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Double-Crested Cormorant ADAPTATIONS FOR LOCOMOTION AND FEEDING IN THE ANHINGA AND THE DOUBLE-CRESTED CORMORANT BY OSCAR T. OWRE ORNITHOLOGICAL MONOGRAPHS NO. 6 PUBLISHED BY THE AMERICAN ORNITHOLOGISTS' UI•ION ORNITHOLOGICAL MONOGRAPHS This series,published by the AmericanOrnithologists' Union, hasbeen establishedfor majorpapers too long for inclusionin the Union'sjournal, The Auk. Publicationhas been made possible through the generosityof Mrs. Carll Tucker and the Marcia Brady Tucker Foundation,Inc. Correspondenceconcerning manuscripts for publication in the series should be addressed to the Editor, Dr. Robert W. Storer, Museum of Zool- ogy,The Universityof Michigan,Ann Arbor, Michigan48104. Copiesof OrnithologicalMonographs may be orderedfrom the Treasurer of the AOU, Burt L. Monroe,Sr., Box 23447,Anchorage, Kentucky 40223. (Seeprice list on insideback cover.) OrnithologicalMonographs, No. 6, 138 pp. Associate Editor: Richard L. Zusi AssistantEditors: Joseph R. Jehl,Jr., and JeanneD. Jehl Issued October 5, 1967 Price$3.50 prepaid ($2.80to AOU members) Library of CongressCatalogue Card Number 67-30457 Printedby The Allen PressInc., Lawrence,Kansas 66044 ADAPTATIONS FOR LOCOMOTION AND FEEDING IN THE ANHINGA AND THE DOUBLE-CRESTED CORMORANT BY OSCAR T. OWRE ORNITHOLOGICAL MONOGRAPHS NO. 6 PUBLISHED BY THE AMERICAN ORNITHOLOGISTS' UNION CONTENTS Page ACKNOWLEIXgMENTS .............................................................................................................. 4 INTRODUCTION ........................................................................................................................ Materials and methods ............................................................................................... 6 AERODYNAMICS AND THE WING ....................................................................................................... 7 Flight characteristics.................................................................................................... 7 Weight and wing loading ...................................................................................................7 Main elementsof the wing skeleton.................................................................................. 9 The carpometacarpusand the digits ............................................................... 10 The pectoral girdle .................................................................................................. 11 Pneumaticity ............................................................................................................... 13 Myology of the wing ..........................................................................................................13 Discussion ........................................................................................................................... 38 Conclusions ........................................................................................................................ 46 THE TAIL ........................................................................................................................................ 48 Size, functions, and molt ................................................................................................ 48 Osteologyof the tail ..............................................................................................................51 Muscles of the tail ........................................................................................................... 54 Summaryand conclusions............................................................................................... 59 THE LEG ............................................................................................................................................ 60 Swimming ...................................................................................................................... 60 Osteologyof the pelvis and leg ......................................................................................63 The pelvis ......................................................................................................................63 The femur ..................................................................................................................... 65 The tibiotarsus and fibula .......................................................................................... 65 The patella ....................................................................................................................66 The tarsometatarsus .................................................................................................... 68 Metatarsal I ............................................................................................................. 69 The digits ..................................................................................................................69 Myology of the leg ..........................................................................................................72 Discussionand summary ......................................................................................................99 Conclusions ............................................................................................................................. 104 THE HEAD ........................................................................................................................................ 106 Method of feeding ...............................................................................................................106 The skull .................................................................................................................................. 107 Certain musclesof the skull and jaws .............................................................................110 Discussion ....................................................................................................................... 121 The musclesof the neck attaching to the skull ...................................................121 Jaw action ...................................................................................................................123 Adduction of the lower jaw ...................................................................................123 Abduction of the lower jaw ......................................................................................126 Action of the upper jaw: abduction .........................................................................126 Action of the upper jaw: adduction ......................................................................126 Phylogeneticimplications of comparisonsof the skull, jaws, and associated muscles ................................................................................ 127 Food ............................................................................................................................................... 129 Summary ...................................................................................................................................133 CONCLVSIONS: TH• ANHINGA AND TH• PHALACROCORAClnA• ............................................... 134 LIT•RATVP. Z CIX•D ........................................................................................................................ 136 ACKNOWLEDGMENTS Inspirationfor this investigationwas derived from coursework and con- versationwith the late Dr. JosselynVan Tyne and Dr. Robert W. Storer. Dr. Van Tyne directedthe studyinto its middlephases. I am greatlyin- debted to Dr. Storer for his counseland direction during the remaining portionof my work. I wishto thankDr. AndrewJ. Berger,then of the Departmentof Anatomy of The Universityof Michigan,for his helpful criticismsof the sectiondeal- ing with the royologyof the wing. Dr. Richard Robins and Mr. Luis R. Rivas of the University of Miami confirmedidentification of certain fish. I am obliged to Mr. Dennis Paulsonfor the collectionof an Anhinga and a cormorant. Mr. Arthur P. Kirk of Goodland, Florida, provided much usefulinformation, gained from his long experiencein the mangroveareas of the State. I am grateful to The University of Michigan and to the University of Miami for facilities extended to me. The Florida Game and Fresh Water Fish Commissiongranted me special permissionto collect in certain re- strictedareas, and Mr. Daniel S. Beard as Superintendentof the Everglades National Park offered me privilegeswhich facilitated my observations. Without the inspirationand assistanceof Harding B. Owre this investiga- tion might neverhave been brought to completion. INTRODUCTION Few contributionsto our knowledgeof the ecologyof the Anhinga, ,4nhingaanhinga leucogaster (Vieillot), havebeen made sincethe observa- tions of Audubon (1838: 136-160). The most importantof theseis the work of Meanley (1954). Current summarizationsof life historymaterial are drawn largely from Audubon'sobservations. Certain aspectsof the anatomyof Anhinga receivedconsiderable investi- gation in the yearspreceding 1900 and, to someextent, shortlyafter that. The osteologyand myologyof the neckwere dealt with by Garrod (1876a: 334-340), Forbes (1882: 210-212), Beddard (1898: 413-415), Virchow (1917), and Boas (1929). Generalreviews of the osteology'were made by Mivart (1879) and Shufeldt (1902)- MorphologY of the alimentarY tract was describedin Audubon (1838), by Forbes (1882: 208-210), Cazin (1884), Garrod (1876a:341-345; 1878:679-681), Beddard (1892: 292- 295), and
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