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 , 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 , on the other hand, involves extremely powerful gripping ; 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.

ACTION OF JAW MUSCLES

There is no lifting action involved in contraction of the depressor muscle in doves for two reasons-- the origin of the muscle is situated much too far posteriorly on the skull, and, more important, the quadrate is not hinged for vertical movement. As will be discussed later, it moves only in a horizontal plane.

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" . 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 anterodorsal direction, and the force is transferred to the upper mandible, which is thereby elevated. A synergetic mechanism is simultaneously 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 and the movement of the quadrate is not so much up and down, or vertical, as it is horizontal . 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 .

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 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 operation 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 relaxed 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 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 .

Ten measurements were taken on each skull, but simple observation 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.

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

The jaw musculature has been known for some time to be highly adaptive 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 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 while investigating pollinating agents of the Saguaro Cactus , and by Gilman 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 noted that both Mourning and White-winged doves were "exceptionally fond of 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 appearance 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.

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 unquestionably 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 relationship of these genera. The original dissections which are discussed in this report are only valuable as one more bit of evidence concerning one characteristic that aids in clarification of generic relationship, and it is only in conjunction with other evidence that any satisfactory conclusion may be forthcoming.

Morphology

"Plumage of head, neck and under parts soft and blended; bare orbital space moderate, broadest beneath eyes. Coloration plain, the proximal secondaries spotted with black; rectrices 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."

Nidification

It has been adequately documented that members of these genera closely resemble one another in their nesting and egg-laying habits. Bent , Davie , Goss and Nice 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 identical habits , and a similar situation exists with the Zenaida Dove .

Like the other species, White-winged Doves lay two white or buffy eggs per clutch and build frail nests of sticks .

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 genotype of the common ancestor.

Interbreeding

Serology

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

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