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THE APPENDICULAR WITH COMPARATIVE MYOLOGY OF THE SANDHILL CRANE, REMARKS ON THE WHOOPING CRANE BY ANDREW U J. BERGER recently, very little had been published on the myology NTIL cranes. Fisher and Goodman (1955) of the described in detail the myology of the Whooping Crane (Grus americana) ; they also dissected one Little Brown Crane (G. c. canadensis). I began a myological study of the Sandhill Crane (G. canadensis tabida) at the suggestion of Dr. L. H. Walkinshaw, whose interest in the biology and taxonomy of the cranes is well known. For the first specimen of this subspecies, I am indebted to Dr. Wallace Grange of Babcock, Wisconsin. After the death of this captive bird, it was frozen immediately; I dissected it during the month of April, 1955. During February, 1956, two additional frozen specimens became available. These birds were killed by hunt- ers during the latter part of October, 1955, in Jasper County, Indiana. these specimens, I am indebted to Dr. Charles Kirkpatrick For of Purdue Uni- versity and to Russell Mumford of the University of Michigan. Through the generosity of Dr. Fisher, I was permitted to study the Whooping Crane manuscript before I began my first dissection. After I had completed this work, Dr. Fisher and I discussed differences in interpretation of certain muscle complexes. Th ese differences will be explained in the descriptions of the individual muscles, inasmuch as it was too late to make changes in the Whooping Crane manuscript. There are two sets of muscle terminology currently in use in this country, that of Hudson (1937) and Hudson and Lanzillotti (1955) and that of Fisher (1946) and Fisher and Goodman (1955) ; I have included both sets of names. The muscles are discussed in the sequence used by Fisher and Goodman; they accepted Montagna’s (1945) conclusions on the numbering of the hand digits, and, consequently, proposed new names for certain muscles (1955 :39). MYOLOGY M. TENSOR Fisher PATAGII and from the dorsal state (p. 42) “comes 1955: Fig. one, to my knowledge, are separate origin” surface 17). (e.g., Fiirbringer biceps and German the cuckoos), only distally. and Gadow . . . .” either In ornithologists, believed arising brevis and is that which of the traditional I believe that longus it is (Fisher that both the tensor patagii of M. deltoideus is a derivative major; no of 111. biceps brachii. In the two tensors have a common origin In some birds, the two muscles are separate 282 belly” longus view as M. tensor patagii brevis muscles were derivatives has suggested that triangular of the tensor patagii of M. by the British “elongately solely to M. tensor patagii to consider the slip from M. biceps brachii and Goodman, birds the as belonging only muscular the antero-palmar longus and the tensor patagii many hlgus) interpret end of the furculum of this complex misleading (prOpatagi& (1955 :43) that “the from treatment LONGUS Goodman WING OF THE and the bellies throughout. Andrew .I. Rerger MYOLOGY Thus, there interpret patagii OF SANDHILL are two possible interpretations the belly arising from the only muscular origin an academic such differences of interpretation eous taxonomic What question, bellies and Goodman longus is that derived but it is important I prefer of Mm. to tensores patagialis”) . If one does 1901: 641, “deltoides in myological 283 in the cranes. to be the fused then it is true, as Fisher for the tensor patagii This is, in part, of this complex furculum longus et brevis (see also Mitchell, not agree with this interpretation, CRANE stated, that the from M. biceps brachii. that the taxonomist recognize studies so that these are not given erron- significance. Fisher and Goodman call the belly of M. tensor patagii longus is actually the biceps slip, a muscular slip widely used in taxonomic diagnoses. The biceps slip of Beddard is the biceps propatagialis of Gadow and Selenka (1891:255) and the tensor accessorius of Parker In the Sandhill and Haswell Crane, (1947 :441), the origin that described for the Whoopin g Crane (6-7 cm. long) arises from the coracoidal and inserts both the tensor propatagialis, to the deltoid patagii (2 cm.) envelope neurosis fuses with bases of the alula in the W’hooping from aponeurosis, quills tendon of M. A second origin brevis muscles. of M. TENSOR PATAGII coverts. belly extends to the middle of the furculum, longus is on the extensor Fisher tensor and Goodman alae digiti II longus. This patagii (= extending (1955:68) abductor The main band M. tendon as described extensor the humerus, (1 cm. wide) the forearm, nare. Fisher which metacarpi is not true in the Sandhill to the elbow; adjacent sending the anterior carpi ulnaris.” toward and then passes proximad to attach to the distal of pars anconalis. over the forearm (1955%) part state that M. rest of the distally, to insert flexor ventrad forearm. through or flexor the it attaches “the ulnaris, and which their coverts are located to insert tendon length wide on the tendon digitorum of end of of to the OS ul- sublimusl does not do so in G. c. tabida and it is difficult secondaries carpi brevis the entire in G. c. canadensis of the wing I= The muscles and extends see how the brevis tendon, located on the dorsal surface of the forearm, sublimus into a thin of pars anconalis superficial tendon Its from the deltoid the elbow and expands slips to the bases of the feathers; and process of the scapula. slips arising of origin over the surface The to the acromion the from the dorsal end the tendon to the origin and Goodman posteriorly such branches and Goodman. It arises primarily above, by tendinous fuses, in part, with passes posteriad continues wide crest. passes distad radialis, that arises of M. pectoralis. of insertion (2 cm. wide), stated pollicis) brevis) of the deltoid crest and from pars propatagialis process of having a belly about 9 cm. long and 2 cm. wide; but has a small attachment main tendon is reinforced, apo- longus and brevis. of the longus tendon developed, This brevis and gives rise into the manus to fuse with its fascia and with the (propatagialis BREVIS This muscle is weakly of pars of the superficial pectoralis. were found in G. americana, but not in G. c. cunadensis, by Fisher M. is a M. pectoralis, is a continuation of the tensor patagii a part of M. abductor I found no branches biceps slip becomes tendinous, to the tendons of insertion the insertion of the tensor patagii and their the same as The fleshy longus tendon. which surrounds but slips extend Crane, the inserting Crane. patagii both of the tensor patagii area of insertion the carpometacarpus, and Goodman. crest, which contributes which and others. longus is essentially tendon of M. biceps brachii, the distal end of the belly to parts of the tendons main (1950:427), longus and the tensor patagii is a wide of the fascial The by Fisher on the elastic part of the tensor small tendon, attached layer Young of M. tensor patagii tendon of origin of M. of flex. for me to could pass postero- on Mm. on the posteroventral flexor digitorum surface of the THE WILSON BULLETIN Fisher “M. and Goodman is single.” strong (1955:43) ... tens. pat. brevis Unilaterally components ponents, in process) of the h ‘ umerus carpi radialis) aponeurotic flexor triceps to the lateral epicondyle metacarpi to the OS ulnare. The extensor of M. extensor expands digitorum origins of Mm. of M. extensor extends meta- into a broad communis to the tendon aponeurosis com- (ectepicondylar and scapulodigitorum the entire length to the origin of its fleshy fibers. of the three components of M. extensor metacarpi of the brevis radialis, about PECTORALIS Crane, into Crane In the Sandhill carina, nearly this is a single muscle and is not divided, superficial however, do insert from and deep layers. by a broad Crane, M. pectoralis the posterior the entire length 1890:70). above Pars Fasciculi aponeurosis arises from of the clavicle. propatagialis from as in the Whooping deep surface of origin approximately of the belly, of M. biceps brachii. the inferior third parts of the body of the sternum, I found no origin surface of the deltoid is entirely from the “tracheal crest (pectoral aponeurotic; its of the and from enclosure.” crest of Shufeldt, attachments were described (p. 283). M. SIJPRACORACOIDEUS M. STERNOCORACOIDEUS CORACOBRACHIALIS POSTERIOR All are similar and Goodman, LATISSIMUS There Crane, in origin In in this complex the origin arises from 1955: 48). fibers differences the latter, pars anterior by fleshy to these muscles in the Whooping (all?) In the Sandhill from the neural between captive bird) head; at midlength. beginning the “thoracic” (= spines of the first head (= Pars posterior, of the last three tached small, (Nos. tendon, Crane, 4, 5, and 6) iliotibialis = immediately insertion and and pars anterior (2 cm. in the (5 cm. wide) of the deltoid to the humeral (Fisher (anteriorly) on the crest and the artic- attachment of M. tri- M. triceps scapularis). arises by an aponeurosis dorsal vertebrae, anterior proximal the Whooping and about 3 cm. wide has a fleshy posterior edge of the ilium. portion In vertebrae four dorsal vertebrae; to the junction is immediately in the Sandhill to the anterior flat anterior M. scapulotriceps edge of M. extensor to the uppermost Pars dorsal) Crane, it arises by an aponeurosis about 4 cm. distal this insertion ceps, scapular terior (Fisher G. americana and G. cana- seems to be less extensive. is a thin fleshy band, about 5 cm. wide at its origin, humerus, Crane DORSI are a few minor Goodman, and insertion 1955:46-48). densis