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First published online August 18, 2005
Journal of Experimental Biology 208, 3409-3419 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01768

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Cranial kinesis in palaeognathous birds

Sander W. S. Gussekloo^* and Ron G. Bout

Institute of Biology Leiden, Evolutionary Morphology, Leiden University, Kaiserstraat 63, NL-2311 GP Leiden, The Netherlands

View larger version (15K): [in a new window]   Fig. 1. Types of cranial kinesis in birds. (A) prokinesis, (B) distal rhynchokinesis and (C) central rhynchokinesis. Skull outlines are given in lateral view. Black triangles indicate the main area of rotation, multiple triangles in a single element indicate the boundaries of an bending zone. The triangle in A indicates the nasal–frontal articulation. Arrows indicate the lateral (nasal) bar of the upper bill, which is incomplete in C. Ligament is shown in grey. (Adapted from Zusi, 1984.)

View larger version (21K): [in a new window]   Fig. 2. Ventral view of the skull of a rhea. A, vomer; B, palatinum; C, rostrum parasphenoidale; D, processus basipterygoideus; E, pterygoid; F, quadrate.

View larger version (16K): [in a new window]   Fig. 3. Experimental set-up in lateral view. A, fixation rods; B, step motor; C, force transducer; D, experimental object.

View larger version (26K): [in a new window]   Fig. 4. Schematic representation of the skull of three species of Paleognathae. (A) Ostrich Struthio camelus, (B) emu Dromaius novaehollandiae, (C) rhea Rhea americana. (D) Model of the rhea drawn into a outline of the actual skull, indicating the relationship between the model and the actual skull. Arrows in A–C indicate the line of action and the approximate length of the muscles: 1, musculus depressor mandibulae; 2, musculus pterygoideus; 3, musculus adductor mandibulae externus; 4, adductor muscle complex of the quadrate; 5, musculus protractor pterygoidei et quadrati. Gray areas represent the quadrate and the lower bill. Circles represent rotation points. Scale bar, 2 cm.

View larger version (16K): [in a new window]   Fig. 5. Bending zones in the upper bill elements of the purple sandpiper Calidris maratima. Relative thickness of the dorsal bar (A) and ventral bar (B) vs the distance from the bill tip. Lines are Lowess fits indicating average relative thickness. In this and subsequent figures, vertical broken lines where shown indicate morphological characters of the bill: (a) caudal border of the rostrum maxillae, (b) rostral point of the rostrum basisphenoideus, (c) position of the lateral bar (b and c not present in this species). Arrows indicate the position of the bending zones as observed during experimental manipulation of the upper bill or from kinematical analysis.

View larger version (13K): [in a new window]   Fig. 6. Bending zones in the upper bill elements of the sanderling Calidris alba. See Fig. 5 for details (b and c not present in this species).

View larger version (17K): [in a new window]   Fig. 7. Bending zones in the upper bill elements of the rhea Rhea americana. See Fig. 5 for details.

View larger version (16K): [in a new window]   Fig. 8. Bending zones in the upper bill elements of the ostrich Struthio camelus. See Fig. 5 for details.

View larger version (16K): [in a new window]   Fig. 9. Bending zones in the upper bill elements of the emu Dromaius novaehollandiae. See Fig. 5 for details.

View larger version (19K): [in a new window]   Fig. 10. Reaction force in the upper bill vs the elevation angle of the upper bill. Forces were measured in the linear phase and are means ± S.D.

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