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The Appendicular Myology of the Labrador Duck (Camptorhynchus Labradorius)

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The Appendicular Myology of the Labrador Duck (Camptorhynchus Labradorius) Condor, 80:407-418 0 The Cooper Ornithological Society 1978 THE APPENDICULAR MYOLOGY OF THE LABRADOR DUCK (CAMPTORHYNCHUS LABRADORIUS) RICHARD L. ZUSI AND GREGORY DEAN BENTZ The Labrador Duck (Camptorhynchus labra- on either humeral element. The remainder of dorius, Gmelin) has been extinct for almost the forelimb bones and muscles were in sur- 100 years, the last specimen having been prisingly good condition. As all the feathers taken in 1878. The species occurred formerly were retained with the skin, M. expansor along the east coast of North America from secundariorum could not be demonstrated. the Ungava Peninsula to the Delaware River Each hindlimb included the distal half of the and perhaps to Chesapeake Bay (Palmer tibiotarsus, the tarsometatarsus, and all pha- 1976). Little is known about its habits or in- langes of all toes except the ungual pha- ternal anatomy, and nothing is known about langes, which remained with the skin. In some its myology. instances complete bellies of muscles were The purpose of this paper is to describe absent and only the tendon of insertion is de- the distal musculature of the wing and leg of scribed. Also, due to the extremely dehy- the Labrador Duck. The unique history of drated condition of all muscles, the presence a study skin of an adult male in the collections of several small muscles and fiber architecture of the Smithsonian Institution (USNM 1972) even of larger ones could not always be dem- gave us the opportunity to do this. About onstrated. Dissection was performed with a twenty years ago, Humphrey and Butsch stereomicroscope and drawings were made (1958) realized that they could study the with the aid of a camera lucida microscope pterylosis and osteology of the Labrador Duck attachment. We referred to one fluid speci- by temporarily dismantling a study skin of men each of Oldsquaw (Clangula hyemalis) the species. On removing the limb bones, and White-winged Scoter (Melanitta de- they also found that the distal musculature glandi) to clarify our interpretation of the dried was “well enough preserved to warrant de- muscles of Camptorhynchus and some points scription,” The limb bones and muscles re- were also checked against the Common Eider moved from that specimen are the basis for this (Somateria mollissima). Bentz wrote the ini- study. Surprisingly, no broad comparative tial descriptions and Zusi made the illustra- survey of hindlimb myology of waterfowl has tions. The entire paper was reviewed and yet been published. Studies on a few species modified by both authors. One leg and one are those of Stolpe ( 1932), Miller ( 1937), wing were skeletonized; their counterparts Frantigek ( 1934) and Ruck ( 1949), and more are retained in alcohol. recently, Raikow ( 1970) analyzed diving We follow the myological nomenclature of adaptations in the stifftail ducks. Even fewer George and Berger (1966). Where their studies of forelimb myology are available. The terminology differs from that being developed most complete work is still that of Fiirbringer by the International Committee on Avian (1888), who dealt mainly with the shoulder Anatomical Nomenclature, we have added the and proximal portion of the wing. Brief de- new name in parentheses. The new terms have scriptions of wing muscles in a few species already been employed in certain studies and a detailed analysis of their functions in (Raikow 1976, 1977). We use the terms relation to flight were presented by Sy ( 1936). “major digit” and “minor digit” for wing Lacking broadly comparative information on digits II and III, and “major metacarpal” anatid myology we made no systematic com- and “minor metacarpal” for metacarpals II and parison of the Labrador Duck with other III, respectively. The term “alular phalanx” species; by contrast our comparison of the refers to the pollex. The directional terms cleaned limb bones with skeletons in the “dorsal” and “ventral,” “cranial” and “caudal” Smithsonian Institution and with informa- as applied to the forelimb, refer to the hori- tion presented by Woolfenden (1961) pro- zontally extended wing. vided some taxonomic clues that conclude this paper. MUSCLES OF THE FORELIMB Only the distal half of each humerus was M. biceps brachii (Not shown). This muscle present and only the distal muscles remained inserts by two parallel tendons that pass dor- [4071 408 RICHARD L. ZUSI A?rTD G. D. BENTZ OS ulnare FIGURE 1. Ventral views of left forearm of Camntorhynchus.