tabida. ular the on the tendon and anterolateral The muscle inserts on the ventral M. The latter the most distal of pars anconalis In the Sandhill M. the tendon three fascia, sends slips to the bases of the feathers, on the tendon 1 cm. proximal with This Crane has three of the epicondyle band. proximally (between radialis). proximal lateral over Mm. fuses with the antebrachial inserts most on the and has attachments of the Whooping in the other birds the tendon of G. c. tabidu, the tendon The being connected passes posteriorly and extensor and attaches M. fascia. to insert (without radialis, of the forearm, tendon in one specimen of insertion; and, in part, fuses with the middle sheet, which and by weak passes proximad metacarpi communis that tendon in one of my specimens connected part, noted has a divided Deeemher 1956 Vol. 68, No. 4 (= Pars from sartorius), posterior to the insertion of the scapulotriceps anchor. from the neural the fascia covering and by an aponeurosis inserts on the humerus of pars anterior, In spines the anat- by a and posterior G. americana pars posterior Andrew Berger J. MYOLOGY “attaches to the deep side of the anterior the fleshy insertion Fisher component (1955:&l) bilaterally of the Sandhill SUPERFICIALIS in from five dorsal the first entire state that the dermal ET two muscles are similar less extensive M. cucullaris, hals pt..” M. rhomboideus Fisher M. rhomboideus spines of the last cervical unilaterally (M. Crane. latissimus I found dorsi a minute in a second specimen, but not Crane. the origins superficialis but arises and Goodman of this muscle “is a caudal of the scapula in the Sandhill beneath PROFUNDUS vertebrae. of origin on the humerus component in the Whooping in the two cranes, G. c. tabida. aponeurosis but also inserts in one specimen, specimen RHOMBOIDEUS These part CRANE portion.” may or may not be present at all in a third MM. of the anterior and Goodman metapatagialis) dermal OF SANDHILL Crane. superficialis and insertions (1955:51) extension are by an aponeurosis point out that the of the aponeurosis of inserts on all but the caudal 3 cm. M. rhomboideus profundus and the six dorsal vertebrae; arises from the neural it inserts on the caudal 10 cm. of the scapula. In both cranes, an unusual M. rhomboideus M. CORACOBRACHIALIS This pect of the shoulder; mostly by fleshy sertion M. In fibers rhomboideus G. americana, M. canal of the coracoid margin muscle (about 6 cm. long) surface by Fisher and Goodman deltoideus minor by two heads: G. c. tabida, on the ventral membrane; as- It arises anterior to The in- (The fibers insert (scapulohumeralis of Mm. it arises in- process (procora- a dorsal head from ventral distal present in some birds; and posterior on the tendon the head corresponds, such an inserts on the tendon of M. supracoracoideus.) minor conceals anterodorsally PROSCAPULOHUMERALIS in G. c. tabida head from the medial process of the scapula. none of the M. deltoideus See the descriptions to the insertion of that muscle. In of M. As in G. the tendon of M. supracoracoideus. anterior) subscapularis and “proscapulohumeralis brevis.” SUBSCAPULARIS The muscle that Fisher proscapulohumeralis M. subscapularis structure tween surface and Goodman is actually is similar in all genera pars externa of the scapula, The external beginning head arises from Insertion is on the capital for G. americana PROSCAPULOHUMERALIS As Fisher the medial and Goodman caudal surface groove (1946:584) external of M. and, in fact, exhibits tendon immediately 5 cm. long. and Fisher (pars In G. c. tubida Th e inserting arises from in detail head in the two cranes is larger, (as described (1955:52-53) external I have dissected. and pars interna. the two heads. the head, which “M. than (1955:51). has a single head; a ventral the two heads fuse and insert supracoracoideus; M. located edge of the humerus. of the head of the coracoid, and from the coracoclavicular of the acromion accessory head, when present, however, M. is larger and from the deep surface of the biceps tendon. in part, to a small accessory head of M. supracoracoideus americana, profunclus MINOR side the triosseal ventral fleshy from the dorsal is as described coid) M. the belly does not cover the anterior of M. biceps brachii, DELTOIDEUS is that ANTERIOR is a well developed the origin feature superficialis. of M. serratus M. the same general it arises by two typical anterior heads: passes be- an area 3 cm. long on the lateral to the glenoid of the scapula and internal by Fisher call subscapularis. lip. tuberosity and Goodman, The internal over an area about of the humerus 1955:53). BREWS” (1955: 53) state, this muscle is “very easily overlooked, for December1956 Vol. 68. No. 4 THE WILSON BULLETIN it lies between scapular “was the posterior head of M. not found one specimen edge of M. delt. major triceps.” uniformly Furthermore, in the Whooping of the Sandhill Crane, mutilated separate muscle was involved In the Sandhill 2 mm. wide. fossa and Crane of insertion (= (p. proscapulohumeralis to the origin the internal these muscle may G~ZLS. He the external (1901644; 1915 :415) M. humerotriceps that (= (= and illustrated in the pneumatic imal and/or fossa. anterior M. either the birds, and on the In to agree that with brevis” origin Fiirbringer the origin previously The muscle in other M. which Fisher M. DORSALIS SCAPULAE in origin (serratus POSTERIOR In the origin of this complex Crane, and a deep layer. primarily and for example, of a in development the genus forked anchor” Figs. in which in the insertion origin” of He noted of M. scap- 58%260) also it does not insert the muscle inserts proxIt seems likely, and Goodman Its area hut its insertion of origin differs there- call “M. proseems slightly from proscapulohumeralis In G. c. tabida (see also M. expansor brevis” in the it arises from the pos- secundariorum and Fig. 1.) posterior) there are minor and, in the latter, differences between the Whooping the muscle is not separated in the Whooping Crane. Crane into a superficial In the Sandhill Crane, the in shape, being about 3 cm. long and 4.5 cm. wide at its origin, from the shafts and uncinate origin “M. posterior) superficialis It is so divided main belly is rectangular is some fascial the group. and insertion. terior 9 cm. of the blade of the scapula. and the Sandhill called in that (scapulohumeralis muscle is typical SERRATUS the to the humeral (1902:547 proscapulohumeralis. (1946:587) is M. proscapulohumeralis M. In reported. Cathartidae This proname it arises pos- No muscle the variation “near in genera birds, M. fossa of the hum- or of M. humerotriceps. gruiform area the he did not investigate humerus and Pelecanus, is actually call Thus, most genera, fore, that in the genus Grus, the small muscle which Fisher scapulohumeralis Goodman as I have seen it in other genera. to most of the origin of M. . The of M. proscapulohumeralis considerable though proscapulohumeralis In Ciconia origin fossa of the humer- humerotriceps. the and is attached scapularis). fibers, muscle discussed here. orders. heads of M. anconaeus humeralis), anconaeus to the humeralis) and inserts in the pneumatic inserts to the glenoid by fleshy subscapularis. the relationships discussed this muscle Fisher of M. of other in gruiform also that “in Otis it is much reduced, ulotriceps a muscle. However, phylogenetically. stated that about 0.5 cm. proximal of the pneumatic which head and external migrate caudal It inserts triceps for the rudimentary specifications. proscapulohumeralis it in by shot or be so not be aware and lateral heads = margin the muscle in representatives meets described pars posterior, of M. scapulotriceps erus, between Mitchell 285), is available cranes be destroyed one might on the humerus and external this muscle does not exhibit as I have seen them that scapulotriceps. Crane, dorsi, internal is actually cranes, however, of M. of M. to the humerotriceps above scapulohumeralis terior origin latissimus lateral As mentioned of the that in a second, and not at all in a third. that it might edge of the scapula just is on the plane of the inferior us, but is entirely part 5, 1955) I found this muscle bilaterally specimen as in the Whooping triceps, May band of fleshy fibers 3 cm. long and only about the ventral to the of M. humerotriceps preserved this is a minute anteroventral to the insertion however, (letter, at all. It arises from and not by a tendon M. a poorly in handling wrote Cranes.” unilaterally The muscle is so small and delicate, and the most proximal Fisher processes of true ribs numbers also from rib number 6. The insertion 3, 4, and 5; there is almost exclusively by an Andrew Berger J. MYOLOGY aponeurosis Crane, on the ventral the superficial external layer of intercostal The humeral belly tract. face of M. expansor This is similar the arising fascia upward by three between two and Sandhill covering In arising the posterior continues margin distad of the along the sur- fleshy the Sandhill it arises by fleshy 1, 2, and 3, and vertebra. there are minor In the latter, slips, one each from The muscles. the two heads of M. Crane of true ribs numbers anterior) muscles, cranes. the intercostal differences the serratus the first three dense aponeurosis suhscapularis and inserts in origin anterior true be- is a small ribs and from of insertion on the ventral passes edge of a short distance caudal to the glenoid lip. muscle has a single belly ular-shaped of Mm. surface BICEPS This the of the coracoid, subcoracoideus and in one wing M. belly the same relative (about near radius and ulna. The 14 cm. long) the distal larger (see M. tensor patagii DELTOIDEUS (Fisher the humerus. tendon inserts on the humerus, In G. c. tabida in the two cranes. proximal two-thirds of the arm. but not at the insertions on the radius. The The on the biceps slip is present major is about The fibers 12 cm. long and extends of insertion The primary from the blade Crane. posterior fibers and Goodman TRICEPS (triceps This complex connection the scapular lip of M. Crane, this aponeurosis of M. I scapulae. did not mention did slightly posteriorly of the deltoideus is a secondary and the origin dorsalis origin, less than with major for G. the latter, the humeral is on the dorso- and Goodman is attached scapulotriceps, the half way by a flat aponeurosis by Fisher not find described In (7 for the about 3 cm. caudal and dorsal to the an- an OS humeroscapulare and it. brachii) is similar or anchor head are in contact origin of the scapula as described In the Sandhill glenoid component is absent in G. c. tabida. 