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Myology of the Limpkin

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Myology of the Limpkin MYOLOGY OF THE LIMPKIN By TED T. ALLEN > A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY > UNIVERSITY OF FLORIDA June, 1962 ACKNOWLEDGMENTS I am sincerely and deeply grateful for the generous help and skillful guidance of ray chairman. Dr. Pierce Brod- korb, and for the use of his reference materials and skeleton collection. I also thank the other members of my committee, Drs. E. C. Bovee, E. S. Ford, J. N. Layne, and J. R. Redmond, for their help and constructive suggestions. Grateful acknowledgment is also due the Florida State Game and Freshwater Fish Commission for the issuance of a special permit for collection of a small number of limpkins. Dr. 0. L. Austin, Jr., Mr. Anthony Austin, and Mr. Earl May contributed needed specimens. Dr. Glen E. Woolfenden and Mr. Robert McFarlane contributed specimens and also, along with Dr. Dale E. Birkenholz, gave various other forms of valuable assistance. I am indeed grateful for their help. One summer of work was completely financed by a Fellow- ship for Teaching Assistants from the National Science Foundation, and partial coverage for another summer was pro- vided by a fellowship from the College of Arts and Sciences of the University of Florida. The monetary assistance was invaluable, since it greatly reduced the time required to complete the dissertation. ii TABLE OF CONTENTS Page ACKNOWLEDGMENTS ii LIST OP ILLUSTRATIONS iv INTRODUCTION 1 DESCRIPTIVE MYOLOGY 4 Muscles of the Skull 5 Muscles of the Hyoid 54 Muscles of the Orbit 88 Muscles of the Wing 98 Muscles of the Tail 197 Muscles of the Leg 217 DISCUSSION 308 CONCLUSIONS 323 SUMMARY 327 LITERATURE CITED 329 BIOGRAPHICAL SKETCH 339 iii LIST OF ILLUSTRATIONS Figure Page 1. Lateral and posterior views of the skull of the limpkin, showing attachments of the muscles in the occipital region 333 2. Lateral views of the skulls of certain birds, showing the position of the cranial attachment (anterior origin) of M. Dermo-temporal is .... 334 3. Lateral view of the skull of the limpkin, showing attachments of the muscles of the jaws and of the orbit 335 4. Ventral view of the skull of the limpkin, show- ing attachments of muscles 33$ 5. Medial view of the left eyeball of the limpkin, showing the muscles of the eye and their attach- ments 337 6. Palmar and anconal views of the left humerus, showing attachments of certain muscles of the wing 333 iv INTRODUCTION The limpkin, Aramus guarauna (Linnaeus), is the sole living member of the avian family Aramidae of the order Gruiformes. It occurs in southeastern Georgia, Florida, the Greater Antilles, and the Neotropical mainland from Mexico to Argentina (American Ornithologists' Union, 1957). The spe- cies is known from a pre-Columbian site in Venezuela (Wet- more, 1935) and from four localities in the Upper Pleisto- cene of Florida (Wetmore, 1931; Brodkorb, 1956; Woolfenden, 1959) . Other members of the family occur in Tertiary depos- its of the Great Plains, Aramus sp. from the Lower Pliocene of Nebraska (Wetmore, 1928), Aramornis longurio Wetmore (1926b) from the Middle Miocene of Nebraska, Gnotornis aramiellus Wetmore (1942) from the Upper Oligocene of South Dakota, and Badistornis aramus wetmore (1940) from the Mid- dle Oligocene of South Dakota. The living limpkin is adapted to a rather unique diet. This consists almost exclusively of large snails of the genus Pomacea , formerly known as Ampullaria , with occasional other gastropod and pelecypod mollusks (Wetmore, 1926a; Cot- tam, 1936) . The bird usually takes the snail to a regular feeding station and with its long, specialized bill pecks through the operculum, extracts, and swallows the soft body of the mollusk. 1 2 The bird occurs primarily in grassy marshes and wooded swamps (Bent, 1926; Wetmore, 1926a? Nicholson, 1928) . Where bushes or trees are available it perches and nests in them, often at some height from the ground. Both the feeding and perching habits of the limpkin are thus quite distinctive and differ from the usual condition of its ordinal allies, the rails and cranes. Adaptations to a specialized way of life would be expected to obscure re- semblances to its nearest relatives and may account for some of the disagreement over the systematic position of the Aramidae. The subordinal position of the limpkin has not been es- tablished with certainty, although all workers have referred it to one of two suborders, the Grues or the Ralli. Some of the difficulties involved may be seen in a brief review of the opinions of other workers on the systematic position of the Aramidae. Audubon (1839) thought Aramus to be allied to the rails on the basis of the viscera. Nitzsch (1867) found that the pterylosis resembles that of Grus and Psophia and no other bird; nevertheless, he placed it with the rails