- ,~ (A and B) Superficial muscles; M. pronator superficialis removed in A. (C) Deep muscles. sal to M. brachialis. One inserts on the proxi- muscle arises by a strong tendon from the mocranial surface of the ulna. The other in- medial surface of the distal end of the humerus serts on the proximoventral surface of the j-ust deep and slightly proximal to the origin radius and sends a branch to the distal end of the humerocarpal band (h c). The fan- of the first tendon and the adjacent portion shaped belly passes distally to a fleshy in- of the ulna. sertion on the ventral surface of the proximal M. brachialis (Fig. 1C; b). This muscle has two-thirds of the radius. a fleshy origin on the distal end of the humerus Ad. flexor digitorum superficialis (Figs. lA, just proximal to the condyles. The fibers of 3A, 4C; f d s). This muscle arises by a short the belly pass distally along the cranial border tendon from the medial condylc of the hu- of the ligamenturn collaterale ventrale (1 c v) merus and is completely cnsheathed by the and insert fleshy on the ventral surface of the humerocarpal band. The spindle-shaped belly proximal end of the ulna. Its insertion is deep passes distally and ends on a narrow tendon to the belly of M. entepicondyloulnaris. that passes into the hand in a shallow groove M. pronator superficiaks (Fig. 1B; p s). on the cranial surface of the OS ulnare. The This muscle arises by a stout tendon from a tendon attaches to the proximal end of the shallow depression near the medial epicondyle first phalanx of the major digit, encloses a of the humerus. The belly fans out to insert sesamoid, and continues distally to insert on by semitendinous fibers on the cranioventral the cranial surface of the proximal end of the surface of at least the proximal half of the second phalanx of the major digit, ventral to radius. the insertion of M. extensor indicis longus. M. pronator profundus (Fig. 1A; p p). This M. flexor digitorum profundus (Figs. lC, MYOLOGY OF THE LABRADOR DUCK 409 ua FIGURE 2. Dorsal views of left forearm of Campto~hynclaus. (A) Superficial muscles. ( B ) M. extensor metacarpi ulnaris removed from its common tendon of origin with M. anconeus. (C) Deep muscles. 3A, 4C; f d p). This large muscle lies super- muscle has a common tendinous origin with ficial to M. ulnimetacarpalis ventralis and M. pronator profundus from the medial epi- arises by two fleshy heads from the cranio- condyle of the humerus. The belly fans out ventral surface of the ulna. The belly extends caudodistally and inserts fleshy on the ven- most of the length of the forearm and ends tral surface of the proximal one-fourth of the on a strong tendon that passes craniad to the ulna, ventral and caudal to the insertion of OS ulnare, around the pisiform process of the M. brachialis. George and Berger (1966: carpometacarpus, and into the hand. The 354) referred to this muscle as the “gallina- tendon passes along the cranioventral edge ceous” muscle, stating that it was known only of the carpometacarpus where it gradually in kiwis ( Apteryx), the Galliformes and tina- crosses the tendon of M. flexor digitorum mous. It is clearly present in Camptorhynchus superficialis caudally. At the distal end of and we found it also in the other ducks we ex- the first phalanx of the major digit the tendon amined. curves abruptly cranially and ventrally to M. flexor carpi ulnaris ( Fig. 1A; f c u) . This pass through a fibrous loop (not shown) and is the most caudal muscle on the ventral side an adjacent bony loop (not shown) on the of the wing. It arises by a strong tendon from proximocranial edge of the second phalanx. the distal end of the medial condyle of the From this loop the tendon continues along humerus, passes caudal and lateral to a large the cranioventral edge of the second phalanx sesamoid and gives rise to a strong, tendinous to insert on the proximoventral edge of the belly. The sesamoid is anchored by ligaments tiny third phalanx. to the medial condyle of the humerus and M. entepicondyZouZnu& (Fig. 1C; e). This to the olccranon process of the ulna. The 410 RICHARD L. ZUSI AND G. D. BENTZ A B FIGURE 3. Muscles of the wrist and manus of the left wing of Camptorhynchus. (A) Ventral view. (B) Dorsal view. Tendon of M. extensor indicis longus crosshatched for clarity. tendon of origin extends distally through the of insertion. This tendon encloses the patella center of the circumpennate belly, sending ulnaris just before it inserts on the coronoid fibers to the superficial ensheathing aponeuro- process of the ulna. A thin sheet of tendinous sis that becomes the tendon of insertion. The fibers connects the two tendons of insertion superficial aponeurosis is developed most and attaches on the ulna between them. strongly along the cranial edge of the belly. M. extensor metacarpi radialis (Figs. 2A, The 45 mm-long belly extends three-fourths 3A, B; e m r). This large muscle consists of the length of the forearm and ends on a strong two parts. The main part arises by both fleshy tendon that inserts on the proximal surface of and tendinous fibers from the lateral aspect the OSulnare.
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