1955:57) surface of the scapula, and there mm. wide) Fisher The tendons to each other in the two species, but the dermal anchor of M. scapulotriceps. teriormost development lies in the end of the humerus, and Goodman, down to the adjacent from the an- MAJOR of deltoideus Whooping insert at its origin of that bone. longus). muscle is similar belly lateral width to the middle they fused at the insertion. is ossified americana In G. c. tabida it is a very small triang- BRACHII tendon This just dorsal and subscapularis muscle exhibits small in the cranes. muscle, 4 cm. long and 1 cm. in maximum teromedial M. metapatagialis) 4 and 5 and from the intercostal SUBCORACOIDEUS This M. superficialis tendon the angle, superficialis preceding the scapula, beginning M. surface, near (serratus of the the Whooping muscle the almost to the elbow. process of the last cervical ANTERIOR is true tween In the Whooping arises from ribs 4, 5, and 6. opposite this area, a fibrous in the two cranes. from the lateral SERRATUS As M. serratus in the metapatagium From secundariorum from the transverse M. end of the scapula. 4, 5, 6, and 7 and from fascia overlying The deep layer (= 287 PROFUNDUS complex fasciculi CRANE surface of true ribs numbers terminates feather SERRATUS muscles.” component from the lateral fascia. M. edge of the posterior layer arises from “ribs I found a large dermal primarily OF SANDHILL (z (1.5 in the two cranes. cm. long In G. c. tubida there is a strong aponeurotic and 1.5 cm. wide) M. scapulotriceps) between and the humerus. the anterior The humeral this band begins about 3 cm. distal to the head of the humerus surface attachment and lies immediately of of an- terior to the insertion from the entire tinctly of pars anterior inferior divided bicipital into almost to the level exclusively margin latissimus margin dorsi. head. by a wide tendon. end of the humerus, The tendon of insertion arises and is not dis- A few fasciculi of M. dorsalis scapulae. of the belly near the middle of the distal M. humerotriceps fossa of the humerus and an internal to the area of insertion discussion of M. anconaeus arise from the An ossified tendon of the arm. Fleshy but the insertion does not contain fibers arise on the ulna is a sesamoid. (See coracoideus.) BRACHIALIS This muscle is typical M. EXPANSOR Fisher and Goodman did not mention G. c. tabida, M. expansor In bow. This numbers relations, muscle about is a smooth 17 through band, connected over one-third the dorsal origin secundariorum which the layer way up the tendinous fibers about 2 cm. from the insertion corner er, dorsal branch passes dorsomesiad the procoracoid. the scapula to the sternum ton (1896: M. ANCONAEUS This 6081 called discussion Crane (Fig. which of M. by Fiirbringer coracoideus of M. dorsalis the tendon scapulae, bifurcates about at- the small- called the tendon internurn” extending ligament; New- ligament.” of M. anconaeus tendon of M. and earlier muscle the proximal resents one tendon In the near For a Sandhill 1 mm. in secun- gives rise to a second tendon, the distal noted by Fiirbringer. Fisher end of the humerus. and Goodman do not Crane. RADIALIS in the two cranes. (about surface a part of the tensor tendons, papers). (1956:159). tendon of M. expansor coracoideus scapulotriceps muscle, as previously METACARPI Two (1902:576, see Berger of this muscle is about 4.5 cm. long, but only about muscle is similar anconalis. at the apex of the belly edge of the scapula, near its (1902:572) described the belly on the spindle-shaped where of the has not been process of the sternum; the “sterno-scapular this muscle in the Whooping EXTENSOR from epicondyle apparently, skin of origin. of the tendon has its major It arises by a tendon from the “scapular” Distally inserts This branch to the ventral anconaeus 11, the belly width. This is a striated mention to attach to the that skin fold and pierces the as a pulley) ventral a fibrous 16. The belly CORACOIDEUS recent dariorum. is formed through In the axilla, Fiirbringer of secondaries attached to the medial the “Sternocoraco-scapulare it simply muscle was first maximum function are tri- to the el- has two tendons this origin, of the sternocoracoidal articulation calami to number fasciculi A second tendon The larger, attaches attached runs proximad (which on the muscle brevis; of that muscle. to the pulley. to the medial pulley, band of M. pronator This tendon semitendinous roughly several of the distal tertials; The for the cranes previously. from primarily where described with inserts arm, is by a flat in the metapatagium. 1) is a well developed, of the metapatagium. distal to the origin 3 cm. proximal (Fig. to the humero-ulnar humerus, lowermost must be present and 10 cm. long and 3 cm. wide at its base posterior muscle, primarily forming distal this muscle, but it certainly 23 and on the skin containing extends tachment and insertion. in G. americana. angular-shaped The in origin, SECUNDARIORUM well developed M. of M. of the pneumatic an external crest anterior forms on the ventral M. December 1956 Vol. 68, No. 4 THE WILSON BULLETIN 288 6.5 cm. long), of the olecranon patagii whose fleshy Crane, fibers process and about brevis tendon interconnected of insertion In the Sandhill fuses with brevis; about 1 cm. distal the tendon by fascia, are present. of M. tensor patagii pars anconalis begin The the posterior to the area of origin anterior is a 5.5 cm. of pars tendon tendon, rep- the ori- MYOLOGY OF SANDHILL CRANE 289 FIG. 1. Ventral view of certain muscles in the proximal region of the wing of Grus tabida to show relationships of Mm. expansor secundariorum and anconaens coracoideus. The distal end of the humerus is distorted in order to show the triceps tendon and the origin of forearm muscles. Not shown is the fascial extension of the flexor digitorum sublimus tendon, which invests the superficial surface of M. flexor carpi ulnaris. Explanation of symbols: Ant., anconaeus coracoideus; Dor. scap., dorsalis scapulae; Exp., expansor secundariorum; Flex. c. u., flexor carpi ulnaris (humeroulnar pulley not shown) ; Flex. dig., tendon of origin of flexor digitorum sublimus; Met., metapatagium; Pro. brev., pronator brevis; Pro. long., pronator longus; T. hum., camdensis humerotriceps; T. scap., scapulotriceps. gin of pars anconalis. Both tendons are attached to the lateral epicondyle of the humerus. Pars palmaris is developed as illustrated for the Whooping Crane by Fisher and Goodman (1955: Fig. 16). The fleshy belly is about 7.5 cm. long. Pars anconalis and pars palmaris each give rise to separate tendons, which fuse to form a single ossified tendon of insertion. In addition to the usual insertion on the extensor process of the carpometacarpus, part of the tendon fuses with the tendon of M. extensor longus digiti II (= extensor pollicis longus), as in the Whooping Crane. M. EXTENSOR DIGITORUM COMMUNIS In the Sandhill Crane, this is a small, spindle-shaped muscle (8-9 cm. long), which is located in a little more than the proximal third of the forearm. The tendon bifurcates near the base of the pollex. The shorter branch inserts on the posterodorsal edge of digit II (= pollex), about 0.5 cm. from the proximal end of that bone. Fisher and Goodman THE WILSON (1955: Fig. 20) as inserting illustrated and Goodman), I found no branch Whooping Crane &I. SUPINATOR There this corner In II differences Sandhill from the lateral RADIALIS (c?XteIISor METACARPI exhibits small, similar spindle-shaped width. to, the anconeus tendon. M. ; It muscle superficial space origin. arises from the the tendon but 10 cm. long) distal Fleshy extends more less than than beginning one-third tabida EXTENSOR LONGUS it arises distance of insertion. from only The proximal and posteriorly. DIGIT1 there In the Sandhill about Crane 0.5 cm. in in common is an aponeurotic (pronator with, and attachment than sublimus and pronator to profundus) as in G. americana. M. pronator M. (belly pronator M. longus brevis pronator it inserts end of the radius pronator II (f?XkIISOr the radius brevis primarily longus 8 cm. long), extends slightly by an aponeurosis, and extending to within 13.5 cm. cm about the same areas in G. c. over an area about 7 cm. long; M. The a distance ulnar (about 1OIlgUS) development origin begins tendon II length) distally, is ossified, metacarpi and but except near radialis; much The the distal (for in about where tendon a biceps muscle-sheet the insertion, the common ulna to the is located is a flat the proxiit fuses inserts on the (I). belly in the two species. larger than (about the In the Sandhill tendons 13 cm. long) of Mm. extends extensor