be- cause of peculiarities of the feathers themselves. Garrod (1876) considered Aramus as being closer to Grus , on the ba- sis of several anatomical features, especially of the skull, sternum, and some aspects of the thigh musculature. Shufeldt (1894) placed it with the cranes but considered the species a perfect link between cranes and typical rails. Clay (1950) found the aramid Mallophaga to be rail-like and strongly 3 different from the type found in cranes. On the basis of the electrophoretic profiles of egg-white proteins, Sibley (1960) thought Aramus to be closer to the rails, and he men- tioned the large hallux, feathered head, and color of the downy young as also being rail-like. From a broader view, these opinions indicate that the Aramidae are definitely related to both the Gruidae and the Rallidae. A major purpose of this paper is to investigate affinities of the three families through study of their com- parative myology. Fisher (1955) remarked that detailed ana- tomical work should solve the question of affinities. DESCRIPTIVE MYOLOGY During the course of this study, six limpkins were available for dissection. Two were dissected in detail and critical points were checked in the others. To represent both wading and swimming types of Rallidae, two specimens of the clapper rail, Rallus longirostris Boddaert, and two of the coot, Fulica americana Gmelin, were dissected. Some features were also examined on a specimen of the purple gal- linule, Porphyrula martinica (Linnaeus) . A specimen of the crowned crane, Balearica pavonina (Linnaeus) , was dissected as representing a generalized gruid, and comparisons were made with Fisher and Goodman's (1955) description of the more specialized whooping crane, Grus americana (Linnaeus), and Berger's (1956a) study of the appendicular myology of the sandhill crane, Grus canadensis (Linnaeus) . All dissections follow the basic plan of the excellent work by Fisher and Goodman (1955), whose descriptions are quite adequate as a guide to dissection of the species used in this study. With very few exceptions, each muscle discussed by those authors is found in each of the other species, and only a few are present that are not discussed by them. In the descriptions of individual muscles comparisons are made on the basis of their proportions, as if all spe- cies studied were of the same size. Attention is called to 4 5 differences In general configuration, points of attachment, and function, at the specific, generic, or family levels, in an effort to determine the relationship of the limpkin to the cranes and rails. Muscles of the Skull 1. M. dermo-temporalis Elongate and sheet-like dermo-osseous muscle. Covers lateral surface of full length of neck. Attaches to cranium, to furculum, to M. tracheohvoideus , and to skin of neck. Lies just beneath skin and deep to lateral edge of sheet of M. constrictor colli in anterior half of neck. Lat- eral to dermal fasciculi of Mm . intertransversarii . Anterior end superficial to posterodorsal portion of M. depressor mandibulae , ventrolateral edge of cephalic part of M. cucul- laris , anterodorsal portion of M. rectus capitis lateralis , side of fourth and fifth fasciculi of M. rectus capitis superior , and dorsal tip of hyoid cartilage. Origin . In two parts, both fleshy, at opposite ends of muscle. Anterior origin from posterolateral edge of skull bordering posterior edge of temporal fossa and extending to base of opisthotic process, in a narrow line 8 ram long, just posteroventral to origins of deep slip of M. adductor mandibulae externa s superficialis and rostral slip of M. ad- ductor mandibulae externus profundus . Anterior origin sepa- rates origins of latter two muscles from both anterior edge of origin of M. depressor mandibulae and anteroventral edge of origin of M. cucullaris , cephalic part. s s 6 Posterior origin is in common with M. tra cheohyo ideu as result of fusion of the two muscles. Common sheet origi- nates from narrow line along anterolateral edge of ventral three- fourths of furculum, extending to mid-ventral line. From this origin the sheet runs anterodorsad onto the neck. Insertion . By connective tissue, onto skin of neck, by all but anterior 10-18 mm of length of muscle. Fuses medi- ally with M. tra cheohyo ideu at ninth or tenth cervical ver- tebra to form common sheet. Posterior to fusion the common sheet covers neck, except in narrow area along middorsal and mid-ventral lines. A few short bundles, arising from furcu- lum, leave dorsomedial edge of muscle near its posterior end and insert on skin of neck near middorsal line. Action . Tenses skin of neck. Comparisons . In the Gruidae the muscle undergoes change. In Balearica it is similar to the condition just described in Aramus, but the belly is much thinner. Grus is apparently unique in lacking the furcular origin and in having a very short belly.
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