distad Crane it arises by a very digitorum slightly more of the ulna, and thus inserts on somewhat more than the proximal M. EXTENSOR Relative In the Sandhill from immediately 13 cm. in overall it is rounded of M. extensor in the two birds. of 6 cm.) (RIICOIIReuS) tendon, supinator. pOlli& (for The belly muscle is similar length also near the base of the intermetacarpal the forearm. the same relative process of metacarpal ANCONEUS large fibers communis. but it is only The two muscles insert of the forearm; the tendon This The muscle some fleshy end of the humerus of the radius; M. 2 cm.). small half extensor M. brevis. one- uhrk) Crane, further the proximal muscle exhibits Crane, with of about fibers arise from this tendon 5 to 6 cm. from LONGUS way down length G. americana: development extends brevis over an area 4 to 5 cm. long. This mally distad end of that bone. and pronator M. half the 4 cm. from of the distal for the is ossified. M. PRONATOR extends species in the in the two cranes. These two muscles have the same relationships (belly and pulley. and illustrated 8 cm. long) digitorum The muscle inserts of insertion BREVIS AND two 13 cm. long, As in the Whooping end of the ulna. PRONATOR a fibrous 20). of the humerus; Carpi relationships maximum the proximal and in- (of Montagna as far distad as M. pronator iV1. FLEXOR its humeral III as described it (p. 60) is ossified passing through (about epicondyle of M. extensor it is a very of digit and Fig. the rhe belly of origin muscle (I) between Crane arise from the tendon This after (1955:60 of the radius, i.e., nearly arises by a tendon but described tendon (SLIpiIIatOr) the third the length Crane, phalanx (of Hudson), and Goodman are no important muscle. II on metacarpal by Fisher BREVIS of the proximal or digit inserting in the Whooping In G. c. tabida the longer finger.” serts on the anterobasal Fisher this branch on the “third December l!JSO Vol. 6% No. 4 BULLETIN LONGUS DIGITI III (extensor to the size of G. americana muscle in genera sidered rudimentary spindle-shaped of other belly families, in the cranes. 12 cm. long M. indicis communis than half half of that bone. longus) and G. c. tabida and to the development extensor longus In the Sandhill but only or the about digiti Crane III might almost of this be con- this muscle has a very small, 3 mm. in maximum width. It arises Andrew J. MYOLOGY Berger from the posterior surface and from the “middle 1955:64). It M. DIGITORUM FLEXOR tion extends is ossified manus; it inserts this muscle torum sublimus. M. FLEXOR and “anterior inserts and cranes, it definitely complex is tendon Fisher extensive in small rudimentary, limus ; not found ulnaris and manus, where the fleshy from the families FLEXOR to the carpi and The ulnaris flexor it as the (1946:598) part” is the flexor rudimentary by in the the distal in edge of this deep begin of the which on the The sublimus tendon The III of the tendon muscle anterior passes distad tendon, and and Fig. 13. part M. digitorum the the anterior edge, about midto the has a more distal Goodman discussion of the “anterior 1946:606 carpi into continuation therefore, Fisher of the and is flexor and along on the anterior tendon. is entirely as it passes of M. muscle) deep to the edge fibrous tendon but there is a fascial sublimus profundus the of this complex sublimus The sub- 4 cm. distal becomes to the primarily (II), in their tendon (anterior palmar digitorum and from sublimns inserts inserts, has several on the flexor by but G. c. tabida. about an accessory insertion ulnare but fascia and of the a thin the tendon aponeurosis fibers humerus, Crane tendon, distally, the of the of insertion of the (OS cuneiform), G. americana fleshy end for the Whooping part” (1955:66 of M. flexor ULNARIS and the anterior (1955:65567), G. c. tabida. (= passes through flexor this muscle exhibits In Gras arises by a very strong tendon immediately digi- and as it is described to the ulna; surface The digitorrrm digitorum from 10 cm. long) and inserts this insertion and Goodman G. americana the (II). because most of the muscle- as described to fuse with ossified. of digit see also Fisher, one excludes Fisher describe “anterior is confused both accessory it becomes the flexor CARPI III flexor and Fisher be considered OS ulnare tendon, of the phalanx also described If of M. they genera posterior (about tendon of birds. surface tip of that phalanx. manus represents of the carpometacarpus M. might to attach anterior attachment again ulnaris; though This the found (humeral) of the distal carpi of of inser- of digit to the insertion manner. arises of the tendon. surface 67) The tendon phalanx (1890:141) it 1) belly ossified main posterior than brevis. half of M. brachialis; end of the ulna and into of the distal muscle, longus, From are humeral anterior (Fig. corner length, insertion ulnaris Shufeldt situation passes posteriad small, the in all the The pass into the latter around this Though (1955:66). arises from of the insertion the distal proximal ulnaris. pronator relationships The separate ulnare of M. anterobasal of the main humerus. discuss of origin bipinnate, muscle surface corner in a similar is present. Goodman These margin carpi poorly developed to less than the proximal and aponeurosis. slips, which surface. flexor phalanx (1891:278). tendon on the of M. carpi complex aponeurosis part, is limited it passes around do not to the origin and Crane and Goodman, (II). muscle as I have seen it in other Selenka strong posterior (Fisher in the two species. The relatively at the inferior on that flexor this sublimus Gadow III in the Sandhill Crane SUBLIMUS of M. also interpreted digitorum origin where Goodman part” of the radius of digit Crane on the anteroventral DIGITORUM Fisher The relationships to the origin except Thus, phalanx in the Sandhill It has a V-shaped the origin half PROFUNDUS (10 cm. long) the ulna. the distal CRANE of this bone in the Whooping on the distal This muscle has similar belly of about third” inserts OF SANDHILL a strong and apparently sublimus) about by in in all from the medial humero-ulnar as described the same development other (internal) pulley. birds, humeral I have never M. flexor condyle seen any THE WILSON 292 departure from this relationship. The bulk of the belly developed. third of the almost the forearm, though to the distal anterior the “attaches thirds of the ulnar (about length” (Fisher M. This and muscle, the palmar M. DIGITI II head. belly In M. from become surfaces. On The inserts This II, muscle fibers DIGITI is well II DIGIT1 this is a strong tendon on both then its dorsal over the of head arises insertion of fans to out and palmar two heads arise from “the tendon of the the tendon of cranes METACARPI (pollex). edge of digit cranes. from In the metacarpal III surface brevis digiti is composed III Hudson II palmar process; it head inserts in its basal half. (II). Crane The II it bulky arises by a belly passes (pollex). In the Sandhill Crane the belly is III) of connective it would be better part of the preceding digiti tissue (Fisher to consider and this not muscle. II) In all birds the carpometacarpus with that digit I have dissected, this structure is III. BREVIS is absent in G. americana and in G. c. tabida. suggest that this name “be dropped represents the extensor The Sandhill of digit mostly that as a distal (abductor seen such a muscle. and 43) arises from II III) (flexor IV DIGITI connecting muscle the inserting the palmar There radialis. arises from 0.5 cm. wide. muscle, but simply muscle II muscle in the cranes. I agree with MINOR I have never (1955:35 DIGITI in the 1955:68). as a separate This the I have dissected, metacarpi the Crane: manus Crane of the ulna. pollicis) in the digiti developed BREVIS M. FLEXOR genera this head from but half profundus. extensor of this muscle on the anterior 1 cm. wide (flexor Iv structure a ligament a each it has both a palmar passes superficially on most of the posterior is a weakly M. ABDUCTOR In radialis. alae digiti (adductor developed about to insert Goodman, two- Crane, In the Sandhill other fibers corner of the base of digit 3 cm. long and less than This in the Sandhill digitorum of M. process, of the fascia head of the abductor ALAE RI. FLEXOR metacarpi deep fascia ulnaris proximal on the base of the ulnare. in G. americana but none of the fibers and tendinous FLEXOR This of insertion extensor this ; carpi the longus. aponeurosis anteriad the the from than the distal as in all by “tendinous extensor on surfaces, on the anterior BI. ADDUCTOR iv. both anconal by fleshy flat with alae digiti patagii of M. longus continuous abductor tensor patagii more This is not true for the Sandhill the tendon tensor and M. ventralis) of M. flexor Crane, state that process” tens. pat. longus.” only 1955:67) in the two cranes. slightly the tendon of arising in the two cranes. (1955:68) extensor As in the Whooping is similar Sandhill on and second fourths flexor pollicis) the tendon forms part” and also inserts to the origin the of insertion of the ulnare. (abductor arises from and Goodman base of the arises from apparently, an anconal Fisher muscle, similar relationships ALAE belly poorly in the proximal accompanies tendon of the first feathers (ulnimetacarpalis BREVIS 12.5 cm. long) ABDUCTOR portion posterior and Goodman, this posterior ULNARIS It has the typical the is relatively is located fibers ossified the junction over the bases of the forms from CARP1 of the belly 15 cm.) of fleshy strong, at about fasciculus This is a well developed belly bundle The Crane about it inserts on the posterobasal to the fascia M. rrkxcm Sandhill length a small of the belly “superficial small tendon the end of the ulna. surface of the forearm; Crane, In (total Deeember 1956 Vol. 68, No. 4 BULLETIN a distal head of M. extensor from Hudson and Lanzillotti the literature” indicis longus inasmuch (extensor as longus MYOLOGY digiti III). Data which use in some families. flexor M. metacarpi hrevis INTEROSSEUS This I have I think, obtained suggest therefore, CRANE that this that it would for indicating the presence head may he convenient he of taxonomic to retain or absence of this the name small muscle. DORSALIS muscle is similar on the OF SANDHILL in the two cranes. base of the distal phalanx In the Sandhill of digit III ; (II) Crane, a small it inserts tendon primarily continues to the tip of the digit. M. INTEROSSEUS In three-fourths phalanx of digit The Crane DIGITI though pollicis about III (abductor in the (1955: species. a few fleshy fasciculi at about the junction insertion is typical. II (fleXOr the FLEXOR and Goodman in the two cranes, This same Sandhill insertion base of the distal as illustrated for the 18). the Sandhill Crane, the muscle of the proximal The very and middle small, deep is process. thirds head, of described in G. c. tabida. is present pollkk) base of the carpometacarpus M. is developed Fig. In the in to the 2, about The arise at the level of the pisiform forms Similar of phalanx 1955:69). indicis) two tendon DIGIT1 aspect brevis) 1.5 cm. long) carpometacarpus; FLEXOR posterior and Goodman, is also anchored and Goodman DIGITI is similar tendinous, the tendon (belly for G. americana by Fisher M. on “the (Fisher (extensor II by Fisher MAJOR muscle ossified the VOhrk) inserts (II). BREWS ABDUCTOR This mostly III very small muscle Whooping M. muscle of the way out its length” EXTENSOR This (iIItWOSSf?Us this in G. c. tabida, but is similar M. VENTRALIS G. americana this small (1.5 and inserts cm. long), fleshy on the posterobasal muscle corner arises from of digit II the (pollex) Crane. METACARPI is a poorly relationships (ulnimetacarpalis POSTERIOR developed muscle with as described G. c. tabida the two smaller by dorsalis) a belly Fisher 3.5 cm. long. and heads are mostly Goodman tendinous In general, (1955:70), it has the except that in bands. MYOLOGY OF THE LEG M. EXTENSOR This ILIO-TIBIALIS extensive muscle by an aponeurosis of the posterior throughout Goodman is similar from the entire iliac a thin (iliotibialis) LATERALIS crest. in the two cranes. anterior Some of the (1955 :76) pointed out, “the part of the muscle is aponeurotic ilio-tibialis of M. Mm. M. lateralis extensor flexor EXTENSOR In the lateralis ILIO-TIBIALIS Sandhill of M. and anterior from the muscle the Crane, the 5 cm. posteriorly Crane is by fleshy edge is the thickest. in the center edges.” superficial cruris dorsi of it arises fibers. It As Fisher is and of the muscle are less than The distal half of the central muscles. view the anterior The extensor and superior half the bellies of medialis. (SRrtOriuS) muscle is no origin same Sandhill biceps femoris) , but it does not conceal and flexor latissimus there exhibits (= this the and is fused with the underlying ANTERIOR Crane, pars posterior Whooping fibers and posterior muscle conceals from ilio-fibularis cruris origin sheet of muscle, but the posterior half as long as those of the anterior In iliac crest and from all but the caudal 1 cm. arises from the primarily the median by an aponeurosis spine of the last dorsal dorsal ridge of the from the last dorsal vertebra. configuration in shared neural the two cranes. synsacrum. In In general, the Little with vertebra In the however, Brown THE WILSON Crane, Hudson (1937:17) The insertion M. found in the Sandhill (gluteus PIRIFORMIS medius As stated by Fisher but it is “more the origin Crane by a flat (1955:79), developed” trochantericus 'M. M. (= M. this area muscle specimen trochantericus anterior this complex femur just continuous There however, M. belly FLEXOR Crane about of that ilio- to et 29 and 30.) 69) reported tabida. muscle and M. This the bellies dissections. toward are parallel. The (biceps fibers fused, so the same area, fusion tendon posterior. of these two inserts Though arise on the there are muscles, there is an almost (femoritibialis complex insert the ilio- of muscles which common of 3 cm., beginning on the trochanter. externus between externus” from The fusion iliotrochantericus in this and are completely arising uni- iliotrochan- fuse distally muscles it in and hip of one specimen, muscle-mass, other femoritibialis found in two specimens medius MEDIALIS that they did not (1955:123) and medius) the two cranes; see, below. femOriS) from all In the Sandhill but the Crane posterior the well developed 1 cm. of the posterior 4 cm. distal to the proximal articular bone. exhibits crest. The continues to become the middle (semitendinosus a similar muscle raphe and accessorius semitendinosi) configuration arises from which downward continuous with separates between in two cranes. of that In the posterior the semitendinosus pars media the tendon is accompanied the approximately of the belly of M. gastrocnemius. muscle, the raphe head of Fisher dorsal medius iliotrochantericus Figs. it bilaterally only; in the two cranes. CRTJRIS LATERALIS iliac gluteus to M. and Goodman The strong tendon inserts on the fibula semitendinosus M. 1955: (193760, tendency of M. VASTUS by fleshy the semitendinosus cnemius (= arise directly of for the three iliotrochanterici of “M. is similar complex posterior relationships In the right in the general differences ILIO-FIBULARIS of that This muscles for a distance AND no significant arises mostly surface origins by a single to the insertion LATERALIS muscle Its Fisher iliotrochantericus of the the discussion iliac crest. M. and line of insertion EXTENSOR This but areas and whose fibers are origin and Goodman, aponeurosis. tendons of insertion VASTUS cranes. at their cm.) by both distal two distinct MM. (1.5 is an example adjacent muscle the and Hudson is represented occupied of this by of G. canadensis are separate wide from two G. c. canadensis. I found and anterior muscles It inserts on the profundus posterior) (See Fisher in the genus Grus, in a third however, gluteus anterior) (1891:142) by a common, that 4 cm. long. of M. MEDIUS and Selenka G. americana tericus in the two cranes, In G. c. tabida it is a triangular- and about pre-empted in the below. ILIOTROCHANTERICUS this only.” (1955 :79). . is similar are described laterally being (iliotrochantericus muscle Gadow find (origin) to the insertion (iliotrochantericus piriformis) ILIACUS medius anterodistal PROFUNDUS acetabulum, This and Goodman this muscle is similar in G. c. canadensis. in the two cranes, none of the fibers minimus edge of the ilium by Fisher posterior). CLUTEUS Similar the tendon the “anterior et minimus) shaped muscle, 2 cm. wide at its base femur from is as described and Goodman strongly December 1956 Vol. 68, No. 4 BULLETIN from and pars interna muscle; Sandhill the accessory of M. the raphe In its course between by a small fleshy the 1 cm. of the gastro- is ossified the two heads belly, the “distal accessory” and Goodman. As in the Whooping Crane, there are two distinct parts to the accessory semitendinosus MYOLOGYOF SANDHILL CRANE Andrew .I. Berger muscle (Fisher fibers (popliteal flexor the region) tendon of in the FLEXOR CRURIS with the just before inserts raphe the beginning found and The the internal to the proximal (1955:85 in this complex. They and found of the Whooping In one dissection, arises they found is a very primarily of the band thin, iliac three from bony notch by a long, width. covering which is well and difference and illustrated two and of M. caudofem.” Goodman’s Figure of variation they pars iliofemoralis in a third in specimen. This Crane. Pars ilio- at its base, where of about the middle 6 cm. in length. It inserts beginning it third by about of M. ischiofemoralis. tendon tendon posterior (3 cm. long, but only 0.5 mm. muscles iliac of the tail; passes through crest; a fibro-osseous the tendon canal. about The I found a bony notch at is held in the muscle 3 cm. distal no inserts by to the trochanter of pars iliofemoralis. In 31. to the in the parts insertion the of the caudofemoralis and Goodman no genus the Sandhill of M. relationships of M. inserts “posterior In In in these cranes. by Fisher state that M. ischiofemoralis parts Crane (ischiofemoralis) developed the and surface the depressor completes 1 cm. proximal is a striking described muscle belly, 8 to 9 cm. long and only about It arises by a small on the pygostyle. ISCHIOFEMORALIS ischiofemoralis The of the bone. amount to the insertion to the insertion There fused on the tibiotarsus, surface of the femur, spindle-shaped end of the projecting about the Crane. In the Whooping end” the is approximately on the lateral medial femur more to gastrocnemins, gastrocnemius. 2 cm. wide the ventral and directly the The is intimately specimens of the Sandhill (2 cm. long and 2 mm. wide) muscle and femur and does not of M. 1.5 cm. wide) emphasized and 1 cm. distal the fascia a ligament, FLEXOR of the the proximal tendon This birds. of insertion pars media head flat M. to the passes lateral but only pars iliofemoralis from has a small attachment the most caudal many head) end of that bone. 123) the belly to the trochanter 6 mm. in maximum direct tendon sheet of muscle, 5 mm. wide) Pars caudofemoralis in diameter) IV, parts to the muscle. triangular crest; (about 3.5 cm. distal of Mm. digiti attachment both pars caudofemoralis Crane, by an aponeurosis posterior a fleshy origin as it does in the Whooping and from I found both parts to this muscle in three femoralis fleshy region (PirifOrmiS) Goodman two specimens by common semimembranosus) pars media, “some five centimeters CAUDOFEMORALIS Fisher (= inserts intercondylar perforatus as in accessory (2.5 cm. long and about 3 cm. distal inserts The vertical, semitendinosus fuses with aponeurosis about the tendon to the and flexor medialis in the two cranes. accessory latter by a thin than part the (semimembranosus) MEDIALIS of the and proximal III, (distal cruris proximal pars media. rather muscle gastrocnemius, muscle is similar more condyle digiti gastrocnemius, flexor of M. The medial immediately direction, of M. the perforatus accessorius on the belly This of femur, flexor of M. of insertion insert M. of the transverse part 1955:83). surface longus, of origin is nearly tendon Goodman, posterior hallucis distal M. and on the 295 have Sandhill (1955:86 tendons in the (see illustrated pars caudofemoralis on above). of insertion Whooping the insertions seen the condition Crane, it inserts of M. Crane and Figs. 30 and 31). to and between I Crane caudofemoralis as They of the two in Fisher inserts medial in the two cranes. to pars iliofemoralis. MM. ADDUCTOR These M. two adductor SUPERFICIALIS muscles profundus exhibit ET PROFUNDUS about is entirely (adductor the same relative fleshy at its origin longus et brevis) development in the Sandhill Crane and I did THE WILSON BULLETIN December 1956 Vol. 68. No. 4 not find a conspicuous “heavy layer of tendon” covering the medial surface of this muscle such as Fisher and Goodman (1955:88) described for the Whooping Crane. The glistening muscular fascia is well developed, however. M. AMBIENS Fisher and Goodman (1955:88-89) called attention to the differences in termination of the ambiens tendon in G. nmericanu and G. c. can&ens&. In the former, the ambiens tendon serves as the “principal, if not sole, origin for M. flex. perf. dig. II, although there is strong fascial interconnection between the origins of Mm. flex. perf. dig. II, III, and IV, and in one instance there is actually a branch of the main ambiens tendon that goes to the tibiotarsus.” In the Little Brown Crane, “M. ambiens connected distally to the small lateral head of M. flex. perf. dig. III. It had little connection with M. flex. perf. dig. II and none with M. flex. perf. dig. IV.” In the Sandhill Crane, I found that the ambiens muscle arises primarily by a flat tendon from the pectineal process. The small, spindle-shaped belly is 6 to 8 cm. long and less than 1 cm. in maximum width. Distally, a very small tendon (1 mm. wide) forms and has the usual course through the patellar ligament. The tendon then passes distad medial to the biceps tendon and serves as the primary origin for the lateral head of M. flexor perforatus digiti III; the ambiens tendon does not give rise to any other muscle. M. FEMORITIBIALIS INTERNUS There are minor differences between the Whooping and the Sandhill cranes in development of this muscle mass and I found variation in the pattern in the three specimens I dissected. In one right leg, the muscle was indistinctly divided into two heads. In another right leg, one long and two short heads were present; each gave rise to a tendon and the three tendons fused for a common insertion. In the other dissections, there were two distinct heads. The posterior or long head arises from the posteromedial surface of the femur, beginning a short distance proximal to the area of insertion of M. iliacus; the origin is fleshy as far as the medial condyle. The short or distal head arises from a small area (about 2 cm. long) on the anteromedial surface of the femur, just above the medial condyle. The small tendon from the latter head fuses with the patellar ligament and with the tendon of the long head; the combined tendon inserts on the medial corner of the tibiotarsus at the base of the inner cnemial crest. “M. FEMORITIBIALIS EXTERNUS” As Fisher and Goodman (1955:89-90) imply, there is some confusion concerning the muscle they consider under this name and their M. vastus lateralis. Fisher (1946: Table 42) stated that his M. vastus lateralis was a synonym for M. femoritibialis externus of Gadow and Selenka (1891:154) and Hudson (1937:20). This muscle was discussed earlier by Fisher and Goodman on page 81. The muscle which Fisher and Goodman call “M femoritibialis externus,” I believe is simply a distal head of their vastus lateralis (= femoritibialis externus). I considered this head a part of M. femoritibialis externus in the cuckoos (Berger, 1953:68 and Fig. 6) ; it was illustrated, but not given a special name. Mitchell (1901:647 and Text-fig. 79) also called attention to this distal part in gruiform birds. In the Sandhill Crane, the distal head of M. femoritibialis externus arises from the posterior and lateral surfaces of the distal half of the femur, posterior to the more distal origin of Fisher and Goodman’s vastus lateralis. The tendon of the distal head fuses, in part, with the patellar ligament, but the strongest portion of the tendon passes distad to insert on the outer cnemial crest of the tibiotarsus. MYOLOGY OF SANDHILL CRANE 297 Fisher and Goodman (LX. cit.) also state that Gadow and Selenka’s (1891:1551 “M. femoritih. medius is apparently lacking . . . or is fused to M. femoritib. ext.” in the Whooping Crane. However, Fisher (1946: Table 42) placed his M. vastus medialis in synonymy with Gadow’s femoritibialis medius, which Fisher and Goodman described on pages 81 and 82. The discussion by Gadow and Selenka (1891:155) of the origin of the femoritibialis complex is not entirely clear, but points out that the femoritibialis medius passes directly to the patella, and, in many birds, the fleshy fibers do insert on the proximal surface of that sesamoid (see Hudson, 1937:20; Berger, 1953 :68). M. OBTURATOR EXTERNUS Fisher and Goodman (1955:90) said that “Hudson (1937:28) stated that M. obt. ext. had two distinct parts in G. canudensis; our dissection of this species showed the separation to be superficial only.” In the Sandhill Crane, also, this muscle may be a single mass or may be partially separated into two heads. It is a broad band of fleshy fibers with a nearly continuous origin from the anterodorsal and anteroventral margins of the obturator foramen. The belly conceals much of the tendon of M. obturator internus. The externus has a broad fleshy insertion (1 cm. wide) on both sides (proximal and distal) of the tendon of M. obturator internus. M. OBTURATOR INTERNUS This muscle is triangular in shape. It does not arise inside the pelvic cavity, as it does in Porzana and COZUZ.The fibers converge to a tendon, which emerges through the obturator foramen and inserts on the posterolateral surface of the femur, less than 1 cm. from the trochanter. M. PSOAS (ikicus) This is a flat, fleshy band about 4 cm. long and 0.5 cm. wide. It arises from the ventral edge of the ilium about 1 cm. anterior to the acetabulum. It inserts by fleshy fibers on the femur for a distance of 1 cm., beginning 1 cm. distal to the neck of the femur, just proximal and somewhat posterior to the origin of M. femoritibialis internus. Fisher and Goodman (1955:90) said that “the condition described by Fisher (1946:670) for the cathartid vultures is found” in the Whooping Crane. M. GASTROCNEMIUS The three heads of this complex are, in general, similar in the two cranes. The following specific points may be mentioned for the Sandhill Crane. Pars externa arises from the lateral condyle of the femur, as described by Fisher and Goodman (1955:91). The tendon of origin is, in part, fused to the biceps loop. Pars interna has an extensive origin from the medial surface of the inner cnemial crest and from the patellar ligament. In fact, the fibers of M. sartorius and part of those of pars interna insert and arise, respectively, from a tendinous raphe separating the two muscles. Pars media arises by a flat tendon from the intercondylar (popliteal) area of the femur, just proximal to the common tendon of origin for the long flexors. Fleshy fibers begin about 3 cm. from the femoral origin of the tendon. In the left leg of one specimen, I found an accessory medial head. This head (5 cm. long, but less than 1 cm. wide at the origin) arises from the posterior surface of the medial condyle of the tibiotarsus. The b e11y passes lateral to the inserting tendon of M. semimembranosus and anterior to the tendon of the distal accessory belly of M. accessorius semitendinosi. The belly tapers to a minute tendon, which fuses with the fascia covering the deep surface of pars externa (gastrocnemius) in the area where this head fuses with pars media. THE WILSON BULLETIN 298 M. PERONEUS LONCUS Similar in the two cranes, this is a M. tibialis anterior. The strong tendon on the superficial surface of the belly. perforatus digiti 111, 4 cm. distal to the Mitchell, 1913:1053.) M. TIBIALIS December 1956 Vol. 68, No. 4 well developed muscle, which conceals all of of insertion forms as ossified radiating bands The tendon inserts on the tendon of M. flexor proximal end of the tarsometatarsus. (See also ANTERIOR Similar in general configuration in the two cranes, the femoral head (whose tendon is ossified) is almost equal in bulk to the tibia1 head in the Sandhill Crane. The common, ossified tendon bifurcates at the insertion on the tarsometatarsus, about 2 cm. from the proximal end of that bone, but the two tendons insert adjacent to each other. M. FLEXOR PERFORANS ET PERFORATUS DIGITI II In general, this muscle is similar in the two cranes. In one specimen of G. c. tabida, the belly was not bipinnate. Fisher and Goodman (1955:94) found this muscle “not clearly bipinnate” on one side of a specimen of G. c. canadensis. The total length of the belly is about 8 cm.; fleshy fibers extend further distad in the anterior half of the belly. The tendon of insertion is calcified in the region of the crus and tarsometatarsus, but not over the intratarsal joint. Fisher and Goodman found fusion between the tendons of Mm. flexor perforans et perforatus digiti II and flexor perforatus digiti II in G. c. canadensis (see also Mitchell, 1901:653), but not in G. americana. I did not find such fusion in G. c. tabida. The tendon of M. flexor perforans et perforatus digiti II perforates the tendon of M. flexor perforatus digiti II and is perforated by the tendon to digit II of M. flexor digitorum longus, as described by Fisher and Goodman (1955:94). M. FLEXOR PERFORANS ET PERFORATUS DIGITI III This muscle has a small (10 cm. long) bipinnate belly. The tendon perforates and is perforated. Hudson (1937:42) and Fisher and Goodman (1955:97) found a vinculum connecting the tendon of this muscle with the tendon of M. flexor perforatus digiti III in G. c. canadensis, but Fisher and Goodman did not find such a vinculum in G. americana. This vinculum is present in G. c. tabida. The vinculum is short (about 0.5 cm. long), but strong, and is located about 3 cm. from the distal end of the tarsometatarsus. M. FLEXOR PERFORATUS DIGIT1 IV The origin and insertion of this muscle in the Sandhill Crane are similar to those described for the Whooping Crane by Fisher and Goodman (1955:9(i). My specimens also exhibited the peculiar lateral head, which passes distad lateral to the biceps tendon before fusing with the medial head. The bulk of the muscle lies medial to the biceps tendon. Mm. flexor perforatus digiti IV, flexor perforatus digiti III, and flexor hallucis longus have a common origin, fleshy and tendinous, from the intercondylar area of the femur. This common origin is located just distal to the insertion of the accessory semitendinosus muscle and lateral to the origin of pars media of the gastrocnemius. M. FLEXOR PERFORATUS DIGIT1 III In the Sandhill Crane, the posterior head arises as described for the Whooping Crane by Fisher and Goodman. The much smaller lateral head is, in part, a direct continuation of the ambiens tendon, but a long (7 cm.), flat tendinous band, attached proximally to the head of the fibula, fuses with the ambiens tendon, just proximal to the origin of the fleshy fibers of M. flexor perforatus digiti III. This tendinous band is intimately associated with the lateral head of M. flexor perforatus digiti II. The fleshy fibers of the lateral head of M. flexor perforatus digiti III begin about 8 cm. distal to the proximal end of the fibola. There is a well developed vinculum between the tendon MYOLOGY of this muscle tendon M. and the of insertion FLEXOR This PERFORATUS Both cm. long) band from is calcified). the tendon inserts fibers tendon FLEXOR extends above. digiti III. The tendons. The ligament and Goodman lateral the same length, of M. tendon flexor of flexor et perforatus (about 4.5 and associated fascia and perforatus II, phalanx, with the ambiens arises by a flat perforatus digiti (1955: head This head is not connected about digiti digiti although III II of the is perforated the bulk as described tendinous (part by of the tendon by Fisher and Good- LONGUS of this muscle in the Sandhill described for the Whooping differs aspect of the crus. in that it does not pass through ; 1937:69) it Goodman, does pass through 1955 :99). Fisher digitorum Ic.1 longus, canodensis (nor did ; we) most of the tendon digitorum branch longus of the ensheathed unable in the hallucis by the tendon to demonstrate the tendon of flexor (representing trated In than the Sandhill the belly were its Crane, of M. the tendon Only connected longus by a weak in the pattern small was not also were In to the vinculum distal end on the hallux. vinculum Mitchell a did not fuse with by a strong inserted throughout. a very in G. americana. 4 cm. proximal tendon Crane, of flexor and Goodman tendon hallucis as was the this tendon were connected tendon), integrity them, and in Grus (2.5 of In a cm. long), (1901:654) illus- of these two deep plantar birds. the flexor belly hallucis head is short and small. (13.5 cm. long) longus (15 The tendon The of this cm.). The muscle lateral is slightly shorter head is the larger; of this muscle alone passes through insertion the is typical. BREVIS developed in the cranes (see Mitchell, 1913 :1053). the belly is 13.5 cm. long, but it is less than 0.5 cm. wide. at the level of insertion is typical of flexor longus of them of the Sandhill hallux; and there LONGUS This muscle is poorly hill Crane, between Fisher (Fisher most birds hallucis fusion tarsometatarsus. brevis. they Crane fused with to the Crane (see also Hudson, “in flexor not find of the brevis variation bony canal in the hypotarsus. PERONEUS of the of the hallucis intergeneric DIGITORUM of Mm. a vinculum directly but hallucis retained in several gruiform the medial single tendon out that longus hallucis longus, of the the two tendons hallucis FLEXOR flexor the remainder the considerable tendons one-fourth digitorum over half specimen, the hallucis such a perforation the tarsometatarsus; third only of G. c. tabida the tendon second specimen Whooping In one of my specimens continued of M. the did muscles in the Sandhill pointed tendons he noted distal of insertion in have (1937:48) of flexor tendon tendon a canal Cranes.” (15 cm. long) but it is one of the best developed the Hudson case in one of our Whooping however, such and Goodman small belly a bony canal in the hypotarsus between” “but is the same as Fisher The relatively The and Goodman is some sort of a connection flexor Crane Crane. about half way down the tibiotarsus, on the posterolateral M. perforatus by Fisher Crane. the patellar side of the proximal HALLUCIS origin (1955:98) M. from Crane Sandhill of origin perforans on the lateral et (1955:98). The but in the The inserting of flexor perforans for the Whooping or deep, head, the femoral flexor I1 band mentioned medial, M. CRANE by both of the deep flexor are present arises by fleshy The tendon M. DIGIT1 heads from the tendinous tendon. of is perforated muscle is as decribed 97-98). man tendon OF SANDHILL (see Fisher of the biceps tendon, and Goodman, as in the Whooping 1955:99-100). In the Sand- The origin Crane. The begins insertion THE WILSON BULLETIN M. EXTENSOR DIGITORUM December Vol. 1956 68. No. 4 LONGUS The belly of this muscle is about 14 cm. long, but only 1 cm. in maximum width at the head of the tibiotarsus. The tendon of insertion passes through a bony canal at the distal end of the tibiotarsus, but it is held in place by a ligament on the proximal end of the tarsometatarsus. The general development is the same as described for the Whooping Crane by Fisher and Goodman (1955:lOO). M. POPLITEUS Typical in origin and insertion in the Sandhill Crane, this small muscle is about 2 cm. long and 1.3 cm. wide. It arises on the fibula, inserts on the tibiotarsus. M. PLANTARIS This is a very small muscle in the Sandhill Crane. Its belly is about 7 cm. long, but only about 0.7 cm. in maximum width at the proximal end of the tibiotarsus. The minute tendon inserts on the medial corner of the tibia1 cartilage. The short toe muscles are very poorly developed in the Cranes (Fisher and Goodman, 1955:102), although I found remnants of the following eight muscles in both legs. M. EXTENSOR HALLUCIS LONGUS This is relatively a very small muscle, 5 to 6 cm. long and with a maximum width of but 2 to 3 mm. It seems to arise exclusively from the fascia covering the anteromedial surface of the intratarsal joint. The tendon inserts on the medial side, and not on the dorsal surface, of the distal phalanx of the hallux. M. EXTENSOR PROPRIUS DIGIT1 III This muscle is rudimentary. I found a few fleshy fibers on the anterior surface of the distal half of the tarsometatarsus, but much of the muscle seems to be represented by semitendinous fibers. M. EXTENSOR BREWS DIGIT1 IV This muscle also is rudimentary. Tendinous and fleshy fibers arise from about the distal third of the tarsometatarsus. The tendon is about 1 mm. wide; it passes through a bony canal between the trochleae for digits III and IV to insert on the medial surface of the base of the proximal phalanx, digit IV. 111.ABDUCTOR DIGITI II The 3 cm.-long belly is composed of fleshy and tendinous fibers. It arises from the medial surface of the distal end of the tarsometatarsus. The tendon inserts on the medial side of the base of the proximal phalanx of digit II. M. FLEXOR HALLUCIS BREWS This is a minute muscle, 3 to 4 cm. long and with a maximum width of about 3 mm. It arises primarily from the medial surface of the hypotarsus. It has a thin, hair-like tendon about 0.3 mm. wide. It inserts on the base of the hallux and is not perforated by the tendon of M. flexor hallucis longus. M. ADDUCTOR DIGIT1 II This muscle is typical in origin and relationships. The belly is 4 cm. long and 4 mm. in maximum width. It arises at the proximal end of the tarsometatarsus, immediately inferior to the hypotarsal area. The hair-like tendon is about 0.3 mm. wide. It inserts on the dorsomedial surface of the base of the proximal phalanx, digit II. M. LUMBRICALIS This muscle shows the poorest development of any of the short toe muscles, and I did not find it in all dissections. When present, it consists of scattered fleshy and Andrew J. MYOLOGY CRANE 301 tendinous fibers, located at the distal end of the tarsometatarsus. on the cartilaginous pads for digits III and IV. These insert primarily Rerger M. ABDUCTOR DIGIT1 OF SANDHILL IV This is another minute muscle, having a length of about 4.5 cm. and a maximum width of 3 mm. The tendon is less than 0.5 mm. wide, but it expands distally and inserts on the lateral side of the base of the proximal phalanx of digit IV. M. ADDUCTOR DIGIT1 IV This muscle is not present in the Sandhill Crane, nor in the Whooping Crane (Fisher and Goodman, 1955 :102). SUMMARY Only two of the 47 major wing muscles are absent in GFZLS americana and in G. canadensis tabida: Mm. flexor metacarpi brevis (see page 292) and entepicondylo-ulnaris. patagii” Pars propatagialis M. cucullaris (= “dermo-tensor of Shufeldt, 1890, but not of Fisher, 1946: 574) also is absent. The OS humeroscapulare similarities (see page 287) is absent in both cranes. The following of wing muscles in the two cranes deserve special mention: The biceps slip arises from the coracoidal tendon of M. biceps brachii and has a typical insertion on the tendon of insertion of M. tensor patagii longus. Mm. tensores patagii longus et brevis are represented by a single fused belly. M. scapulotriceps has a strong aponeurotic anchor extending from the anterior edge of the belly to the humerus. M. flexor digitorum profundus has a more proximal insertion on digit III (of Fisher) or digit II (of Hudson) than M. flexor digitorum sublimus. The latter muscle, though its belly is very small, is present and similar in each. M. p roscapulohumeralis is present in both cranes. M. subcoracoideus has a single head. The tendon of insertion of M. serratus anterior passes between the two heads (pars externa and pars interna) of origin of M. subscapularis. M. deltoideus major has the usual origin from the dorsolateral surface of the scapula and also has an accessory tendinous origin from the scapula caudal to the glenoid fossa. M. expansor secundariorum is well developed in G. c. tabida and undoubtedly in G. americana. The rudimentary M. anconaeus coracoideus is present in G. c. tabida and probably in G. americanu. M. adductor alae digiti II (adductor pollicis) exceptionally well developed muscle, whereas Mm. abductor major digiti III are poorly developed. (abductor indicis), flexor digiti IV is an (III’), and flexor metacarpi posterior M. latissimus dorsi metapatagialis is inconstant in both species. The following differences between G. americana and G. c. tabida were noted in the wing muscles: M. pectoralis is divided into a superficial and a deep layer in G. americana, but not in G. c. tabida. The origins of Mm. latissimus dorsi, rhomboideus superficialis, and rhomboideus profundus are less extensive in G. c. tabida than in G. americana. There is a dermal component to M. deltoideus major in G. americana, but I did not find one in G. c. tabida. M. THE 302 WILSON BULLETIN deltoideus minor apparently has a single head in G. americana; heads in G. c. tabida. it has two M. serratus posterior is divided into a superficial and a deep layer in G. americana, hut not in G. c. tabida. There are minor differences in the origins of all three serrati muscles. M. abductor alae digiti II (abductor pollicis) has two heads in both cranes; the anconal head arises from the extensor process of the carpometacarpus, the palmar head from the tendon of insertion of M. extensor metacarpi radialis. Fisher and Goodman (1955:68) state that in G. americana the palmar head arises, in part, by “tendinous fibers from the inserting tendon of M. tens. pat. longus”; it does not do so in G. c. tabida. I doubt the existence of any separate muscle which might be called “M. proscapulohumeralis brevis.” I agree with Hudson and Lanzillotti (1955 :43) that M. flexor brevis digiti IV (III) probably is best considered simply a dis- tal part of M. flexor digiti IV (III) ; that M. abductor indicis brevis is a deep (abductor indicis) ; and that M. fasciculus of M. abductor major digiti III abductor minor digiti III (abductor digiti II) is a ligament-I know of no bird in which there is a muscle in this position. The leg muscle formula in the Sandhill Crane is ABC (t iliotrochantericus medius some specimens. Hudson (C) and iliotrochantericus (1937:69) ) DXYAmV. anterior Mm. are fused in gave the formula for the Little Brown Crane (G. c. canadensis) as ABDXYAmV; I assume that the two iliotrochan- terici muscles were fused in his specimen. In two specimens of the Whooping Crane, Fisher and Goodman found the formula to be ABCDXYAm; though they added V to the formula on page 124, they stated (page 97) : “we did not find any vinculum between the tendons of M. flex. perf. dig. III perf. et perf. dig. III.” In a third Whooping and M. flex. Crane, the formula was BCDXYAm. The following features common to the Whooping deserve mention: and the Sandhill cranes The accessory semitendinosus muscle has two well devel- oped heads. M. iliotibialis has a large aponeurotic portion in the center of its distal half. ialis (= Mm. vastus lateralis (= femoritibialis femoritibialis externus) and vastus med- medius) are similar in both cranes; M. vastus lateralis has two distinct heads of origin, one proximal and one distal. In the Sandhill Crane the tendon of M. ambiens gives rise only to the lateral head of M. flexor perforatus digiti III; in the Whooping Crane the ambiens tendon serves as the “principal, if not sole, origin for M. flex. perf. dig. II.” In the Sandhill Crane, M. flexor ischiofemoralis inserts on the femur proximal to the areas of insertion of both parts of M. caudofemoralis. In the Whooping Crane M. flexor ischiofemoralis inserts “posterior to and between the insertions of the two parts of M. caudofem.” (Fisher and Goodman, 1955:86) ; this is a most unusual relationship for these tendons. Fisher and Goodman Andrew .I. BWgW MYOLOGY (1955:94) digiti II OF SANDHILL CRANE 303 found fusion of the tendons of Mm. flexor perforans et perforatus and flexor perforatus digiti americana; II in G. c. car&e&s, G. canadensis tabida. In the Sandhill Crane, only the tendon of M. flexor digitorum longus passes through Whooping Crane the tendons of flexor digitorum a bony canal in the hypotarsus; I found considerable variation the tendons of Mm. flexor digitorum page 299). Unilaterally in the longus and flexor hallucis longus pass through bony canals in the hypotarsus 1955:99). hut not in G. I did not find fusion of these two tendons in three specimens of (Fisher and Goodman, in the amount of fusion between longus and flexor hallucis longus (see in one Sandhill Crane, I found an accessory medial head, arising from the proximal end of the tibiotarsus, of pars media, M. gastrocnemius. LITERATURE CITED BERGER,A. J. 1953 On the locomotor anatomy of the Blue Coua, Coua caerulea. Auk, 70:49-83. 1956 The expansor secundariorum muscle, with special reference to passerine birds. Jour. Morph., 99:137-167. FISHER, H. I. 1946 Adaptations and comparative anatomy of the locomotor apparatus of New World vultures. Amer. Midl. Nat., 35~545-727. FISHER, H. I., AND D. C. GOODMAN 1955 The myology of the Whooping Crane, Grus americana. Illinois Biol. Monog., 24, 127 pp. F~~RBRINGER, M. 1902 Zur Vergleichenden Anatomie des Brustschulterapparates und der Schnltermuskeln. Jena. Zeitschr. f. Naturwiss., 26:289-736. GADOW, H., AND E. SELENKA Viigel. In Bronn’s Klassen und Ordnungen des Thier-Reichs. Leipzig. 1891 H~JDSON,G. E. 1937 Studies on the mnscles of the pelvic appendage in birds. Amer. Midl. Nat., 18:1-108. HUDSON, G. E., AND P. J. LANZILLOTTI 1955 Gross anatomy of the wing muscles in the family Corvidae. Amer. Midl. Nat., 53 :144. MITCHELL, P. C. 1901 On the anatomy of Gruiform birds; with special reference to the correlation of modifications. Proc. Zool. Sot. London, 1901:629-655. 1913 The peroneal muscles in birds. Proc. Zool. Sot. London, 1913:1039-1072. 1915 Anatomical notes on the Gruiform birds Aramus giganteus Bonap., and Rhinochetus kagu. Proc. Zool. Sot. London, 1915 :413-423. MONTACNA, W. 1945 A re-investigation of the development of the wing of the fowl. Jour. Morph., 76 :87-113. NEWTON, A. 1896 A dictionary of birds. Adam and Charles Black, London, 1088 pp. December 1956 Vol. 68, No. 4 THE WILSON BULLETIN PARKER, 1947 SHLJFELDT, T. J., AND W. A. HASWELL A text-book of zoology. 6th ed., vol. 2. Macmillan R. & Co., London, 758 pp. W. 1890 The myology of the Raven (Corvus coraz sinuutus). Macmillan don, 343 pp. YOUNG, J. Z. 1950 The life of vertebrates. Oxford Univ. Press, London, 767 pp. DEPARTMENT OF ANATOMY, UNIVERSITY OF MICHIGAN MEDICAL & Co., Lon- SCHOOL, ANN ARBOR, MICHIGAN, MARCH 5, 1956 NEW LIFE MEMBER Karl E. Bartel was born in Blue Island, Illinois, October 12, 1913, and now is employed by General Biological Supply House, Chicago. He has been interested in natural history since 1928. His bird watching developed into bird banding, and since 1933 he has banded over 30,000 individuals of 159 species. His interest is shifting now to wild flower photography, and to the building up of a series of wild flower slide lectures for rental purposes. Mr. Bartel is a member of the American Ornithologists’ Union, Illinois Audubon Society, Wisconsin Society of Ornithology, Nature Conservancy, Friends of our Native Landscape, Eastern, Western, and Inland Bird Banding associations, Wm. I. Lyon Bird Banding Council and the Chicago Ornithological Society. He is now President of the latter organization. In the picture he is holding a Red-bellied Woodpecker that he had just banded.